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JOURNAL
OF THE
ARNOLD ARBORETUM
VOLUME 68 JANUARY 1987 NUMBER |
PHYLOGENETIC IMPLICATIONS OF LEAF ANATOMY IN
SUBTRIBE MELITTIDINAE (LABIATAE) AND
RELATED TAXA
Monks S. ABU-ASAB AND Puitip D. CANTINO!
Leaf anatomy was surveyed in 39 species of Labiatae, including represen-
tatives of all SIX genera of subtribe Melittidinae. When subjected to cladistic
other apparently derived states, diallelocytic stomata with four subsidiary cells
and subsessile glandular trichomes with partial radial walls, suggest that the
sister group of the Macbridea-Physostegia-Brazoria clade is Galeobdolon or
Synandra. Leaf anatomy provides no evidence that subtribe Melittidinae is
monophyletic
There is relatively little published information on the anatomy of the La-
biatae, a rather surprising situation given the size and economic importance
of the family. We are aware of only a few works on leaf anatomy in particular.
The broadest in taxonomic scope are Solereder’s (1908) general anatomical
survey and Inamdar and Bhatt’s (1972) study of ee types in the family.
Other works, more intensive but narrower in taxonomic scope, are those of
Bokhari and Hedge (1971) on tribe Meriandreae, Rudall (1979, 1980) on sub-
tribe Hyptidinae, Azizian and Cutler (1982) on Phiomis L. and Eremostachys
Ledeb., and Shah and Naidu (1983) on “tribe Ocimoideae.
The primary focus of this paper is the leaf anatomy of subtribe Melittidinae,
but the study collection was selected to include a variety of other Labiatae so
‘Department of Botany, Ohio University, Athens, Ohio 45701.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 1-34. January, 1987.
ps JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
that it would be possible to evaluate the systematic significance of characters
that vary within the Melittidinae. The work was undertaken with two objec-
tives: first, to seek anatomical evidence for the monophyly (sensu Hennig, 1966)
of subtribe Melittidinae and/or its suprageneric subgroups; and second, to
contribute to the body of information available on the leaf anatomy of the
Labiatae.
TAXONOMIC BACKGROUND
LABIATAE
The most widely used classification of the Labiatae today is that of Briquet
(1895-1897), which 1s heavily based on a series of comprehensive treatments
of the family by Bentham (1832-1836, 1848, 1876). Briquet’s classification
differs from Bentham’s mainly in the ranking and interrelationships of supra-
generic groups rather than in the content of those groups (Cantino & Sanders,
1986). The suprageneric groups of both authors are based principally on gross
floral morphology.
An alternative classification of the Labiatae was proposed by Erdtman (1945)
on the basis of palynological features. He subdivided the family into two
subfamilies: Lamioideae, with tricolpate pollen that is shed in a two-celled
stage; and Nepetoideae, with hexacolpate pollen shed in a three-celled stage.
A variety of other characters have since been found to correlate with these
(Wunderlich, 1967; Zoz & Litvinenko, 1979; Cantino & Sanders, 1986).
Subfamily Lamioideae is characterized by albuminous seeds containing a spat-
ulate embryo, the production of iridoid glycosides, the absence of rosmarinic
acid, a low volatile terpenoid content (the leaves hence usually not aromatic),
moderately unsaturated seed oils, and a nonmucilaginous pericarp. Subfamily
Nepetoideae is characterized by exalbuminous seeds containing an “investing”
embryo (terminology of Martin, 1946), the absence of iridoid glycosides, the
production of rosmarinic acid, high volatile terpenoid content (the leaves hence
aromatic), highly unsaturated seed oils, and a frequently mucilaginous pericarp.
Erdtman’s subfamilial classification (1945), with its strong character support,
conflicts markedly with Briquet’s (1895-1897) widely used system but is highly
congruent with Bentham’s (1876) tribal classification (Cantino & Sanders, 1986).
In the present study, Erdtman’s subfamilial classification has been adopted.
TRIBE LAMIEAE
Inasmuch as a primary objective of this study is to investigate whether
subtribe Melittidinae is monophyletic, it is necessary to delimit a monophyletic
study group that includes (but is not limited to) the subtribe. The Melittidinae
fall within Erdtman’s subfamily Lamioideae. Although this subfamily is a
primary phenetic subgroup of the Labiatae, it has not been possible to dem-
onstrate its monophyly through the identification of synapomorphies (Cantino
& Sanders, 1986). There is, however, a less inclusive group that includes sub-
tribe Melittidinae and appears to be monophyletic. It is composed of Bentham’s
(1876) tribes Lamieae and Prasieae, excluding Anisomeles R. Br., Scutellaria
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 3
L., and probably Salazaria Torrey. This group is similar in circumscription to
Wunderlich’s (1967) subfamily Lamioideae (“Stachyoideae’’; corrected no-
menclature follows Sanders & Cantino, 1984), but if it is recognized at the
tribal level it must be called Lamieae. Except where otherwise stated, all future
use of the name Lamieae will refer to the group thus circumscribed.
The monophyly of tribe Lamieae is supported by one clear synapomorphy,
one probable synapomorphy, and a third congruent character, the polarity of
which cannot currently be assessed. Hagemann and co-workers (1967) found
an allenic component, which they presumed to be laballenic acid, in the seed
oils of all examined members of Bentham’s Lamieae and Prasieae except An-
isomeles and Scutellaria. The allenic component was not found in these two
genera or in the other examined members of Erdtman’s subfamily Lamioideae,
and it was found in only four of 122 examined species of subfamily Nepetoideae.
The polarity of this character can be assessed by outgroup analysis (Watrous
& Wheeler, 1981; Maddison et al., 1984), using suprageneric taxa of the Ver-
benaceae as outgroups. (It is generally believed that the Labiatae evolved from
the Verbenaceae, which would make the latter at best paraphyletic, but it is
unclear which members of the Verbenaceae are the closest relatives of the
Labiatae. All members of the Verbenaceae must therefore be included among
the outgroups in the assessment of character polarity within the Labiatae.) In
an unpublished study, Robert Kleiman (pers. comm.) found the allenic com-
ponent to be absent from the seed oils of all 24 species of Verbenaceae ex-
amined, including representatives of three subfamilies and eight tribes. Oc-
currence of the allenic component therefore appears to be a derived trait within
the Labiatae and represents a synapomorphy of a monophyletic group com-
posed of Bentham’s tribes Lamieae and Prasieae (excluding Scutellaria and
Anisomeles). Because Salazaria appears to be closely related to Scutellaria on
morphological (Epling, 1942) and chemical (Kooiman, 1972) grounds, it should
perhaps be excluded from the Lamieae as well, although its seed oils have not
been investigated.
Embryological peculiarities of the Lamieae offer two other possible synapo-
morphies. The mature embryo sac in the Labiatae tends to be two-lobed, with
distinct micropylar and chalazal sections. Genera differ in the relative size and
shape of these lobes. Wunderlich (1967) reported that the micropylar lobe is
much longer and broader than the chalazal one in Bentham’s Lamieae and
Prasieae (except Scutellaria and Anisomeles), whereas the micropylar lobe is
shorter than or equal to the chalazal in the rest of the Labiatae, except two
genera of Nepetoideae. Sa/azaria was not examined. Embryo-sac shape has
been reported for ten genera of Verbenaceae representing three subfamilies and
six tribes (Junell, 1934; Misra, 1939; Tatachar, 1940; Pal, 1951; Maheshwari,
1954; Khaleel & Nalini, 1972; Spies & Stirton, 1982; Spies, 1984a, 1984b;
Thirumaran & Lakshmanan, 1984). In only one species, Clerodendrum ugan-
dense Prain, does the embryo sac resemble those found in the Lamieae (Junell,
1934). In all other Verbenaceae examined, including four other species of
Clerodendrum L. (Junell, 1934; Misra, 1939), the micropylar end of the embryo
sac is usually little if at all broader (in some species narrower) than the chalazal
end; if it is much broader, it is shorter than the chalazal end. The characteristic
4 JOURNAL OF THE ARNOLD ARBORETUM [voL. 68
embryo-sac shape of the Lamieae is thus probably derived, although more
Verbenaceae need to be studied before character polarity can be assessed with
confidence.
Wunderlich (1967) reported the presence of what Schnarf (1918) called
““‘Lamium-type” glandular trichomes (identical to our “type 4’’; see trichome
classification below) on the outside of the integument in recently fertilized
ovules of all examined genera of Bentham’s Lamieae and Prasieae except Scu-
tellaria and Anisomeles, no such glandular trichomes were found in other
Labiatae (but Sa/azaria was not examined). It is not possible to assess the
polarity of this character because of lack of data for the Verbenaceae, but its
distribution in the Labiatae closely parallels that of the other two characters.
Although a strong case can be made for the existence of a monophyletic tribe
Lamieae (as circumscribed above), one must remain aware that the characters
delimiting the group have been examined in a minority of its members. Seed-
oil chemistry was studied (Hagemann ef a/., 1967) in 18 of the 42 genera of
Bentham’s (1876) Lamieae and Prasieae, and the two embryological characters
cited above were studied in 16 genera of these tribes (Wunderlich, 1967). There
are 11 genera for which data are available for all three characters. Because the
congruence between the three characters is perfect in these genera, we are
assuming that the characters are highly correlated in the group as a whole.
Examination of more genera may demonstrate, however, that others besides
Scutellaria and Anisomeles are not members of the monophyletic group. Our
tentative inclusion of all of Bentham’s Lamieae and Prasieae (except Scutel-
laria, Anisomeles, and possibly Salazaria) reflects our confidence in Bentham’s
usually excellent taxonomic judgment—1.e., we are assuming that those genera
not yet examined for embryology and seed-oil chemistry really are closely
related to those that have been.
SUBTRIBE MELITTIDINAE
The historical changes in the circumscription of subtribe Melittidinae have
been summarized by Cantino (1985a). As currently circumscribed, the subtribe
comprises six genera, four of them (Brazoria Engelm. ex A. Gray, Macbridea
Elliott ex Nutt., Physostegia Bentham, and Synandra Nutt.) North American,
one (Chelonopsis Miq.) Asian, and one (Melittis L.) European. The group is
delimited on the basis of a set of calyx and corolla characters that were proposed
by Bentham (1876) and adopted by Briquet (1895-1897): calyx broadly cam-
panulate, membranaceous or herbaceous, 3- or 4-lobed or 5-toothed, with
venation scarcely visible; corolla tube long-exserted from calyx, site at base
or markedly dilated distally, with upper lip broad and scarcely conc
A survey of these characters in subfamily Lamioideae (Cantino, ened
data) revealed that none is diagnostic of subtribe Melittidinae. Three states
used by Bentham and Briquet (calyx broadly campanulate, calyx membrana-
ceous or herbaceous, and corolla tube long-exserted from the calyx) are present
throughout the Melittidinae but are also common elsewhere in the subfamily.
The other character states cited by these authors are not only found elsewhere
in the subfamily but also occur in only some members of the Melittidinae.
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 5)
Weak calyx venation at anthesis is characteristic of Physostegia, Brazoria, and
(to a degree) Synandra, but not the other three genera. The upper lip of the
corolla is broad and only barely concave in Physostegia, Chelonopsis, and three
species of Brazoria but markedly concave in Macbridea, Synandra, and Bra-
zoria scutellarioides. The number of calyx lobes varies from three to five, with
no two genera having the same calyx morphology. Corolla-tube shape is sim-
ilarly variable. It is, of course, insufficient to consider characters only singly.
In groups in which parallel and/or reticulate evolution have been common,
taxa are often distinguished by combinations of character states, with no single
state unique to any taxon (“kaleidoscopic variation”; see Cantino, 1982). How-
ever, the combination of the three character states that occur throughout the
Melittidinae is also found in some or all species of at least ten other genera in
subfamily Lamioideae (Colguhounia Wallich, Gomphostemma Bentham, La-
mium L., Microtoena Prain, Phyllostegia Bentham, Scutellaria, Stenogyne Ben-
tham, Tetraclea A. Gray, Thuspeinanta T. Durand, and Trichostema L.), seven
of them in tribe Lamieae as circumscribed above.
We are unaware of any morphological feature or combination of features
that would distinguish subtribe Melittidinae from the rest of tribe Lamieae, let
alone a clearly derived feature. Nor does cytology provide evidence for the
monophyly of the subtribe. Chromosome number is extremely variable among
the genera, chromosome size is moderately variable, and other karyotypic
features are restricted to particular species or species groups (Cantino, 1985a).
The present study was undertaken to investigate whether leaf anatomy might
provide evidence for the monophyly of subtribe Melittidinae, where mor-
phology and cytology have not.
STOMATAL TERMINOLOGY
Because of the variety of stomatal classifications now available and the
sometimes conflicting use of terms contained therein, a brief review of the
situation is necessary if the reader is to understand our adopted terminology.
For a more comprehensive and very enlightening review, see Rasmussen (1981).
Stomata have been classified on the basis of three criteria: the configurations
of neighboring and subsidiary cells in mature stomata (Vesque, 1889; Metcalfe
& Chalk, 1950; Payne, 1970), stomatal ontogeny (Pant, 1965; Stevens & Martin,
1978; Payne, 1979), and a combination of the above (Fryns-Claessens & Van
Cotthem, 1973; Stevens & Martin, 1978).
The first criterion is relatively uncomplicated and has the advantage that it
can be applied when one is working with mature leaves. Its principal disad-
vantage is that the same stomatal morphology may develop through different
ontogenetic pathways in different plants and may therefore not be homologous
(Rasmussen, 1981, and references cited therein). Classifications based partly
or completely on stomatal ontogeny are more difficult to apply, and some of
the terms used are defined differently by different authors.
Pant (1965) classified stomata on the basis of their ontogenetic pathways:
mesogenous stomata, in which the guard-cell mother cell and all subsidiaries
are derived from the same meristemoid; perigenous stomata, in which all
6 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 68
neighboring and subsidiary cells are derived from protodermal cells other than
the meristemoid that produces the guard-cell mother cell; and mesoperigenous
stomata, in which the surrounding cells are of dual origin, some mesogenous
and others perigenous.
The guard-cell mother cell is the immediate progenitor of the guard cells.
Subsidiary cells surround the guard cells and clearly differ from other epidermal
cells; neighboring cells immediately surround the guard cells but do not differ
in shape from the remaining epidermal cells (Fryns-Claessens & Van Cotthem,
1973; Rasmussen, 1981). Unfortunately, the ambiguity of the term ‘“meri-
stemoid”’ has rendered Pant’s and other ontogenetic classifications difficult to
use.
Stomatal ontogeny starts with the unequal division of a protodermal cell.
The smaller daughter cell, which contains a denser cytoplasm, divides again
unequally or directly produces (by an equal division) the pair of guard cells
(Fryns-Claessens & Van Cotthem, 1973; Payne, 1979; Rasmussen, 1981). The
term “‘meristemoid” was used by Fryns-Claessens and Van Cotthem (1973)
and Rasmussen (1981) to refer to the smaller daughter cell of the original
protodermal cell, whereas Payne (1979) referred to the protodermal cell itself
as the meristemoid. If the latter usage is adopted, there is always at least one
neighboring or subsidiary cell that is derived from the meristemoid (i.e., me-
sogenous), sO a true perigenous type cannot exist (Fryns-Claessens & Van
Cotthem, 1973; Payne, 1979). A consequent disadvantage of Payne’s termi-
nology is that it is less precise; i.e., a wider variety of ontogenetic pathways
is necessarily subsumed under the same term, mesoperigenous (see fig. 3 in
Rasmussen, 1981). For this reason, and because the meristemoid sensu Payne
can only be recognized after it has divided and hence no longer exists (Ras-
mussen, 1981), the ontogenetic terminology of Fryns-Claessens and Van Cott-
hem (1973) rather than that of Payne (1979) is adopted in this study. The more
complex system of Stevens and Martin (1978) is even more precise but is not
used here because of the difficulty in distinguishing ‘‘agene”’ cells (Rasmussen,
1981) from perigene cells sensu Rasmussen
MATERIALS AND METHODS
Leaf material was obtained from 53 specimens representing 39 species (see
APPENDIX 1), including all species of Brazoria, Macbridea, Melittis, and Syn-
andra, seven of the 12 species of Physostegia, and two of the approximately
16 species of Chelonopsis. Leaf material of most species was collected from
living plants, with herbarium specimens prepared as vouchers. Leaf material
of Chelonopsis, Melittis, and some species of Physostegia was obtained directly
from herbarium specimens.
Fresh leaves were fixed in Carnoy’s solution (3 parts ethanol to | part acetic
acid). Dried leaves were revived by soaking them in 5 percent sodium hydroxide
for three days at room temperature. Both types of material were then stored
in 70 percent ethanol. To prepare the material for study, we used the whole-
mount method as well as transverse sectioning of the lamina. In the former
method leaves or leaf pieces were stained with ferric tannate (2.5% tannic acid
1987] ABU-ASAB & CANTINO, LEAF ANATOMY i
in 50% ethanol, followed by 2.5% ferric chloride in 50% ethanol; modified
from Berlyn & Miksche, 1976) and mounted in surface view. In the latter,
leaves were infiltrated with and embedded in paraffin (Cutler, 1978) and sec-
tioned at 10-um thickness with an AO rotary microtome. After sectioning, the
leaves were stained with toluidine blue or with safranin O and fast green FCF.
The procedure using toluidine blue is outlined in Sakai (1973). The double-
staining procedure, adapted from Johansen (1940), required deparaffination of
the sections, staining with safranin (1% in 50% ethanol) and fast green (0.1%
in 50% ethanol), dehydration through a series of ethanol, xylene: ethanol
(1:1), and xylene, and mounting in Permount.
A set of permanent slides has been deposited in the Bartley Herbarium of
Ohio University (BHO). Drawings were prepared by means of a microprojector
or the camera-lucida attachment of an Olympus BH-2 microscope.
RESULTS
STOMATA
Based on shapes and arrangements of mature subsidiary and neighboring
cells, the following types of stomata were found in the species examined (see
FiGureE 1) (definitions follow Payne, 1970, 1979; and Wilkinson, 1979): an-
omocytic (stoma surrounded by a limited number of cells that are indistin-
guishable from other epidermal cells); paracytic (stoma bordered on both sides
by one or more subsidiary cells whose long axes lie parallel with the long axis
of the guard cells; subsidiary cells sometimes meeting over the poles and some-
times laterally elongated); anisocytic (stoma surrounded by three cells, one of
which is markedly smaller than the other two); diacytic (stoma enclosed by a
pair of subsidiary cells whose common radial walls are at right angles to the
guard cells); and diallelocytic (stoma enclosed by three or more C-shaped cells
at right angles to the guard cells).
Two subtypes of diallelocytic stomata were found in the species examined,
one with three subsidiary cells and the other with four. The two have not been
distinguished by previous authors, including Payne (1970), who discussed the
ontogeny of diallelocytic stomata. Since the two types do not always occur
together (see TABLEs 1, 2), they are worth distinguishing. The three-celled type
will be referred to as diallelocytic-1 and the four-celled type as diallelocytic-2.
The ontogenetic pathways of several stomatal types were documented through
examination of young leaves in various stages of development (see FiGuRE 1).
The diallelocytic-1 type was studied in Scutellaria lateriflora, Stachys riddellii,
and Stachys tenuifolia and is mesoperigenous in all. Because the diallelocytic-2
type occurs only with the diallelocytic-1 type in the species examined (although
the latter may occur without the former), and the former differs from the latter
in having one more subsidiary cell, the ontogenetic pathway reported by Payne
(1970) for the diallelocytic-2 type is presumed to occur in the taxa examined
in this study. The ontogenetic pathway for the diacytic type was also adopted
from Payne (1970). The ontogeny of the anomocytic type was studied in Scu-
tellaria lateriflora, Stachys tenuifolia, and Stachys riddellii and is perigenous
8 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
@ 62
mene Paracytic
DPMS)
\ eroe
a 5b \ 6b
4c
Diacytic Sc 6c
Diallelocytic1 Diallelocytic2
Figure |. Stomatal ontogenetic pathways: anomocytic (la), paracytic (2a, b), aniso-
cytic (3a-c), diacytic (4a—c), diallelocytic-1 (5a—c), and diallelocytic-2 (6a—c). M = meri-
stemoid (sensu Fryns-Claessens & Van Cotthem). Diacytic and diallelocytic-2 ontogenies
adopted from Payne (1970).
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 9
in all. The ontogeny of the paracytic and anisocytic types was studied in 771-
chostema dichotomum. Both are mesoperigenous, and they share a common
initial step in their ontogenetic pathways.
Other stomatal features examined included distribution (viz., both leaf sur-
faces or abaxial surface only), position in relation to general level of epidermis
(viz., sunken or raised), and presence of stomatal ledges. Stomatal ledges are
elevated extensions of the cuticle that rise from the guard-cell surface “like an
incompletely roofed dome” (Wilkinson, 1979, p. 97). They extend over the
stomatal pore, delimiting an outer cavity (see fig. 10./ in Wilkinson, 1979). In
some taxa similar inner ledges project from the guard cells, forming an inner
cavity (Wilkinson, 1979), but only outer ledges were found in the present study.
Published data on stomatal types in Labiatae and Verbenaceae are scarce,
and the authors do not always clearly indicate the taxonomic distribution of
stomatal types. Data from Inamdar (1969), Ramayya and Rao (1969), Payne
(1970), and Inamdar and Bhatt (1972) are summarized in TABLE 1; our own
observations are shown in TABLE 2.
Diacytic, diallelocytic-1, and anomocytic stomata are all common in the
Labiatae. Of the 39 species we examined, diallelocytic-1 stomata were found
in 31, diacytic in 33, and anomocytic in 24. Diacytic and anomocytic stomata
occur widely in both the Labiatae and the Verbenaceae, but diallelocytic-1
stomata are apparently rare in the Verbenaceae, having been reported only
from Lippia lanceolata.
Diallelocytic-2 stomata have been observed in seven genera of Labiatae and
one of Verbenaceae. Specifically, we observed this type in six of seven examined
species of Physostegia, both species of Macbridea, one of the four species of
Brazoria, the single species of Galeobdolon Adanson, and two of the seven
examined species of Scutellaria; it has also been reported from Ocimum L.,
Plectranthus L’Hér., and Lippia L.
Anisocytic omnia appear to be rare in the Labiatae. me fee ae only
in Trichostema and Prostanthera Labill. In previous studies anisocytic stomat
have been reported from three species of Verbenaceae te not in any Dabiatae.
We found paracytic stomata only in Trichostema, Prostanthera, and Melittis,
and in the latter two they are rare; they have also been reported from three
species of Verbenaceae. Parallelocytic stomata (Payne, 1970), which resemble
the diallelocytic-2 type but have the subsidiary cells parallel to the guard cells,
have been found in Lippia lanceolata (Abu-Asab, 1984) but notin any Labiatae.
Helicocytic stomata (Payne, 1970) were included in the drawing of Lavandula
gibsonii in Inamdar and Bhatt (1972, fig. 13).
Leaves are amphistomatic in the North American Melittidinae, Scutellaria
integrifolia, and the examined species of Ajuga L., Trichostema, Lamium,
Marrubium L., and Prostanthera. They are hypostomatic in the rest of the
Labiatae examined, including Chelonopsis and Melittis of subtribe Melittidinae.
Most species of Labiatae and Verbenaceae investigated in previous studies
have hypostomatic leaves. We found intrageneric variation in this character
in Scutellaria, and such variation has also been reported in Leucas R. Br. and
Ocimum (Inamdar & Bhatt, 1972), Eriope Humb. & Bonpl. ex Bentham (Ru-
dall, 1979), and Phiomis (Azizian & Cutler, 1982).
JOURNAL OF THE ARNOLD ARBORETUM
[VOL. 68
TABLE |. Published data on stomatal types in Labiatae and Verbenaceae.*
Species
Stomatal TypesP
3. 4 5 6 7 8
Sst.
Loc.© Ref.4
Labiatae
Subfamily Lamioideae
e Lamieae
eonotis Bees Seca R. Br.
el
Congas ae ae en
biflora
Saieioe es _sptengel
linifolia
urttcife Li
fae 17
ae
eo 8 @ 6
zey oe
mie ie Bee Royle ex
Be am
Other Lamioid
Anisomeles heyneana Bentham
A. scenes B
Dysophyl Aenea Blume
Pogostenon parvi oS Bentham
Micro meria a capitata Bentham
Tribe ts
Acrocephalus capitatus Bentham
Coleus ome (Bentham)
Lour.
- gibsonii aha
Cree eee achyun Bentham
imum a illd
silicum
a
basilicum a
canum en
ratiss
kilimandscharicun Gurke
sanctum
ee ole
sanctum L.
t enoetpnon oaks Royle ex
t
incanus Lin
P. mollis (Burman) Kuntze
Tribe Salvieae
Salvia plebeia R. Br.
S. santolinifolia
Verbenaceae
Subfamily Verbenoideae
Lantana camar
Lippia lanceolata Michaux
Stachytarpheta jamaicensis
-) M van
venosa eee & Hooker
t++itt bette tet
++
++
tee eeeeteeteeees
tebe bl
+r tts
1++4+4+1
\ 1
t++etet tee eteet
1 1
\ 1
' 1
1 \
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1
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PVEnBRoCoST Ge aa
v6 ja sits oge ore >}
mm
rrr mY
jo 04
Promo
WWWW WW WwW
WWW WW
WOWWWWOW OW WWW Ww
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 1]
TABLE | (continued ).
Stomatal Types St.
Species 2 3 4 5 6 7 8 Loc. Ref.
(continued)
Subfamily viticoideae
(++i
{
fo)
Nek
Ke
je)
be
Ic
I
ao)
o
+++
'
!
Holmskioldia Sanguinea Retz.
oe ibe eg iceae
melina arborea Roxb. - - = + ©= = - =
Vitex negundo L. - +
1
!
joomroo mms omrccmrerarecwiocane sy
8piallelocytic types in eae oes and Inamdar and Bhatt Nees
am
dp
Sen 1972; 4 = Payne: 1970; 5 = Ramayya & Rao,
Outer stomatal ledges were present in all genera except Trichostema. Stomata
were found to be at the same level as the epidermis in 24 species, slightly
elevated in 12, and markedly elevated in four. Stomatal position varied within
a number of genera and within one species (Physostegia virginiana). It is prob-
ably of little taxonomic value at the generic level and of none in delimiting
suprageneric groups.
Our data do not support the observations of El-Gazzar and Watson (1968,
we examined. All 16 investigated species of Bentham’s Lamieae (including
Scutellaria but not the North American Melittidinae) had anomocytic stomata;
however, all but one also had diacytic stomata, and 12 of them had diallelocytic
stomata as well. In most species in which both diacytic and anomocytic stomata
were found, the former type was more abundant. El-Gazzar and Watson’s
(1970) generalization about Bentham’s Lamieae thus appears to be incorrect.
(On the other hand, anomocytic stomata are at least present in all examined
Lamieae except three genera of North American Melittidinae.) Our data also
disagree with regard to particular genera in table J of El-Gazzar and Watson
(1968). They included Ajuga, Galeobdolon, Teucrium L., and most species of
Pogostemon Desf., Scutellaria, and Stachys L. in their list of taxa whose stomata
JOURNAL OF THE ARNOLD ARBORETUM
TABLE 2. Stomatal characters in Labiatae examined.’
[vVoL. 68
Taxa
Stomatal Types?
4 5 6
St.
Loc.© Ldg.@
St.
Pos.©
sre ge ae aac
Tribe La
eubeeiye Meee eines
Brazoria arenaria
pulcherrima
B.
B. scutellarioides
B. truncata
Chelonopsis forrestii
r+te++
1++ i
tr terteeetrttetet
t++te¢etee ti
t++¢teu¢ti¢ti
1+
r++ +++
+ +
pepe et
\
teeter
++
In
FIA [51M | | [in |
Ke |D + © © ©
re iC
0 |Q. 10 |a
omen amit)
O | |R-\K |
Q |W |3 Ie IR ect |<
© |ct | |B |p
© |o ct
28 lea |p
m |m | |
ct 3
JE [O.1o |
b- 2
a fv)
on =)
fe) n
ct (00)
fe)
3
Cc
5
lanc
Subfamily Newared iene
Blephilia hirsuta
narda fistulosa
Peet teeetute
+r tetete+eteties
t+ettteeeetgetit
nm PPM TUIM ere roe rye TRoP cer Pe FPP PPPoE Poco Se ey
t++eetetetee+erettt+tetet
poeororooonoapoaoase
+++
yom
t+tte¢+
soooe D
r+ttteetetetetest
epvyangwaHaPAraAHaADS
++
au
“subfamilial classificati
on follows Cantino and
€stomatal position:
vated,
ele
subfamilies.
5 oe
sea anen tp anata
nisocytic, 6
os H
sent; -, absent.
level with the epidermis,
strongly Cedar ad.
b
Sanders's (1986)
Tribe Lamie
ae
‘pa eracytic,
hypostomatic
slightly
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 13
are predominantly anomocytic and/or anisocytic. We found the stomata of all
of these genera to be predominantly to entirely diacytic and/or diallelocytic.
TRICHOMES
NONGLANDULAR TRICHOMES. Simple, uniseriate trichomes were found on the
leaves of most species. Only Physostegia (all species examined), Macbridea
(both species), and Prostanthera rotundifolia lacked nonglandular foliar tri-
chomes and could be described as having glabrous leaves if the minute, sub-
sessile glands were ignored (see below). Of these three genera, only Physostegia
consistently has glabrous leaves (Cantino, 1982). Species of Prostanthera not
examined in this study have pubescent leaves (Conn, 1984), as do some in-
dividuals of both Macbridea alba (Godfrey & Wooten, 1981; Kral, 1983) and
M. caroliniana (Godfrey & Wooten, 1981). The leaves of Brazoria are nearly
glabrous, with the usually sparse trichomes concentrated toward the base of
the blade (the leaves of B. scutellarioides are essentially glabrous throughout).
The only other nonglandular trichomes observed were dendritic in form and
confined to Marrubium vulgare. These are stalked and basally branched, with
the stalk composed of several cells, a ray arising from each stalk cell, and each
ray composed of one to six cells. Similar trichomes were reported by Solereder
(1908) and Theobald and colleagues (1979) from other species of Marrubium
and were illustrated in the latter publication.
The simple trichomes vary in cell number (see TABLE 3). Unicellular tri-
chomes were found in eight genera and 12 species, while multicellular ones
were observed in all species. The variation in cell number may prove on further
study to be of taxonomic use within genera or in distinguishing among closely
related genera, but it appears to be of no value in delimiting suprageneric taxa
in the Labiatae.
GLANDULAR TRICHOMES. Two distinct classes of glandular trichomes were ob-
served. Clavate glandular trichomes (see FiGurE 2), found in 14 species, consist
of a unicellular or multicellular head resting on a relatively long, multicellular,
uniseriate stalk, the uppermost cell of which is usually discoid. Subsessile
glandular trichomes (see FiGurE 3), found in nearly all species, consist of a
unicellular or multicellular head borne on one (rarely two) short, discoid stalk
cell(s) resting on one or more foot cells. The foot cells are generally sunken
below the level of the adjacent epidermis, the gland as a whole lying in a tiny
depression on the leaf surface. The cuticle is fused to the wall of the stalk cell
but appears to separate from the wall of the head, leaving a space in which
secretions accumulate. (For excellent photographs, plus evidence that the sep-
arated cuticle is provided with a noncellulosic framework derived from the
outermost wall layer of the head cells, see Bruni & Modenesi, 1983.)
There is considerable variation in size and morphology of clavate glandular
trichomes (see FiGuRE 2). Those of Synandra hispidula and the four species of
Scutellaria in which clavate glandular trichomes were observed (S. e/liptica, S.
nervosa, S. ovata, and S. serrata) are quite similar, with a four-celled head atop
a more or less discoid stalk cell, and three to six elongate stalk cells. Clavate
14 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 68
TABLE 3. Simple, nonglandular trichomes in Labiatae examined.*
Trichomes?
Species al 2 3 4
Subfamily oo
Tribe Lami
Subtribe nat ee baeaae
Brazoria arenaria -
B. pulcherrima -
B. scutellarioides ~
B. truncata =
Chelonopsis forrestii -
Cc. moschata =
Melittis melissophyllum -
Synandra hispidula =
Other Lamieae
Galeobdolon luteum -
++ ett¢t
t+t+ttti +s
b++tut
t
ay
o
O
3s
Cc
H
G
7)
a
se)
R
QO.
=
m
Q
ie)
|
t++++44
Scutellaria elliptica -
S. incana +
. integrifolia +
+
. lateriflora
nervosa
rn ir
|4 5 ITA [TA |~
167)
a
“TR
!
ata
eucrium oe
t+++tt+eetpe+p+e¢es
++t+tteeeeeeest
+++eeeeeu tees
Subfamily Nepetoideae
Blephilia hirsuta
Monarda fistulosa
+ +
++
++
+ +
4species with glabrous leaves omitted; see text.
Classification of suprageneric taxa as in TABLE 2.
Otypes of trichomes: 1, one-celled; 2, two-
celled; 3, three-celled; 4, with more than three
cells; +, present; -, absent.
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 15
trichomes with heads composed of more than four cells were found only in
Brazoria truncata and B. scutellarioides but resemble those on the calyx and
inflorescence axis in Physostegia (Cantino, 1979, 1982). Clavate trichomes with
single-celled heads were observed in Marrubium vulgare and Trichostema lan-
ceolatum.
Subsessile glandular trichomes are very characteristic of the Labiatae and
occur in many Verbenaceae as well (Solereder, 1908; Metcalfe & Chalk, 1950).
They have been referred to by a variety of names, including shortly-stalked
bladder-like glands (Metcalfe & Chalk, 1950), sunken glandular dots (Huang
& Cheng, 1971), glandular scales (Bosabalidis & Tsekos, 1982), and glandular
capitate sessile trichomes (Shah & Naidu, 1983). The term subsessile seems
appropriate to us since the glands appear to be sessile unless examined very
closely. Because of their nearly universal occurrence in the Labiatae and the
extensive variation in their complexity, subsessile glandular trichomes offer
considerable potential as a taxonomic character in the family. They have been
little used for this purpose, perhaps in part due to lack of a satisfactory clas-
sification of the glands on which to base taxonomic comparisons. We have
developed such a classification (see APPENDIX 2), based on number of cells and
cell-wall configurations (FIGURE 4) in the head of the gland.
Terms used to describe cell-wall configurations are adopted from Stace (1973).
A primary radial wall originates from the center of the head of a gland and
ends at the periphery. A secondary radial wall originates on a primary radial
wall and ends at the periphery. A tertiary radial wall originates on a secondary
radial wall and ends at the periphery. A tangential wall connects two radial
walls. A partial radial wall originates on a tangential wall and ends at the
periphery.
Subsessile glandular trichomes (see FiGuRE 5) were found on the leaves of
all species examined except Trichostema lanceolatum (see TABLE 4). Gland
types 4 and 5 were encountered in nearly all species, both within and outside
of subtribe Melittidinae. Types 2 and 3 are rare and type | absent in the
Melittidinae. Type 1 was found only in Pogostemon, while type 2 was com-
monest in Teucrium and Pogostemon.
The more complex gland types (6-10) were most frequently encountered in
Scutellaria and the North American Melittidinae. Glands with partial radial
walls (types 7 and 10) were restricted to subtribe Melittidinae, where they were
found in all species of Brazoria, Macbridea, and Synandra, as well as in two
species of Physostegia. Type 7 was found only in Synandra. The systematic
value of the complex gland types in subtribe Melittidinae is discussed below.
LEAF HISTOLOGY
All species examined have a uniseriate epidermis composed of unsclerified,
thin-walled cells and a midrib consisting of an arcuate collateral bundle; all
lack a hypodermis. Of the other characters investigated, two (number of cell
layers in palisade parenchyma and shape of palisade cells) are too variable on
individual specimens to be of any taxonomic use. Histological characters that
16 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
a IG
e
Ficure 2. Clavate glandular trichomes: a, Brazoria arenaria; b, B. scutellarioides, c,
B. truncata; d, Melittis melissophyllum, e, Stachys riddellii: f, Synandra hispidula; g,
Scutellaria nervosa; h, S. elliptica; i, Trichostema lanceolatum: Jj, Marrubium vulgare.
Scale bars = 15 wm
1987] ABU-ASAB & CANTINO, LEAF ANATOMY ie,
Cut. Head
Epidermal
cells
Stalk cell
Foot cells
FiGuRE 3. Subsessile glandular trichome (Macbridea alba), transverse section. Cut. =
cuticle.
may be of some taxonomic value are listed in APPENDIX 3, and the distribution
of their character states is summarized in TABLE 5.
Most of these characters vary too much within genera to be of much value
in circumscribing suprageneric groups. They may prove useful, however, in
distinguishing species or species groups within certain genera (i.e., characters
2 and 6 in Brazoria; 1, 2, 4, 5, 6, and 7 in Physostegia; 2 and 4 in Scutellaria;
and 1 in Trichostema). A much more extensive sample will be necessary before
even tentative conclusions can be drawn at this taxonomic level.
In the assessment of phylogenetic relationships above the genus level, pres-
ence of idioblasts in the mesophyll appears to be the character with the greatest
potential because it varies among but not within genera. Two kinds of idioblasts
were observed. One of them, seen only in Pogostemon, resembles a glandular
trichome but occurs inside the leaf (FiGuRE 6h). These structures were also
noted by Solereder (1908, p. 1022), who described them as “internal glandular
hairs” provided with a short stalk of two or three suberized cells and a uni-
cellular, cuticularized head projecting into the intercellular spaces. He also
noted that a secretion accumulates under the cuticle, which is raised like a
bladder, just as in an external trichome.
The second kind of idioblast is a large, saclike cell, presumably secretory in
function (FIGURE 6a-g). These were observed in all species of Brazoria and all
investigated species of Physostegia. They were also noted by Solereder (1908)
in Physostegia intermedia, a species that we did not examine. Our observations
and those of Solereder suggest that within the Lamiales such saclike idioblasts
are unique to these two genera. They vary in shape and thus may offer a good
18 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
2ry rad. wall
Tan. wall
3ry rad. wall
Partial rad. wall
Cut Iry rad. wall
Ficure 4. Subsessile glandular trichome, surface view, showing cell-wall configura-
tions in head. Cut. = cuticle, tan. = tangential, rad. = radial. (Adapted from Stace, 1973.)
taxonomic character at the species level, in addition to providing evidence for
a close relationship between Physostegia and Brazoria.
DISCUSSION
Discussion will center on the question of whether leaf anatomy provides
evidence for the monophyly of subtribe Melittidinae as a whole and/or of
subgroups within it. The existence of shared, derived character states (synapo-
morphies) would constitute such evidence (Hennig, 1966; Wiley, 198 1).
The identification of synapomorphies is a two-step procedure. First, an evo-
lutionary transformation series (Wiley, 1981) is hypothesized for each char-
acter, usually on the basis of ontogeny and structural complexity of the character
states. (This step is trivial when the character is binary.) Second, the evolu-
tionary polarity of the characters must be assessed. Of the many criteria that
have been used to determine polarity (see review by Stevens, 1980), outgroup
comparison is now the most widely accepted (see, for example, Eldredge &
Cracraft, 1980; Stevens, 1980; Arnold, 1981; Wiley, 1981: Farris, 1982) and
is the sole criterion used here. Outgroup comparison, in its simplest form, can
be explained as follows: “For a given character with 2 or more states within a
group, the state occurring in related groups [the outgroups] is assumed to be
the plesiomorphic state” (Watrous & Wheeler, 1981). (For a thorough discus-
sion of the underlying logic of outgroup comparison, which is based on the
principle of parsimony, see Maddison et al., 1984
Because the monophyly of the Melittidinae is in question, the ingroup must
be a demonstrably monophyletic group that includes (but is not limited to)
this subtribe. The least-inclusive such group is tribe Lamieae, as circumscribed
above (i.e., Bentham’s tribes Lamieae and Prasieae minus Anisomeles, Scu-
tellaria, and probably Salazaria; see ““Taxonomic Background” for evidence
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 19
al 2
: b1 b2
a7 a8 a9
: c1 c2
b3 63
d1 d2 d3
e1
d4
f1 f2 f3
e2 e3
\ h1 h2
91 g2 93 h3
30pm
———_—_—1
Ficure 5. Subsessile glandular trichomes, surface view: al—a9, Brazoria arenaria;
bl-b3, Physostegia virginiana subsp. praemorsa; cl-c3 Macbridea alba, d1-d
caroliniana; el-e3, Synandra hispidula; f1-f£3, Teucrium canadense. gl-g3, Marrubium
vulgare, h1-h3, Scutellaria incana.
JOURNAL OF THE ARNOLD ARBORETUM
TABLE 4. Distribution of subsessile glandular trichomes.*
[VOL. 68
Taxa
Trichomes?
4 5 6 7
su
ae re
ribe La
pe eerie See eee
Brazoria arenaria
B. pulcherrima
B. scutellarioides
B. truncata
Chelonopsis forrestii
Macbridea alba
M. carolinia
Melittis neice ophyllum
Physosteqia angustifolia
Gigitalis
subsp. virginiana
Ajuga re
eee ere ant
Prostanthera rece Te re
Scutellaria elliptica
incana
inteqrifolia
lateriflora
[CA 12 [tA [tn | |ta
errata
Teucrium canadense
Monarda fistulosa
'
'
t+ttttetete+eetes
tttteeettere++tet
I++
1
1++H+ 1
ee one ous oe
t++tteee Ft
t+tt+e+ +44
+++
1
P++4+
11a
++tteteteteeust
Pirtttt+eteti
1
I !
ToOToOvV IO!
I
I
l
+
'
I
P+rtitst
fh do ae al
P+ti tees
Peete e ttt
he a ie ee
Ptr tee tees
Aclassification of suprageneric
byNumbers (1-10) and letters
cla
ssification in APPENDIX 2;
a,
+,
taxa as in TABLE 2.
b) refer to oo
abs
present; -,
nt.
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 2
TasLe 5. Leaf histological characters as recorded from transverse sections.
Characters?
Taxa 1 2 3 4 5 6 7
Subfamily Lamioideae
Tribe Lamieae
Subtribe Melittidinae
Brazoria arenaria
B. pulcherrima
B. scutellarioides
B. truncata
Chelonopsis forrestii
ea a
carolini
ae me1issophy1lum
Physostegia anaustifolia
De digital’s
o
~ oO
o
i
Ito Iho Iho Iho Ir |
ke Ik
-\S Jo jo lo
3 fo |o
ct |
- Ir-ls [P-|o
3 IK | fo |
-lo | |
ey) <
3 Ip |e
9 [ei
o |p
lowe OME OME CHROME OC} oun OME On fommo mows Mowiomiememi om om cmemonl ey
gpoonnonanpaAaDATs
yronvwnaannaAAAD hp
popnpnnanaaraATrns
wy
Q
subsp. virginiana
Synandra hispidula
Other Lamiea
Canepa luteum
Lamium purpureum
9oO oF wrouvrororonwwaaraorvroT
90 agvoannagaoaoaaooa pp
00D voorrwandaoroan
~
o 9
aq a
wopone om
yoo Dp
opapp p
vouradna na
qgqaqaadna
ooo o &O
ther Lamioideae
©)
Scutellaria elliptica
Poa ene
3
©
K
<
oO
n
se}
|
@
re
1)
KR
| oo
rh
| oo
ie)
nh
i)
avvrvrurrrovroooUT
wpoanpnorwawrovwnAoMD Dm
opoonoomnponoaonnan D
vwonpoonaaonaAanADD
opoopoanoanpaaaAAD
avannananaanays
2020020202009 0 &
Blephilia hirsuta
Monarda fistulosa
omen
oY
@
@
io
Q
()
a a a a
Q
2
@classification of suprageneric taxa as in TABLE 2.
characters explained in APPENDIX 3
22 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
‘wa iy ( toe a h
2 oe ee
FiGurE 6. Idioblasts in mesophyll. a-d, saclike idioblasts, surface view: a, Brazoria
arenaria (Kessler 5771), b, Brazoria pulcherrima (Kessler 5862). ¢, Physostegia godfreyi
(Cantino 1054), d, Physostegia leptophylla (Cantino 971). e-g, saclike idioblasts, trans-
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 23
supporting the monophyly of this group). The ingroup is represented in our
study by all six genera of subtribe Melittidinae plus five others (Galeobdolon,
Lamium, Leonurus L., Marrubium, and Stachys). Anatomical data for two
additional genera (Phlomis and Eremostachys) have been provided by Azizian
and Cutler (1982). For stomatal characters the ingroup sample includes Leono-
tis (Pers.) R. Br. and Leucas, as well (Inamdar & Bhatt, 1972).
Choice of outgroups is constrained by both the uncertainty about cladistic
relationships within the family and the paucity of anatomical data for the
Labiatae. We have selected as outgroups those few non-ingroup taxa of subfam-
ily Lamioideae for which we have collected anatomical data (Ajuga reptans,
Pogostemon cablin, Prostanthera rotundifolia, Scutellaria [7 spp.], Teucrium
[2 spp.], and Trichostema [2 spp.]), plus subfamily Nepetoideae as a whole.
The latter must be included because, as discussed above, there is no evidence
that subfamily Lamioideae is monophyletic. If it were paraphyletic by exclusion
of subfamily Nepetoideae, the latter might be more closely related to tribe
Lamieae (the ingroup) than are some of the other selected outgroups. Subfamily
Nepetoideae is represented by our own data for M onarda fistulosa and Blephilia
hirsuta and by published data for Zhumeria Rech. f. & Wendelbo (Bokhari &
Hedge, 1976), tribe Meriandreae (Bokhari & Hedge, 1971), and subtribe Hyp-
tidinae (Rudall, 1979, 1980). For stomatal characters (see TABLE 1) ten other
genera can be added as representatives of subfamily Nepetoideae, two other
species can be added as representatives of Pogostemon, and Dysophylla Blume
and Anisomeles can be added to the list of outgroups. For subsessile glands,
nine other genera can be added as representatives of subfamily Nepetoideae
(Bruni & Modenesi, 1983; Werker, Putievsky, & Ravid, 1985; Werker, Ravid,
& Putievsky, 1985). Each of the outgroups is thought to be monophyletic, and
no two of them can be combined into a more inclusive monophyletic group.
For example, there is no evidence that tribe Ajugeae sensu Bentham, repre-
sented in this study by Ajuga, Teucrium, and Trichostema, is monophyletic.
The outgroups must be used in combination, because even the more distant
outgroups may affect polarity assessment in the ingroup (Maddison et a/., 1984).
The analysis is complicated, however, by the lack of resolution of phylogenetic
relationships among the outgroups and by uncertainty about which outgroups
are most closely related to the ingroup. If a state that occurs in the ingroup
occurs in none of the outgroups, it is clearly derived within the ingroup, but
if it occurs in some outgroup taxa (the most frequent situation), polarity as-
sessment is more problematic. The outgroup-substitution approach (Donoghue
& Cantino, 1984) is applicable to this situation but difficult to apply here
because of the large number of plausible outgroup combinations that must be
considered. Moreover, both this approach and the global parsimony approach
of Maddison and colleagues (1984) require a full cladistic analysis using all
verse section: e, Brazoria pulcherrima (Kessler 5865); f, Physostegia angustifolia (Cantino
1058); g, Physostegia godfreyi (Cantino 1054). h, internal glandular trichome, transverse
section, Pogostemon cablin (Cantino 1262). Scale bars = 60 wm (a-g) and 20 ym (h).
24 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
available characters, whereas the intent here is simply an evaluation of the
possible phylogenetic significance of a few specific characters.
An alternative method will therefore be used to evaluate polarity of characters
that vary within the outgroups. This approach, developed by Frohlich (1983,
1987), involves calculation of the probability that the commonest state among
the outgroups could parsimoniously be treated as ancestral in the ingroup if
the cladistic relationships of the outgroups to each other and to the ingroup
were known. Frohlich has developed an algorithm that considers all possible
arrangements of the outgroups, determines for each arrangement which state
ofa binary character it is most parsimonious to consider as ancestral within
the ingroup, and then calculates the percentage of arrangements that assign
each state as ancestral in the ingroup. This can be converted to probability if
all outgroup combinations are assumed to be equally probable, a necessary
assumption when one is ignorant of the true outgroup relationships. Thus,
according to Frohlich, if a state occurs in only one of seven outgroups, the
probability is 0.909 that the alternative state could parsimoniously be treated
as ancestral within the ingroup if outgroup relationships were known (1.e., 90.9%
of the outgroup arrangements yield this polarity assessment, while the rest yield
an equivocal one). Frohlich’s “‘tree-count method” turns out to be helpful in
determining the polarity of several characters (see below).
A derived character state that occurs in some, but not all, members of a
monophyletic group is called a nonuniversal derived state (Cantino, 1985b).
A nonuniversal derived state shared by two or more groups, each known to
be monophyletic on the basis of other characters, provides evidence that these
groups together constitute a clade, but it is weaker evidence than if monophyly
is inferred on the basis of a synapomorphy that occurs in all members of the
clade it delimits (Cantino, 1985b). Both synapomorphies and shared nonuni-
versal derived states are used in the following analysis.
TRA MATION SERIES
Most characters examined in this study are binary. Of the multistate char-
acters only two, stomatal type and subsessile glandular trichomes, display vari-
ation of phylogenetic significance at the suprageneric level.
Based on ontogenetic studies (see FiGure 1), a transformation series for
stomatal types is proposed (FiGurE 7a). The anomocytic type is the simplest
ontogenetically. The diacytic and diallelocytic stomata form a transformation
series from the anomocytic type. The anisocytic and paracytic types, which
form a second transformation series from the anomocytic type, share the initial
step in their ontogenies (FIGURE 1) but diverge after that point.
Bosabalidis and Tsekos (1984) studied the ontogeny of subsessile glandular
trichomes in Origanum L. They found that a single initial protodermal cell
divides to give in succession what we have called trichome types 2, 4, 5, and
6. Based on this study, as well as on a comparison of the structural complexity
of the mature trichomes, a transformation series for the subsessile glandular
trichomes is hypothesized (FIGURE 7b). Type | is the simplest structurally and
ontogenetically, while type 10 is the most complex. Tangential walls occur in
the heads of types 6, 7, 9, and 10. Partial radial walls occur only in types 7
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 25
Anisocytic
-2-@-@)
a Diacytic Dialellocytic-1 Dialellocytic-2
Anomoc ytic
2
O-D 3-8
b 8 9 10
FicureE 7. Hypothesized transformation series: a, stomatal types; b, subsessile glan-
dular trichomes.
and 10. Types 5, 6, and 7 differ from types 8, 9, and 10 in that the former
have no more than one secondary radial wall on a given side of any primary
radial wall and lack tertiary radial walls, while the latter have more than one
secondary radial wall on a given side of at least one primary radial wall and/
or have tertiary radial walls. Types 9 and 10 trichomes can develop by more
than one ontogenetic pathway and are therefore not necessarily homologous
in all taxa in which they occur.
CHARACTER POLARITY
Anomocytic and diacytic stomata are widely distributed in both the Ver-
benaceae and the Labiatae (see TABLES |, 2). Diallelocytic-1 stomata are wide-
26 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
spread among the Labiatae, including the outgroups to tribe Lamieae. Dial-
lelocytic-2 stomata are known to occur in four genera of tribe Lamieae plus
four species among the outgroups (Scutellaria elliptica, S. ovata, Ocimum basi-
licum, and Plectranthus australis). Both diallelocytic types appear to be rare
in the Verbenaceae. The anisocytic and paracytic types occur mainly in the
Verbenaceae and the primitive Labiatae (i.c., tribes Prostanthereae and Aju-
geae).
The above distribution suggests that although both diallelocytic types are
probably derived within the Labiatae, the diallelocytic- | type 1s plesiomorphic
in tribe Lamieae. Based on Frohlich’s (1987) probability table, there is a prob-
ability of over 0.984 that the diallelocytic-2 type can parsimoniously be hy-
pothesized to be derived within tribe Lamieae. This calculation is based on its
occurrence in two of the seven examined species of one outgroup (Scutellaria)
and in two of the four examined species of another (subfam. Nepetoideae), and
on its absence from the other five outgroups. The many other species of subfam.
Nepetoideae in TABLE | (in none of which were diallelocytic-2 stomata reported)
are ignored in this analysis because the sample for each consisted only of
published drawings. If these species were to be included, the probability that
the diallelocytic-2 type is derived in the Lamieae would be even greater.
Among the subsessile glandular trichomes (see TABLE 4), types 4 and 5 are
common throughout the Labiatae and thus plesiomorphic within tribe Lamie-
ae. Types 1, 2, and 3b are of scattered occurrence but apparently do not occur
in the North American Melittidinae. Of the more complex glands, types 6, 8,
and 9 appear to be too common in the outgroups, particularly in subfamily
Nepetoideae in the case of types 6 and 9 (Werker, Putievsky, & Ravid, 1985:
Werker, Ravid, & Putievsky, 1985), to permit polarity assessment in the in-
group. Glands with partial radial walls (types 7 and 10) were found only in the
Lamieae, however, where they apparently represent a derived state.
The saclike idioblasts in the mesophyll of Brazoria and Physostegia appear
to be unique to these genera and thus represent a synapomorphy. Undiffer-
entiated mesophyll has been observed only in Physostegia godfreyi and may
represent an autapomorphy of the species. Bundle-sheath extensions are absent
(state a of character 6, TABLE 5) in some Lamieae (one species of Brazoria and
five of Physostegia) but are present in all but one outgroup. Similarly, keels on
the secondary veins are absent (states a—c of character 7, TABLE 5) in some
Lamieae (Brazoria and some species of Macbridea and Physostegia) but present
in all but one outgroup. According to Frohlich’s (1987) probability table, there
is a 0.909 probability that a state occurring in six of seven outgroups can be
parsimoniously hypothesized to be ancestral within the ingroup. If this level
of probability is deemed acceptable, absence of bundle-sheath extensions and
of secondary-vein keels can tentatively be treated as derived in the Lamieae.
PHYLOGENETIC HYPOTHESES
Since the samples of both ingroup and outgroup taxa are small and only leaf
anatomy is being considered, phylogenetic hypotheses must be considered very
preliminary. The characters that offer apparent synapomorphies should be
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 21
MACBRIDEA BRAZORIA PHYSOSTEGIA
SACLIKE IDIOBLASTS
— LOSS OF BUNDLE-SHEATH
EXTENSIONS
— TYPE LO GLANDULAR TRICHOMES
nuf 2 2° VEINS LACK KEELS
FicurE 8. Cladogram showing hypothesized phyl lationships between Bra-
zoria, Physostegia, and Macbridea. Solid bar = Sa dashed bars = shared
nonuniversal derived states.
examined in a broader survey of both tribe Lamieae and the outgroups. The
latter may force reassessment of character polarity in some cases, while ex-
pansion of the ingroup sample may increase the membership of certain clades.
Moreover, other sets of characters may support conflicting hypotheses. At the
very least, however, this analysis should help focus future investigations on
particular characters and taxa
Shinners (1953) suggested that Brazoria and Physostegia are close relatives.
In the numerical phenetic analysis of El-Gazzar (1969), these two genera paired
on the phenogram at a very high similarity level. Until now, however, no strong
evidence that they form a monophyletic group (i.e., the occurrence of synapo-
morphies) has been reported. In the present study an apparent synapomorphy—
the occurrence of saclike idioblasts in the mesophyll of all examined species
of both genera—has been documented. No other taxon in the Lamiales is
known to have this feature. Weaker additional support for the monophyly of
this clade is provided by a shared nonuniversal derived state (Cantino, 1985b),
absence of bundle-sheath extensions. As discussed above, there is a 0.909
probability that this state can parsimoniously be hypothesized to be derived
since it occurs in one of seven outgroups (Prostanthera).
Cantino (1982) suggested that Brazoria, Physostegia, and Macbridea may
form a monophyletic subgroup within the Melittidinae. No synapomorphy was
found to corroborate this hypothesis, but it is supported by two nonuniversal
derived states (FIGURE 8). Type 10 glandular trichomes, the most complex
subsessile glands, occur in all species of Brazoria and Macbridea and two species
of Physostegia, but they were not observed in any other taxa of either the
ingroup or the outgroup. Weaker additional support for the Brazoria-Physoste-
gia-Macbridea clade is provided by another nonuniversal derived state that
28 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
occurs 1n one outgroup as well as in this clade, but nowhere else in the ingroup.
Secondary veins lack keels in three species of Brazoria (and may or may not
lack them in the fourth), Macbridea alba, most species of Physostegia, and the
outgroup species Prostanthera rotundifolia. This is probably a reflection of the
relatively succulent nature of the leaves of these taxa. As discussed above, there
is a 0.909 probability that the loss of secondary-vein keels can be parsimoni-
ously hypothesized to be derived in the ingroup. However, the independent
evolution of this character state in the outgroup and ingroup indicates that it
may be particularly subject to parallelism, which reduces its value as a phy-
logenetic indicator (Gosliner & Ghiselin, 1984). If absence of secondary-vein
keels is indeed a function of leaf succulence, it can be expected in other suc-
culent Labiatae that have not yet been examined. Such a character state can
provide only weak support for the Brazoria-Physostegia-Macbridea clade.
Leaf anatomy has provided strong support for a Physostegia- Brazoria clade
and weaker support for a clade composed of these plus Macbridea. The question
still remains whether there is any anatomical evidence to link these three genera
to the rest of subtribe Melittidinae or to other genera within tribe Lamieae.
Two ingroup genera, Galeobdolon and Synandra, are suggested as possible
relatives of the Physostegia-Brazoria-Macbridea clade on the basis of shared
nonuniversal derived states; an expanded survey of the Lamieae may reveal
other relatives. Diallelocytic-2 stomata are shared by Physostegia, Brazoria,
Macbridea, and Galeobdolon. Subsessile glands with partial radial walls (types
7 and 10) occur in Physostegia, Brazoria, Macbridea, and Synandra
Leaf anatomy has provided no evidence that subtribe Melittidinae ; is Mono-
phyletic. The four North American genera may form a clade, but Galeobdolon
(which has never been treated as belonging to the subtribe) is no less strongly
implicated than Synandra as the sister group of the Physostegia-Brazoria-
Macbridea clade. No anatomical characters suggest a relationship between
Chelonopsis or Melittis and the rest of the Melittidinae. Since leaf anatomy,
floral morphology, and karyology (Cantino, 1985a) do not provide any con-
vincing evidence that subtribe Melittidinae is monophyletic, nor does any other
character we are aware of, its abandonment should be seriously considered.
ACKNOWLEDGMENTS
We would like to thank Robert Kleiman, of the U. S. Department of Agri-
culture, for permission to cite his unpublished data on the fatty-acid compo-
sition of the seed oils of the Verbenaceae, and Michael W. Frohlich for allowing
us to draw heavily on his unpublished manuscript in our cladistic analysis. We
are also grateful to Willard W. Payne and Hanne Rasmussen for discussing
with us their contrasting viewpoints on stomatal ontogeny. This research was
supported by National Science Foundation grant BSR 83-06878.
LITERATURE CITED
Asu-Asas, M. S. 1984. Phylogenetic implications of leaf anatomy in subtribe Melit-
peng (Labiatae) and related genera. Unpubl. M. S. Thesis, Ohio Univ., Athens,
Ohi
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 29
ARNOLD, E. N. 1981. ents phylogenies at low taxonomic levels. Z. Zool. Syst.
Evol.-Forsch. 19: 1-35.
AZIZIAN, D., & D. F. sei 1982. Anatomical, cytological and phytochemical studies
on Phlomis L. and Eremostachys Bunge (Labiatae). J. Linn. Soc., Bot. 85: 249-281.
BENTHAM, G. 1832-1836. Labiatarum genera et species. Ridgway & Sons, London.
——. 1848. Labiatae. Pp. 27-603 in A. DE CANDOLLE, = Prodromus systematis
naturalis regni vegetabilis. Vol. 12. Treuttel et Wirtz, Pan
1876. Labiatae. Pp. 1160-1223 in G. BENTHAM & 7, D. Hooker, Genera
plantarum. Vol. 2. Reeve and Co., London
BERLYN, G. P., & T. P. MIKSCHE. 1976, Botanical microtechnique and cytochemistry.
Iowa State Univ. Press, Ames, Iowa.
Bokuarl, M. H., & I. C. HEDGE. 971. Observations on the tribe Meriandreae of the
Labiatae. Notes Roy. Bot. Gard. Edinburgh 31: 53-67.
1976. Zhumeria (Labiatae): anatomy, taxonomy and affinities. Iran.
t. 1: pale 10.
Lane A., & I. Tsekos. 1982. Glandular scale development and essential oil
secretion in Origanum dictamnus L. Planta 156: 496-504.
& Glandular hair formation in Origanum species. Ann. Bot.
(London) 53: ale.
Briguet, J. 1895-1897. Labiatae. Pp. 183-375 in A. ENGLER & K. PRANTL, eds., Die
natiirlichen Pflanzenfamilien. Vol. 4, part 3a. W. Engelmann, Leipzig
Brunt, A., & P. Mopenesi. 1983. Development, oil storage and sere of peltate
trichomes i in Thymus vulgaris (Lamiaceae). Nordic J. Bot. 3: 245-251.
CANTINO, P. D. 1979. ea godfreyi (Lamiaceae), a new species from northern
Florida. Rhodora 81: 409-4
82. A monograph i - genus Physostegia (Labiatae). Contr. Gray Herb.
211: 1-105.
_ 1985a. Chromosome studies in subtribe Melittidinae (Labiatae) and systematic
eta Syst. Bot. 10:
1 . Phylogenetic inference from nonuniversal derived character states. Ibid.
1 19-12
—— &R. a “SANDERS. 1986. Subfamilial classification of Labiatae. Syst. Bot. 11: 163-
185
Conn, B. J. 1984. A taxonomic revision of Prostanthera Labill. section Klanderia
(F. V. Muell.) Benth. (Labiatae). J. Adelaide Bot. Gard. 6: 207-348.
CuTLer, D. F. 1978. Applied plant anatomy. Longman, London
DONOGHUE, M. J., & P. D. Cantino. 1984. The logic and rere of the outgroup
substitution approach to cladistic analysis. Syst. Bot. 9: 192-202
ELDREDGE, N., & J. CRACRAFT. 1980. Phylogenetic patterns and the evolutionary pro-
cess. Columbia Univ. Press, New York.
E_-Gazzar, A. 1969. A taxonomic study of Labiatae and cae genera. Unpubl. Ph.D.
dissertation, Southampton Univ., Southampton, Engla
_ WATSON. 1968. Labiatae: taxonomy and aca nie to Puccinia menthae
Pers. New Phytol. 67: 739-743.
& ——. 70. A taxonomic study of Labiatae and related genera. Ibid. 69:
451-486.
EpLinc, C. 1942. The American species of Scutellaria. Univ. Calif. Publ. Bot. 20: 1-
ERDTMAN, G. 1945. Pollen morphology and plant taxonomy. IV. Labiatae, Verbenaceae
and Avicenniaceae. Svensk Bot. Tidskr. 39: 279-285.
gee J.S. 1982. Outgroups and parsimony. Syst. Zool. 31: 328-334.
FROHLICH, M. W. 1983. The common-is- ea rule: how common is common?
ee J. Bot. 70(5, part 2): 113,
1987. Common-is-primitive: a partial dion by tree counting. Syst. Bot.
12: in press.
30 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
FRYNS-CLAESSENS, E., & W. VAN COTTHEM. 1973. Anew classification of the ontogenetic
types of stomata. Bot. Rev. (Lancaster) 39: 71-138.
Goprrey, R. K., & J. W. Wooten. 1981. Aquatic and wetland plants of southeastern
United States. Vol. 2. Dicotyledons. Univ. Georgia Press, Athens, Georgia.
Gos.iner, T. M., & M. T. GuiseLin. 1984. Parallel evolution in opisthobranch gas-
tropods and its implications for phylogenetic methodology. Syst. Zool. 33: 255-274.
HAGEMANN, J. M., F. R. EARLE, & I. A. WoLrr. 1967. Search for new industrial oils.
XIV. Seed Gils of Labiatae. Lipids 2: 371-380.
HENNIG, W. 1966. Phylogenetic systematics. Univ. Illinois Press, Chicago.
HOLMGREN, P. K., W. KEUKEN, & . SCHOFIELD. 1981. Index herbariorum. Part 1.
Herbaria of the world. ed. 7. Bohn, Scheltema, & Holkema, Utrecht.
Huana, T. C., & W. T. CHENG. 1971. A preliminary revision of Formosan Labiatae
(1). Taiwania 16: 157-174.
INAMDAR, J. A. 1969. pear nea some Verbenace-
ae. fee Bot. (London) 33: 55-66.
D.C. BHaTr. 1972. en and development of stomata in some Labiatae.
Ibid. 36: 335-344.
JOHANSEN, D. A. 1940. Plant microtechnique. McGraw-Hill, New
JUNELL, S. 1934. Zur ene und Systematik der Wren und
Labiaten. Symb. Bot. Upsal. 4: 1-219.
KHALEEL, T. F., & A. S. NALINI. 1972. Embryology of Lantana aculeata L. var. nivea
Bailey. Curr. Sci. 41: 491-494.
Koorman, P. 1972. The occurrence of iridoid glycosides in the Labiatae. Acta Bot.
Neerl. 21: 417-427.
KRAL,R. 1983. A report on some rare, threatened, or endangered forest-related vascular
plants of the South. Vols. 1, 2. U.S.D.A. Forest Serv. Techn. Publ. R8-TP2. Atlanta,
Georgia.
Mapoison, W. P., M. J. DonoGuug, & D.R. MADDISON. 1984. Outgroup analysis and
03
kh ]
SI
MaHEsHwari, J. K. 1954. Floral morphology and yology of Lippia nodiflora Rich.
Phytomorphology 4: 217-230.
Martin, A. C. 1946. The comparative internal morphology of seeds. Amer. Mid],
Naturalist 36: 513-660.
Metcacre, C. R., & L. CHALK. 1950. Anatomy of the dicotyledons. Vol. 2. Oxford
4
Pant, D. D. 1965. On the ontogeny of stomata and other homologous structures. Plant
Sci. Ser. Allahabad 1: 1-24.
Payne, W. W. 1970. Helicocytic and allelocytic stomata: unrecognized patterns in the
Dicotyledonae. Amer. J. Bot. 57: 140-147.
Stomatal Rene in embryophytes: their evolution, ontogeny and in-
terpretation Taxon 28: 117-132.
RamayyYA, N., & J. RAo. om Range of structural and ontogenetic stomatal variations
in an species of Ocimum (Labiatae). Curr. Sci. 38: 79-82.
Se H. 1981. Terminology and cae oe of — and stomatal devel-
ment—a critical survey. J. Linn. Soc., Bot. 83: 212.
nea P. 1979, way ne anatomy of Eriope, a mie genus of Labiatae.
J. Linn. Soc., Bot. 78: 157-180.
1980. Leaf cute of the subtribe Hyptidinae (Labiatae). bid. 80: 319-340.
Sakal, W. S. 1973. Simple method for differential staining of paraffin embedded plant
material using toluidine blue O. Stain Technol. 48: 247-249.
1987] ABU-ASAB & CANTINO, LEAF ANATOMY 31
SANDERS, R. W., & P. D. CAnTINO. 1984. Nomenclature of the subdivisions of the
Lamiaceae. Taxon 33: 64-72.
ScHNARF, K. 1918. Beitrage zur Kenntnis der Samenentwicklung der Labiaten. Kaiserl.
kad. Wiss. Wien, Math.-Naturwiss. KI., Denkschr. 94: 211-2
Suan, G. L., & A. C. Narpu. 1983. Trichomes on leaves of some Lamiaceae. Geo-
phytology 13: 165-176.
SHINNERS, L. H. 1953. Synopsis of the genus Brazoria (Labiatae). Field & Lab. 21: 153,
154.
SOLEREDER, H. 1908. of the dicotyled Clarendon Press, Oxford.
Spies, J. J. 1984a. ae sac development in some South African Lantana species
(Verbenaceae). Bothalia 15: 161-166.
mbryo sac eee in some South African Lippia species (Ver-
benaceae) S. African J. Bot. 3: 120-124.
& C. H. Stirton. 1982. Embryo sac development in some South African
cultivars Es Lantana camara. Bothalia 14: 113-117.
Stace, C. A. 1973. The significance of the leaf Nea in the taxonomy of the
Combretaceae. IV. The genus Combretum in Asia. J. Linn. Soc., Bot. 66: 97-115.
STEVENS, P. F. 1980. Evolutionary polarity of character states. Annual Rev. Ecol. Syst.
11: 333-358.
STEVENS, R. A., & E. S. Martin. 1978. A new ontogenetic classification of stomatal
types. J. Linn. Soc., Bot. 77: 53-64
TATACHAR, T. 1940. The development of the embryo sac and formation of haustoria
in Lantana indica Roxb. and Stachytarpheta indica Vahl. J. Indian Bot. Soc. 19:
D
THEOBALD, W. L., J. L. KRAHULIK, & R. C. Rotiins. 1979. Trichome description and
Acheter Pp. 40-53 in C. R. MetrcaLre & L. CHALK, eds., Anatomy of the
dicotyledons. ed. 2. Vol. 1. Clarendon Press, Oxford.
THIRUMARAN, K., & K. K. LAKSHMANAN. 1984. Embryological studies on Priva cor-
difolia. Acta Bot. Indica 12: 103-106.
VesqueE, M. J. 1889. De l’emploi des eae anatomiques dans la classification des
végétaux. Bull. Soc. Bot. France, II. 36: 41-77.
Watrous, L. E., & Q. D. WHEELER. 1981. The out-group comparison method of
character analysis. Syst. Zool. 30: 1-11.
Werker, E., E. Putievsky, & U. Ravip. 1985. The essential oils and glandular hairs
in different chemotypes of eae” vulgare L. Ann. Bot. (London) 55: 793-801.
WERKER, E., U. RAvip, & E. PUTI y. 1985. Structure of glandular hairs and iden-
tification of the main cree of their secreted material in some species of the
Labiatae. Israel J. Bot. 34: 31-45.
Wier, E.O. 1981. Phylogenetics: the theory and practice of phylogenetic systematics.
John Wiley and Sons, New York.
WILKINSON, H. P. 1979. The plant surface (mainly leaf). Pp. 97-165 in C. R. METCALFE
& L. CHALK, eds., Anatomy of the dicotyledons. ed. 2. Vol. 1. Clarendon Press,
rd.
WuNDERLICH, R. 1967. Ein Vorschlag zu einer natiirlichen Gliederung der Labiaten
auf Grund der Pollenkérner, der Samenentwicklung und des reifen Samens. Oesterr.
Bot. Z. 114: 383-483.
Zoz, I. G., & V. I. Litvinenko. 1979. On the division of the family Lamiaceae Juss.
into natural groups. (In Russian.) Bot. Zurn. (Moscow & Leningrad) 64: 989-997.
32 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
APPENDIX |. Abbreviated collection data for
voucher specimens.
Ajuga reptans L. Ohio, Athens Co., Athens, Cantino 1217.
Blephilia hirsuta (Pursh) Bentham. Ohio, Vinton Co., Lake Alma State Park, Cantino
& Abu-Asab 1249.
Brazoria arenaria Lundell. Texas: oe Co., Aransas National Wildlife Refuge, Kess/er
5773; Refugio Co., Kessler 577
Brazoria eilehennma Lundell. a Leon Co., Kessler 5862, 586
Brazoria scutellarioides Engelm. & Gray. Texas, Travis Co. aneaee 761 68 (TEX), 76179
(TEX
Brazoria truncata (Bentham) Engelm. & Gray. Texas, Live Oak Co., 2 km SW of
tsett, Sanders 76122 (TEX
C oe forrestii Anthony. China, Szechwan Prov., Rock 5515 (A).
ere moschata Miq. Japan, Prov. Iwashiro, Pref. Fukushima, Furuse s.n., 7-IX-
957 (a).
Galeobdolon luteum Hudson. Ohio, Athens Co., greenhouse plant from commercial
source, Cantino 1271.
Lamium purpureum L. Ohio, Athens Co., Athens, Cantino 1214.
Leonurus cardiaca L. Michigan, Ingham Co., East Lansing, Cantino 1224.
Macbridea alba Chapman. Florida, Bay Co., E of Callaway, Godfrey 79884.
Macbridea caroliniana (Walter) Blake. North Carolina, Pender Co., near Long Creek,
Cantino 4.
Marrubium vulgare L. Ohio, Athens Co., Athens, in garden, Cantino 1242.
nes ae eats be Czechoslovakia, Brinn [Brno], Piskoi 667 (Gu); France, be-
Capendu and Moux, Neyraut s.n., 12-VI-1888 (GH
iene L. Ohio, Vinton Co., Vinton Twp., Cantino & Abu-Asab 1251.
Physostegia angustifolia Fern. on aa, St. Tammany Parish, 10 mi SW of Covington,
Cantino
as digitalis Small. Louisiana, Rapides Parish, 3 mi N of Elizabeth, Cantino
1070 (G
ie: sodfreyi Cantino. Florida, Gulf Co., 12 mi S of Wewahitchka, Cantino 1054.
Ph He leptophylla Small. North Carolina, Hertford Co., 4 mi W of Winton, Cantino
71
a longisepala Cantino. Louisiana, Lafayette Parish, garden plant transplanted
from vicinity of Mauriceville, Orange Co., Texas, Vincent 42
Physostegia purpurea (Walter) Blake. North Carolina, Harnett Co., 3 mi SE of Bunnlevel,
Physostegia virginiana (L.) Bentham subsp. praemorsa (Shinners) Cantino. North Car-
olina: Transylv ania Co., 4 mi SW ae ke Toxaway, Cantino 946; Montgomery Co.,
0.5 mi N of Blaine, Canine 943 (G
ge aie virginiana (L.) Bentham has virginiana. Ohio, Athens Co., York Twp.,
Cantino 1260.
Pogostemon ae Bentham. Ohio, Athens Co., greenhouse plant from commercial
urce, Cantino
i 5 rv R. Br. Ohio, Athens Co., greenhouse plant from commercial
aria gia no 1261,
elliptica Muhlenb. Ohio, Jackson Co., Lake Alma State Park, Cantino & Abu-
yee 1222.
*Vouchers at BHO unless otherwise indicated. Herbarium abbreviations follow Holmgren ef al.
(1981)
1987] ABU-ASAB & CANTINO, LEAF ANATOMY a3
Scutellaria incana Biehler. Ohio: Athens Co., Athens, Cantino & Abu-Asab 1236, Hock-
ing Co., Ward Twp., Cantino & Abu- Asab 1247
Scutellaria integrifolia L. Ohio, Vinton Co., Lake Alma State Park, Cantino 1227.
Scutellaria lateriflora L. Ohio. Vinton Co.: Lake Alma State Park, Cantino & Abu-Asab
- Lake Hope State Park, Cantino & Abu-Asab 1257.
Scutellaria nervosa Pursh. Ohio: Athens Co., Athens, Cantino 1231; Perry Co., Monroe
oung s.n. (no voucher
Scutellaria ovata Hill. Ohio, Athens Co., Athens Twp., Cantino 1232.
ciate serrata Andrz. Ohio. Vinton Co.: Brown Twp., Cantino & Abu-Asab 1219;
Lake Alma State Park, Cantino & Abu-Asab 1221.
Stachys riddellii House. Ohio, Vinton Co., Lake Alma State Park, Cantino 1229, 1230.
Stachys tenuifolia Willd. Ohio. Athens Co.: Athens, Cantino 1235, Waterloo Twp.,
Cantino & Abu-Asab 1253. Vinton Co., Lake Hope State Park, Cantino & Abu-Asab
1256.
Synandra hispidula (Michaux) Baillon. Ohio, Morgan Co., Union Twp., Cantino 1151.
Teucrium canadense L. Ohio, Athens Co., Dover Twp., Cantino & Abu-Asab 1243,
1244.
Teucrium chamaedrys L. Ohio, Athens Co., Athens, in garden, Cantino 1240.
Trichostema dichotomum L. Ohio, Perry Co., Monroe Twp., J. Young s.n., 24-VIUII-
Trichostema lanceolatum Bentham. Ohio, Athens Co., greenhouse plant from commer-
cial source, Cantino 125
APPENDIX 2. Classification of subsessile glandular
trichomes in the Labiatae.
Type 1. | Head composed of one cell.
Type 2. Head composed of two cells (FiGuRE 5f1).
Type 3. Head composed of three cells.
3a. Head divided by two transverse walls (FIGURE 5a6).
3b. Head divided by three radial walls (Figure 5f2, g1).
Type 4. Head composed of four cells (FicureE Sal, bl, dl, el, f3, g2, hl).
Type 5. | Head of more than four cells, usually divided by four primary radial
walls that are more or less perpendicular to each other; tertiary and
tangential walls absent; no more than one secondary radial wall
arising on a given side of any primary radial wall (FIGURE 5a2-5,
b2-cl-c24d2.:d3.ie2, 63. NJ).
Type 6. Asin Type 5, but with tangential walls present (FIGURE 5d4).
Type 7. As in Type 6, but with partial radial walls present (FIGURE 5e3).
Type 8. | Head of more than four cells; tertiary radial walls present and/or
more than one secondary radial wall arising on the same side of at
least one primary radial wall; tangential walls absent (FIGURE 5a7).
Type 9. Asin Type 8, but with tangential walls present, partial radial walls
absent (FIGURE 5a8, c
Type 10. As in Type 9, but with partial radial walls present (FIGURE 5a9,
b3).
*Cell-wall configurations are defined in the text and illustrated in Ficure 4.
34 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
APPENDIX 3. Leaf histological characters of
possible taxonomic value.
Mesophyll differentiation: a, undifferentiated; b, bifacial; c, isobilateral.
. Compactness of palisade cells: a, compact; oose
. Idioblasts in mesophyll: a, absent; b, saclike: ¢ resembling internal glandular tri-
chomes.
. Fibers associated with midrib: a, absent; b, present only on adaxial side of midrib:
c, present only on abaxial side of midrib; d, present on both sides of midrib.
. Collenchyma associated with midrib: a, absent; b, present only on abaxial side of
midrib; c, present on both sides of midrib.
. Bundle-sheath extensions: a, absent; b, present only on abaxial side of bundle; c,
present on both sides of bundle.
. Prominence of keels associated with vascular bundles as viewed in transver ul
of lamina: a, keels absent; b, secondary veins lacking keels, midrib keel protruding
slightly; c, secondary veins lacking keels, midrib keel protruding greatly; d, secondary
veins keeled, midrib keel protruding slightly; e, secondary veins keeled, midrib keel
protruding greatly.
WN —
lon way aa
—
Erratum—The Pogostemon used in this study was P. heyneanus Bentham, not
P. cablin Bentham.
1987] ROSATTI, PONTEDERIACEAE ye)
THE GENERA OF PONTEDERIACEAE IN THE
SOUTHEASTERN UNITED STATES!
THOMAS J. ROSATTI’
PONTEDERIACEAE Kunth in Humboldt, Bonpland, & Kunth, Nova Gen.
Sp. Pl. 1: 211 (folio ed.); 265 (quarto ed.). 1816, ‘““Pontedereae,” nom. cons.
(PICKEREL-WEED FAMILY, WATER-HYACINTH FAMILY)
Submersed, emersed, or floating aquatic herbs, sometimes on wet ground
because of lowered water levels. Stems sympodial, successive axes terminating
in inflorescences, stout or elongate. Juvenile leaves, especially if submersed,
usually sessile and linear. Adult leaves simple, alternate [or whorled in Hy-
drothrix]; stipulate or exstipulate; the bases mostly sheathing, either open or
fused basally around the stem; sessile or petiolate, the petioles sometimes
inflated: the blades linear [filiform in Hydrothrix] to orbicular, sometimes
sagittate or cordate, the veins parallel, usually arching. Inflorescence a terminal
spike, raceme, panicle, or single flowered; sessile or pedunculate, subtended
and enclosed to various degrees by a sheathing bract that is sometimes sur-
mounted by a variously reduced petiole and/or blade, each flowering stem (1.e.,
that which is not part of the sympodium) also bearing a single leaf that some-
Prepared fo ic Fl fthe Sout! United States, a long-term project made possible
by grants aan the National Science Foundation and at this Neh supported by BSR-8303100 and
BSR-8415637 (Norton G. Tse ie investigator), under which tt
BSR-8415769 (Carroll E d, Jr., ipal investigator). ae omen itis 113th i in ‘the series,
follows the format ee ae in the ae one (Jour. Arnold Arb. 39: 296-346. 1958) and continued
primarily on the plants of this area, with info rmation about extraregional members of a family or
genus in brackets [ ]. References I have not verified are marked with an asterisk.
I am indebted to Norton Miller and Carroll Wood for the many ways they contributed to this
resent.
Florida, Tennessee, Alabama, Mississippi, Arkansas, and Louisiana. The descriptions are based
and herbaria pee with the New a State Museum and Harvard Universit
materials collected by Carroll Wood and Richard J. Eaton and dissected by Wood.
This treatment is published as Contribution Number 491 of the New York eee Science Service.
Biological Survey, New York State Museum, The State Education Department, Albany, New York
12230.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 35-71. January, 1987.
36 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
times differs from the others. Flowers perfect (some species of Eichhornia and
Pontederia tristylous); perianth petaloid, biseriate, usually funnelform io sal-
verform [parts nearly distinct in Monochoria], zygomorphic, subactinomor-
phic, or actinomorphic, the 6 [4, rarely 3, in Scholleropsis] lobes imbricate,
often unequal. Stamens usually 6 (in 2 series of 3) or 3 [4 in Scholleropsis], |
in some cleistogamous flowers [and in Hydrothrix], staminodes sometimes
present when stamens fewer than 6; filaments inserted on the perianth tube at
various levels; anthers held at various levels, basifixed, auriculate and some-
what movable on filaments (and therefore appearing dorsifixed), bilocular, with
introrse, longitudinal dehiscence [rarely terminal pores]; pollen bi- or trinu-
cleate when shed, with | to 3 distal or subequatorial colpi. Gynoecium of 3
united carpels; style single, of various lengths (i.e., some species tristylous):
stigma terminal, entire or variously toothed and/or lobed (often 3- or 6-parted);
Ovary superior, with 3 locules, each locule with an axile placenta, or with |
locule and 3 intrusive parietal placentae, or with 1 locule (through abortion of
2 locules) and a terminal placenta (in Pontederia); ovules in each locule nu-
merous (solitary in Pontederia), anatropous, crassinucellar, and bitegmic; nec-
taries septal (lacking in Heteranthera). Fruit a many-seeded, loculicidal capsule
or a |-seeded utricle (in Pontederia). Seeds small, ovoid, at least those in
capsules with longitudinal ridges or ribs; endosperm starchy; embryo axile,
cylindrical, with a terminal cotyledon and a lateral plumule. (Including Het-
erantheraceae J. G. Agardh, Theoria Syst. Pl. 36. 1858, “Heteranthereae.”’)
Type GENUS: Pontederia L.
A small family of fresh-water aquatics comprising about six genera and 30
species, mostly pantropical but extending into the temperate zones as well.
Although of diverse habit (submersed, emersed, or free floating; erect or pros-
trate; rhizomatous, stoloniferous, or neither), the plants are all more or less
obviously sympodial in structure: each successive axis terminates in an inflo-
rescence that may appear to be axillary.
Plants in the family are most readily distinguished by a combination of
characters, including (in addition to the sympodial structure) herbaceous stems
enveloped to various extents by sheathing leaf bases: inflorescences subtended
by a single sheathing bract; usually six petaloid, nongreen tepals in two series
of three, variously connate basally; stamens adnate to the perianth; and superior
ovaries.
Three tribes, each represented in the Southeast, were recognized in the Pon-
tederiaceae by Schwartz (1927, 1930). Although both the Eichhornieae Schwartz
and the Heteranthereae Schwartz have three-locular ovaries (appearing one-
locular at maturity in Hydrothrix Hooker f.) and many-seeded capsules, the
Pontederieae (Pontederia L. and Reussia Endl.) have a single fertile locule and
a one-seeded, indehiscent fruit. Mostly funnelform perianths and six stamens
characterize the monogeneric Eichhornieae; flowers in the Heteranthereae nor-
mally have either mostly salverform perianths and three or fewer stamens
(Heteranthera Ruiz & Pavon, with three stamens of two more or less distinct
kinds; Scholleropsis H. Perr., with three stamens of two kinds; and Hydrothrix,
with one stamen and two staminodes) or initially campanulate but ultimately
1987] ROSATTI, PONTEDERIACEAE 37
spreading to almost rotate perianths of nearly separate tepals and six stamens
(Monochoria Presl). Extraregional genera of Heteranthereae include Scholler-
opsis (one species in Madagascar with four or rarely three tepals), Hydrothrix
(one or two Brazilian species with filiform leaves), and Monochoria (perhaps
five species ranging from northeastern Africa to Manchuria, Japan, and Aus-
tralia; one of these established in experimental rice plots in California, according
to Mason). The monotypic genera Eurystemon E. J. Alex. (Texas and northern
Mexico) and Zosterella J. K. Small (widely distributed in Mexico and North
America) are included here in Heteranthera. Reussia (two or three species in
South America) is treated as a subgenus of Pontederia.
The systematic position of the Pontederiaceae has long been a subject of
controversy. The group is considered by many to be most closely related to
the Philydraceae (e.g., Casper & Krausch, Dahlgren & Clifford, and Thorne)
and/or the Haemodoraceae (e.g., Cronquist; Dahlgren; Simpson, in press) and
has been variously allied at higher levels with, among other families, the Bro-
meliaceae and/or Commelinaceae by some and the Liliaceae by others.
In Thorne’s system the Pontederiineae (Pontederiaceae and Philydraceae)
were included as one of seven suborders in the Commelinales; the Commelini-
florae and the Liliiflorae were placed as far apart as possible among the five
superorders of monocotyledons recognized. Thorne further proposed (p. 100)
that the Pontederiineae and Bromeliineae share a “rather close common origin”
and thought that misplacement of the former with groups included in his
Liliiflorae was probably due to the presence in the Pontederiineae of a petaloid
(although biseriate) perianth that is often connate at the base and mostly zy-
gomorphic, as well as to a cylindrical embryo centrally placed in abundant,
starchy endosperm. Castellanos, on the other hand, considered the Pontede-
riaceae to be related to the Commelinaceae because both exhibit zygomorphy
and androecial reduction. On the basis of starch grains, Czaja recognized three
groups of monocots, one including the Bromeliaceae, Commelinaceae, Hae-
modoraceae, Philydraceae, and Pontederiaceae, and another the Liliaceae and
their close relatives. Likewise, Huber suggested that the superorder Pontederii-
florae (Pontederiales and Philydrales) had more in common with the Brome-
liiflorae, Haemodoriflorae, and Commeliniflorae than with the Liliiflorae.
A number of systematists have considered the Pontederiaceae to be more
closely allied to the Liliaceae than to either the Bromeliaceae or the Comme-
linaceae. Although Bentham (in Bentham & Hooker) thought that the flowers
indicated a close relationship with the Liliaceae, Baillon and Solms-Laubach
(1883a) were among the first to suggest unification with that family. Such a
roposal was later at least tacitly accepted by Schwartz (1930), who nevertheless
considered the floral zygomorphy, androecial reduction, and mealy endosperm
to indicate a relationship with the Commelinaceae and Philydraceae. Hamann
(in Melchior) suggested on embryological (starchy endosperm) and anatomical
(unspecified) evidence that the Pontederiaceae should be separated from the
Liliaceae but maintained as one of five suborders (including the Philydrineae)
comprising the order Liliiflorae (Liliales), although he pointed out that in some
characters the family is in agreement with the Commelinaceae (Commelinales)
and Philydraceae.
38 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
Takhtajan stated that the Pontederiaceae probably originated from liliaceous
stock, because of the presence of septal nectaries and similarities in vegetative
anatomy and embryology (neither specified). Accordingly, he included a uni-
familial Pontederiineae as one of six suborders (between the Haemodorineae
and a unifamilial Philydrineae, the latter considered to be somewhat isolated
but related to the Pontederiineae) in the Liliales, one of 14 orders (the Bro-
meliales and Commelinales among them) comprising the subclass Liliidae.
Cronquist included the Pontederiaceae in his Liliales (subclass Liliidae), far
removed from both the Bromeliaceae (Zingiberidae) and Commelinaceae
(Commelinidae). Dahlgren, on the other hand, incorporated a unifamilial Pon-
tederiales (between the Haemodorales and the Philydrales) as one of 12 orders
with the Bromeliales in his Liliiflorae and placed the Commelinaceae in a
separate superorder (Commeliniflorae). Dahlgren & Clifford envisioned a series
of taxa, the members of which (e.g., Haemodorales, Philydrales, Pontederiales,
Bromeliales, Commelinales) formed a gradual transition between the Liliiflorae
and the Commeliniflorae and combined significant features of both. The Pon-
tederiales were indicated to have substantially more attributes in common with
“core” Liliiflorae (11 of 21) than with ‘“‘core’” Commeliniflorae (3 or 4 of 21)
and a profile of features agreeing most closely with that of the Philydrales.
What little is known about the chemistry of the Pontederiaceae has been
compiled and reviewed by Gibbs, and the following is based largely on that
account. The plants are among only six monocot families (including none of
those discussed here) for which the Maule test (which is positive for all but
weakly lignified tissues) is negative or doubtful. Calcium oxalate crystals, usu-
ally raphides, are present. Although cyanogenesis has been reported in Mono-
choria, Gibbs obtained negative results for Eichhornia speciosa Kunth, Het-
eranthera dubia (Jacq.) MacM., and Pontederia cordata L. He determined that
mucilage was present in Pontederia, doubtfully so in Heteranthera, and absent
in Eichhornia. Gibbs observed strong reactions between ferric ammonium
citrate and the leaves of E. speciosaand P. cordata, indicating the likely presence
of tannins or tanninlike substances; Cronquist (p. 1202) characterized the family
as having “scattered tanniferous cells containing proanthocyanins.” Saponins
are reportedly absent or probably absent from Ejichhornia and Pontederia.
Lipids of Eichhornia crassipes (Mart.) Solms have been analyzed by Laksh-
minarayana and colleagues.
Various kinds of phenolic compounds are represented in the Pontederiaceae.
Caffeic acid, cyanidin, and ferulic acid have been reported in Eichhornia spe-
closa and Pontederia lanceolata Nutt., while p-coumaric and synaptic acids
are evidently known only from the latter (Bate-Smith). Leucoanthocyanins
(which produce anthocyanidins when heated with mineral acids) were indicated
for P. cordata by Gibbs and considered to be abundant in the family by Bate-
Smith. The anthocyanins cyanidin, malvidin, delphinidin, and eichhornin have
been reported in E. crassipes (see Krishnaveni et al.), as has been delphinidin
in P. lanceolata (Bate-Smith). A number of other flavonoids, including the
flavones apigenin and luteolin as well as the flavonols quercetin and isorham-
netin, were isolated from various species of Heteranthera by Horn (1985a).
1987] ROSATTI, PONTEDERIACEAE a9
Lowden compared the phenolic profiles of several genera of Pontederiaceae in
his revision of Pontederia (see discussion of that genus).
Cheadle studied the vessel elements in a number of species (including five
occurring in the Southeast) belonging to several genera of Pontederiaceae and
found that they normally had long, obliquely oriented, scalariform perforation
plates with many perforations and were mostly confined to the roots. Some,
however, indicated a more advanced condition, because of either their structure
(five or fewer perforations in nearly transverse plates) or their location (stems
of Eichhornia crassipes, Heteranthera limosa (Sw.) Willd., and possibly Pon-
tederia cordata). Cheadle concluded that while relatively unspecialized vessel
elements are typical of aquatic angiosperms in general, the more advanced
structure of some in the Pontederiaceae suggested that the family was primi-
tively terrestrial. Nevertheless, he also allied the Pontederiaceae with the Phily-
draceae, even though the vessel elements in this group, which is terrestrial, are
somewhat less specialized.
Anthers in the Pontederiaceae are tetrasporangiate (the normal condition
among angiosperms); they are bisporangiate in the Philydraceae according to
Bhandari, but are tetrasporangiate by Cronquist’s account. The microspore
mother cells undergo successive divisions to form either isobilateral or decus-
sate tetrads (Davis), and the pollen grains are binucleate when shed (Brewbaker;
see, however, Cronquist, who indicated that they are sometimes trinucleate).
The Pontederiaceae were not well known palynologically before Simpson’s
recent electron micrographic (both TEM and SEM) studies of the group (pers.
comm.; 1986), which featured comparisons with the pollen of both the Hae-
modoraceae and the Philydraceae. Despite earlier reports to the contrary (Erdt-
man; Rao & Rao; Simpson, in press), pollen with two furrow-shaped apertures
(orientation yet to be determined) appears to be the only type represented in
the family. In part because this condition is unknown in either the Haemo-
doraceae or the Philydraceae (and presumably elsewhere), its derivation was
considered to have been uniquely shared by members of the Pontederiaceae.
Internal exine structure is variable within the Pontederiaceae, according to
Simpson (in press), but the variation does not correspond well to the tribes
recognized here. For example, what was termed a “modified tectate-columel-
late” exine is shared by species of both the Heteranthereae (Monochoria va-
ginalis (Burman f.) Presl, Scholleropsis lutea H. Perr.) and the Pontederieae
(Pontederia cordata), a “two-layered” exine characterizes members of both the
Heteranthereae (Zosterella dubia (Jacq.) Small = Heteranthera dubia) and the
Pontederieae (Reussia rotundifolia (L. f.) Castell. [here put in Pontederia subg.
Reussia]), and a “one-layered” exine corresponding to the outer layer of the
two-layered type was depicted for genera of the Eichhornieae (Eichhornia cras-
sipes) and the Heteranthereae (Hydrothrix Gardneri Hooker f.). A condition
described as intermediate between the modified tectate-columellate type and
the two-layered type was reported for Heteranthera reniformis Ruiz & Pavon
(see, however, Simpson, 1984).
On the basis of palynological evidence, Simpson also concluded that the
Pontederiaceae are more closely related to the Haemodoraceae than to the
40 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Philydraceae. The distinctive verrucate sculpturing reported for all the Pon-
tederiaceae studied (except Pontederia cordata, which has psilate to scabrate
pollen) is identical to that found in tribe Haemodoreae of the Haemodoraceae.
In addition, two genera of Haemodoreae have a one-layered exine identical to
that of Eichhornia and Hydrothrix, while four others in the Haemodoreae and
all six genera of the Conostylideae have a two-layered exine very similar to
that reported by Simpson for Reussia and Zosterella. The Philydraceae gen-
erally have reticulate grains and typical tectate-columellate internal exine struc-
ture.
The pontederiaceous ovule is anatropous, bitegmic (with both integuments
forming the micropyle), and crassinucellar (Davis). The chalazal megaspore
(see, however, Ono) of a linear tetrad develops into a Polygonum-type mega-
gametophyte in which the synergids (at least in Monochoria hastifolia Presl)
may have a filiform apparatus; the degree to which the three antipodals persist
has been controversial (see Coker, R. W. Smith, W. R. Smith). Endosperm
formation is, according to Davis, helobial, with free-nuclear divisions and
subsequent wall formation normally taking place in both the micropylar and
chalazal chambers (the chalazal chamber remains free-nuclear in Monochoria).
Two haustorial arms are developed laterally in the micropylar chamber in
Monochoria and presumably the remainder of the family as well (Davis). Em-
bryogeny in the Pontederiaceae is of the Asterad type (see, for example, Souéges).
The embryos, along with those of Amomum Roxb. (Zingiberaceae), are re-
portedly unique in their complete extension to both poles of the seed (Martin).
According to information compiled by Davis, the most substantial embryo-
logical differences between the Pontederiaceae and the Philydraceae are that
the former have an amoeboid (vs. glandular) tapetum and unhooked (vs. hooked)
synergids.
Tristyly has been reported in the Pontederiaceae and three other flowering
plant families (Lythraceae, Oxalidaceae, and Rubiaceae: see Vuilleumier). Species
with this condition are characterized by individuals with one of three floral
forms (morphs), each differing in the arrangement of anthers and stigmas. (For
the condition in Pontederia cordata, see FIGURE 1, c-e.) Three levels (short,
medium, and long) are occupied by two groups of anthers and the single stigma;
thus, pollen is partitioned on a pollinator in such a way that its delivery to the
stigma of another flower of the same morph is unlikely. For example, pollen
from short and long stamens of mid-styled flowers normally would not be
delivered to the stigmas of other mid-styled flowers. Transfers of pollen from
anthers to stigmas of the same level are termed legitimate pollinations, while
all intraform and some interform pollinations are illegitimate. Populations of
tristylous Pontederiaceae (and perhaps other families) vary from isoplethy, a
condition in which the three morphs are equally represented, to monomorphy,
in which only a single floral form is present.
The mechanical barrier to pollinations leading to either self-fertilization or
assortative (like genotype) crossing effected by the tristylous condition is nor-
mally accompanied by a physiological self-incompatibility system, as well as
by a marked pollen trimorphism, and since the time of Darwin it has been
thought to promote animal-mediated cross-pollination and subsequent out-
1987] ROSATTI, PONTEDERIACEAE 4]
crossing. Under one argument tristyly and self-incompatibility are mutually
reinforcing, while another holds that the former is secondarily reinforced by
the latter, even though self-incompatibility would appear to be superfluous if
the pollen partitioning effected by tristyly were as effective as 1t appears to be.
It may be more reasonable to suppose that in groups with both conditions,
self-incompatibility, which is relatively widespread in plants in general, evolved
first and is secondarily reinforced by tristyly in the sense that pollen partitioning
would minimize the wasteful placement of pollen on incompatible stigmas.
The selective advantage of tristyly evidently does not involve reduction of
interference on the stigma by illegitimate pollen or adjacent stamens, according
to experiments on Pontederia cordata by Barrett & Glover. The pollen tri-
morphism accompanying tristyly in the Pontederiaceae, which involves dif-
ferences in both pollen size and degree of self-incompatibility, appears to be
physiological and/or developmental in nature and dependent on anther level
(see discussions of Eichhornia and Ponteder
our species of Pontederia (P. cordata, P. outdo f., P. sagittata Presl,
P. subovata (Seub.) Lowden) and three of Eichhornia (E. azurea (Sw.) Kunth,
E. crassipes, E. paniculata (Sprengel) Solms) are tristylous (Barrett, 1978a,
979: Lowden; Richards & Barrett). The condition in Pontederia and E. azurea
is accompanied by physiological self-incompatibility, strong pollen trimor-
phism, and populations in which all three floral morphs are usually represented,
but it is characterized in F. crassipes and E. paniculata by a high degree of
self-fertility, weakly developed pollen trimorphism, and populations that are
frequently monomorphic. Self-fertilizing, semihomostylous (upper set of an-
thers adjacent to stigma) races of each of the tristylous species of Eichhornia,
including F. azurea, have been reported (see also discussions of the genera).
Progeny tests have indicated that the determination of floral morph in Eich-
hornia crassipes is governed by two diallelic loci ee 1977). While the M
locus determines whether styles are midlength (MM or Mm) or long (mm), the
S locus ether they are short (SS or Ss) or nonshort (ss) and 1s epistatic
to the M locus. eae Price, & Shore have assumed that this pattern of
inheritance characterizes all tristylous Pontederiaceae. (See further discussion
under Eichhornia and Pontederia.)
The economic significance of the Pontederiaceae lies chiefly with Eichhornia
crassipes, possibly the world’s most serious aquatic weed; other members of
the family also occur as weeds, especially in rice fields. A number of species
have ornamental value, and many are used in one way or another by fish,
waterfowl, aquatic mammals, and humans.
REFERENCES:
ALEXANDER, E. J. Pontederiaceae. N. Am. Fl. 19: 51-60. 1937. [Eichhornia (4 spp.),
Eurystemon (1 sp.), Heteranthera (5 spp.), Pontederia (5 spp.), Zosterella (2 spp.).]
eS oe The phyllode theory of the monocotyledonous leaf, with special reference to
mical evidence. Ann. Bot. 32: 465-501. 1918. [Leafanatomy of Pontederiaceae
discussed, illustrated; inverted bundles indicate the phyllodic (petiolar) nature of
both pseudolaminate (Eichhornia speciosa, Heteranthera reniformis, Pontederia cor-
data) and linear (H. zosterifolia) leaves.
. Water plants. 2 unnumbered + xvi + 436 pp. Cambridge, England. 1920.
42 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
[Numerous references to Eichhornia, Heteranthera, Pontederia, underwater ripening
of fruit.
ASCHERSON, P. Bemerkungen tiber das Vorkommen gefarbter Wurzeln bei den Pontede-
riaceen, Haemodoraceen und einigen Cyperaceen. Ber. Deutsch. Bot. Ges. 1: 498-
502. 1883. [Blue or pale-lilac roots in several genera of Pontederiaceae.]
Aston, H. I. Aquatic plants of Australia. xv + 368 pp. Carlton, Victoria; London;
Portland, Oregon. 1973. [Pontederiaceae, 263-270, 346; Eichhornia crassipes sue
Pontederia cordata introduced and naturalized.]
Austin, D. F. Exotic plants and their effects in southeastern Florida. Environ. Conserv
5: a 34, 1978. [Eichhornia crassipes often found growing with Pistia Stratiotes
(water lettuce).]
Backer, C. A. Pontederiaceae. Jn: C. G. G. J. VAN STEENIS, ed., Fl. Males. I. 4: 255-
261. 1951. [Eichhornia crassipes introduced and widely naturalized in Malesia;
Heteranthera reniformis cultivated; disagrees in many ways with SCHWARTz’s (1930)
BalLey, L. H., E. Z. BAtLey, & BAILEY Hortorium STAFF. Hortus third. xiv + 1290
pp. New York and London. 1976. [Eichhornia, 418; Heteranthera, 558, Pontederia,
900.
BaiLton, H. Monographie des Liliacées. Hist. Pl. 12: 403-600. 1894. [Série des Pon-
tederia, 459-461; Pontederieae, 576-578.]
BARRETT, S. C. H. Breeding systems in Eichhornia and Pontederia, tristylous genera of
the Pontederiaceae. 189 pp. Unpubl. Ph.D. dissertation, Univ. California, Berkeley.
1977.* (Diss. Abstracts B. 38(8): 3526. 1977 [1978?].)
. Floral biology of Eichhornia azurea (Swartz) Kunth (Pontederiaceae). Aquatic
Bot. 5: 217-228. 1978a.
. Pontederiaceae. Pp. 309-311 in V. H. HEywoop, ed., Flowering plants of the
world. New York. 1978b.
. The evolutionary breakdown of Bre in Lichhornia crassipes (Mart.) Solms
hee hyacinth). Evolution 33: 499-510. 1979.
& D. E. Glover. On the Darwinian eed: of the adaptive significance of
tristyly. Evolution 39: 766-774. 1985.
. Price, & J.S. SHore. Male fertility and anisoplethic population structure
in ‘tristylous Pontederia cordata (Pontederiaceae). Evolution 37: 745- ie ae
BaTE-SMITH, E. C. The phenolic constituents of plants and their taxonomic significan
II. Monocotyledons. Jour. Linn. Soc. Bot. 60: 325-356. 1968. (Eichhornia ae
Pontederia lanceolata. |
BEAL, E. O. A manual of marsh and aquatic vascular plants of North Carolina with
habitat data. N. Carolina Agr. Exper. Sta. Tech. Bull. 247. iv + 298 pp. Raleigh,
North Carolina. 1977. [Pontederiaceae, 147-151; Eichhornia crassipes, Heteranthe-
ra dubia, H. reniformis, Pontederia cordata; illustrations of each.
BENTHAM, G., & J. D. Hooker. Pontederiaceae. Gen. Pl. 3: 836-839. 1883. [Treatment
BHANDARI, N. N. The dope eee as Pp. 53-121 in B. M. Jourt, ed., Embryology
of angiosperms. New York (and several other cities). 1984
BOESEWINKEL, F. D., & F. BoUMAN. The seed: structure. Pp. 567-610 In B. M. Jourt,
ed., Embryology of angiosperms. New York (and several other cities). 1984. [Bro-
meliaceae, Commelinaceae, Philydraceae, Pontederiaceae among 25 families of
monocots with opercula (seed lids).]
BOLKHOVSKIKH, Z., V. GriF, T. MATVEJEVA, & O. ZAKHARYEVA. Chromosome numbers
of flowering plants. nN A. FEepERov, ed. (Russian and English prefaces.) 926 pp.
Leningrad. age [Pontederiaceae, 585.]
BREWBAKER, J. L. The distmbution and phylogenetic significance of binucleate and
trinucleate pollen grains in the angiosperms. Am. Jour. Bot. 54: 1069-1083. 1967.
Casper, S. J., & H. D. KRAuSCH. Pteridophyta ae ye eat 1. Teil: Lycopodiaceae
bis Orchidaceae. Band 23 in H. Ettt, J. GERLorF, & H. HEynic, Siisswasserflora
1987] ROSATTI, PONTEDERIACEAE 43
von Mitteleuropa. 403 pp. Stuttgart and New York. 1980. [Pontederiaceae, 372-
376; Eichhornia crassipes naturalized in Portugal, Heteranthera reniformis natural-
ized in northern Italy, Pontederia cordata occasionally introduced and naturalized
in central ratte aa common southw an
CASTELLANOS, A eae de Brasil. Arq. Jard. Bot. Rio de Janeiro 16: 147-
236. 1958. Teen a eo eet (6 spp.), Pontederia (4 spp.), Reussia
pp.).]
CHARLESWORTH, D. The evolution and breakdown of tristyly. Evolution 33: 486-498.
1979. [Eichhornia crassipes and Pontederia cordata included in discussion.]
CHAUVEAUD, M. G. Recherches sur le mode de formation des tubes criblés dans la
cine des monocotylédones. Ann. Sci. Nat. VII. 4: 307-381. p/. 8. 1896. [Ponte-
deriaceae (Pontederia cordata), 367 and pl. 8, fig. 35 (drawing of transverse section
ot).
CHEADLE, V. I. Vessels in Pontederiaceae, Ruscaceae, Smilacaceae and Trilliaceae. Jn:
N. K. B. Rosson, D. F. CUTLER, . Grecory, eds., New research in plant
anatomy. Jour. Linn. Soc. Bot. 63(Suppl. 1): 45-50. 1970. [Vessel elements of Eich-
hornia crassipes and Pontederia ge illustrated. ]
Cuesters, K. I. M., F. R. GNauck, & N. F. HuGuHes. Angiospermae. Pp. 269-288 in
W. B. HARLAND et a eds., The fossil record. London. 1967. [Pontederiaceae (Het-
eranthera) from the Cretaceous, 522.]
CLAPHAM, A. R., T. G. TuTin, & E. F. Warsurc. Flora of the British Isles. ed. 2. xlviui
1269 pp. Cambridge, England. 1962. [Pontederiaceae, 983; Pontederia cordata eae
in gardens, rarely naturalized.]
CoKER, W. C. The development of the seed in the Pontederiaceae. Bot. Gaz. 44: 293-
301. pl. 23. 1907. [Observations on Eichhornia, Heteranthera, Pontederia: illustra-
tions of H. limosa and P. cordata; antipodals found to persist in Heteranthera and
Ue but see R. W. Smitu, W. R. Scans
Cook, C. D. K., B. J. Gut, E. M. Rix, J. SCHNELLER, & M. Seirz. Water plants of the
world. viii a 561 pp. The Hague. 1974. rpomeden ae 482-492; nine genera, line
drawings. ]
H. B. CorreLt. Aquatic and wetland plants of southwestern United
Q
ie)
fe
2
C
is
ev)
n
[Pontederiaceae 597- 604; line drawings of Eichhornia crassipes, Eurystemon mexi-
canum (not known from the Southeast), Heteranthera dubia, H. Liebmanii, H.
limosa, H. arate Pontederia cordata.
& M. C. JoHNsTON. Manual of the vascular plants of Texas. xv + 1881 pp.
Renner, Texas. 1970. [Pontederiaceae, 366-368; Eichhornia (2 spp., both intro-
duced), Eurystemon (monotypic), Heteranthera (4 spp.); forms ene vs. elongate
stems) of H. limosa thought possibly to represent two species
Cronaquist, A. An integrated system of classification of flowering plants. Frontisp. +
Xvili + 1262 pp. New York. 1981. [Pontederiaceae between Philydraceae and Hae-
modoraceae, one of 15 families in the Liliales; pollen indicated as sometimes trinu-
cleate (other accounts mention only binucleate).]
Czasa, A. T. Structure of starch grains and the classification of vascular plant families.
Taxon 27: 463-470. 1978. [Three groups of monocots based on starch grains and
their carbohydrate substitutes: Pontederiaceae with Bromeliaceae, Commelinaceae,
Philydraceae, Haemodoraceae, and others in “true” monocots (mature seeds with
“highly compound starch grains,” vegetative organs with more than one type of
‘“derived’”’ monocots (mature seeds seldom with starch grains, vegetative organs with
starch and many other carbohydrates); the third group irrelevant here.]
DAHLGREN, R. M. T. A revised system of classification of the angiosperms. Bot. Jour.
Linn. Soc. 80: 91-124. 1980.
T. CuirForD. The monocotyledons: a comparative study. xiv + 378 pp.
44 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
London (and several other cities). 1982. [Liliiflorean attributes of Pontederiaceae
include paracytic stomata, girdle type of endothecial thickenings, starchy (but not
mealy) endosperm, and (possibly) absence of steroid saponins. ]
, 5. ROSENDAL-JENSEN, & B. J. NIELSEN. A revised system of classification of the
angiosperms with comments on correlation between chemical and other characters.
Pp. 149-204 in D.A. baths & D.S. SEIGLER, eds., Phytochemistry and angiosperm
phylogeny. New York.
DAUMANN, E. Das eee bei den Pontederiaceen und die systematische Stel-
reniformis, lacking in H. dubia and H. zosterifolia, placement of family near Liliaceae
(which lack septal nectaries) nevertheless favored; Pontederiaceae considered more
primitive than Liliaceae, although descendant from a common ancestor. ]
Davis, G. L. Systematic embryology of the angiosperms. x + 528 pp. New York, London,
and Sydney. 1966. eee 218, 219.)
ECKENWALDER, J. E., & S.C. H. BARRETT. Phylogenetic systematics of Pontederiaceae.
Syst. Bot. 11: 373-391. 1986.* [South American origin for family, with several east-
ward dispersals; Monochoria and Pontederia monophyletic, aces es Het-
eranthera paraphyletic; heterostyly as a synapomorphy of only one line
EIcHLerR, A. W. Bliithendiagramme. Erster Theil. 348 pp. Leipzig. 1875. onc aces
164-166.]
ERDTMAN, G. Pollen morphology and plant taxonomy. Angiosperms. Frontisp. + xii +
9 pp. Uppsala. 1952. [Pontederiaceae, 335, 336; pollen of Pontederia cordata
illustrated; pollen of family said to have two or three sulculi.]
Eyies, D. E., & J. L. RoBERTSON, JR. ae and key to the aquatic plants of the
southeastern United States. U. S. p. Int. Fish Wildlife Serv. Bur. Sport Fish.
Wildlife Circ. 158. 151 pp. 1963. pee of U. S. Publ. Health Bull. 286. 1944.)
[Pontederiaceae, 106, 107.]
— N.C. A ma nual of aquatic plants. Revised ed., with eee Appendix by
E. C. OGDEN. 1x + 405 pp. Madison and Milwaukee, Wiscon and London. 1957.
[Pontederiaceae (Heteranthera, Pontederia), 171-173; oe including those
of several forms of P. cordata.
Gisss, R. D. Chemotaxonomy of flowering plants. Vols. 1-4. 2372 pp. Montreal and
London. 1974. [Vol. 4 includes bibliography, index, and addendum; numerous
references to Pontederiaceae. }
Goprrey, R. K., & J. W. Wooren. Aquatic and wetland plants of southeastern United
States. Monocot yledons. xii + 712 pp. Athens, Georgia. 1979. [Pontederiaceae, 534-
541; perpen crassipes, Heteranthera dubia, H. reniformis, Pontederia cordata
tr :
HEGNAUER, Chemotaxonomie der Pflanzen. Band 2. Monocotyledoneae. 540 pp.
Basel ue ‘Stuttgart. 1963. [Pontederiaceae, 419-421: Eichhornia crassipes with al-
kaloids, hydrocyanic acids, and possibly triterpenes by one account, lacking alka-
loids, saponins, and tannins by another; neither alkaloids nor saponins detected in
Pontederia cordata.
HE.Liquist, C. B., & G. E. Crow. Aquatic vascular plants of New England: part 5.
Araceae, Lemnaceae, Soares Eriocaulaceae, and Pontederiaceae. New Hamp-
shire Agr. Exper. Sta. Bull. 523. iii + 46 pp. 1982. [Pontederiaceae, 35, 38-46;
1987] ROSATTI, PONTEDERIACEAE 45
ne dubia, H. reniformis, Pontederia cordata; illustrations, distribution
eee Teer Y., & K. R. Sutvanna. The receptive surface of the angiosperm
stigma. Ann. Bot. II. 41: 1233- na 1977. [Eichhornia, Pontederia said to have
unicellular paige on dry stigm
Hooker, J. D. Hydrothrix, a new genus of Pontederiaceae. Ann. Bot. London 1:
89-94, pl. 7. ae (Helpful illustrations of this aberrant genus
Horn, C. N. Anatomical adaptations to the aquatic environment in ‘the Pontederiaceae,
its taxonomic usefulness. (Abstr.) ASB Bull. 31: 62. 1984a. [Eichhornia, Heteran-
thera, Hydrothrix, Pontederia, and Zosterella considered; all seedlings initially pro-
duce linear, nonpetiolate leaves; anatomical variation adaptive, of little taxonomic
value. ]
. A systematic revision of the genus Heteranthera (sensu lato, Pontederiaceae).
xiv + 260 pp. Unpubl. Ph.D. dissertation, Univ. Alabama, University. 1985a. (Diss.
Abstracts B. 46(7): 2174. 1986.)
Huser, H. The treatment of the saree in = vemoneyy, ua of classi-
fication. Jn: K. KUBITZKI f higher
categories. Pl. Syst. Evol. eucel: 1: 285- 298, 1977. [Pontederiiflorae (Pontederiales,
Philydrales) one of five ae of monocotyledons in which dicotyledonous
features are rare or absen
Hunter, C. os Vee of Arkansas. viii + 296 pp. Ozark Society Foundation,
Little R Arkan 1984. [Pontederiaceae, 32, 33; Heteranthera limosa and
pein ire fihocsied in color.
Hurtcuinson, J. The families of flowering plants. ed. 3. xx + 968 pp. Oxford. 1973.
[Pontederiaceae, 761-764.
Jones, S. B., JR. Mississippi flora. I. Monocotyledon families with aquatic or wetland
species. Gulf Res. Rep. 4: 357-379. 1974. [Pontederiaceae, 372-374; Eichhornia
crassipes, Heteranthera dubia, H. Liebmannii, H. limosa, H. reniformis, Pontederia
cordata; need for ane collections of Heeaniae indi cated.
KRISHNAVENI, M., M. VIVEKANANDAN, & S. NAGARAJAN. Pigment studies on Eichhornia
labellum. ee Te Bot. 30: 207-209. 1981. Cnsrensis Gane and beta-caro-
tenes), ee eae Saas cyanidin) in F. crassipes.]
LAKSHMINARAYANA, : DAR Rao, A. J. PANTULU, & G. THYAGARAJAN. Com-
position of lipids 1 in roots, erik ane and flowers of Eichhornia crassipes (Mart.)
Solms. Aquatic Bot. 20: 219-227.
Lona, R. W., & O. LAKELA. A flora a tropical Florida. xvii + 962 pp. Coral Gables,
Florida. 1971. [Pontederiaceae, 274, 275; Eichhornia crassipes, Pontederia lanceo-
lata (= P. cordata var. lancifolia (Muhl. ) Torrey).]
LoveLL, J. H. The flower and the bee. xvii + 286 pp. New York. 1918. [Observations
on Heteranthera reniformis, 200; on Pontederia cordata in southern Maine, 105-
107 (fig. 53)—Bombus vagans with about 70 floral visits per minute, the larger B.
borealis (see Asa Gray Bull. 6: 60-65. 1898) with a lesser rate.]
LowpeNn, R. M. Revision of the genus Pontederia L. Rhodora 75: 426-487. 1973.
MacRoserts, D. T. The vascular plants of Louisiana. Bull. Mus. Life Sci. Louisiana
State Univ. 6. 165 pp. Shreveport, Louisiana. 1984. [“‘Pontedariaceae,” 53; Eich-
hornia crassipes, ee dubia, H. Liebmannii, H. limosa, H. reniformis,
Pontederia corda
Martin, A.C. The ee internal morphology of seeds. Am. Midl. Nat. 36: 513-
660. 1946. [Pontederiaceae, 550, 551; Eichhornia crassipes, Heteranthera dubia,
Pontederia cordata.
Mason, H. L. A flora of the marshes of California. ix + 878 pp. + errata. Berkeley and
Los Angeles. 1969. [Pontederiaceae, 343-347; Eichhornia turalized and
locally abundant, mostly in San Joaquin and Sacramento valleys; Feeesnihera dubia
46 JOURNAL OF THE ARNOLD ARBORETUM [voL. 68
known from few localities, perianth tubes much shorter than those elsewhere in
.8.; Monochoria vaginalis locally established in experimental rice plots, native to
India and southeastern Asia); line drawings.
MCATEE, W. L. Wildfowl food plants. ix + 141 pp. Ames, Iowa. 1939. [Pontederiaceae,
46-48; seeds of Heteranthera dubia and Pontederia cordata eaten by various wild
ucks.
sae ona, H. Reihe Lilliiflorae. 7”: H. MELcHior, Engler’s Syllabus der Pflanzenfami-
lien. ed. 12. 2: 513-543. 1964. [Pontederiaceae, 534, 535, by U. HAMANN.]
Mee W.C. Storage and germination of seeds of aquatic plants. New York State
Agr. Exper. Sta. Bull. 652. 17 pp. 1936. [Seeds of Heteranthera dubia and Pontederia
cordata should be stored in water, at 1-3°C; dry storage prevented germination of
seeds of H. dubia, contrary to an earlier report.]
A ic plants of the United States. x + 374 pp. Ithaca, New York. 1944.
[Pontederiaceae, 199-206; Eichhornia (2 spp., both introduced; E. crassipes thought
possibly to be native to Florida), Heteranthera (4 spp.), Pontederia (1 sp.); illustra-
tons, distribution maps, comparison of leaves of Pe/tandra virginica, Pontederia
cordata, and Sagittaria latifolia.]
MuLter, J. Fossil pollen records of extant angiosperms. Bot. Rev. 47: 1-142. 1981.
Naar 104; report of Pontederia cordata (D. M. JARzEN, Palynology 2:
29-38. 1978) in Maestrichtian (Upper Cretaceous) rejected.
NETOLITzKY, F. Anatomie der Angiospermen-Samen. Handb. Pflanzenanat. II. Arche-
gon. 10. vi + 365 pp. 1926. [Pontederiaceae, 74.]
OGDEN, E. C. Anatomical patterns of some aquatic vascular plants of New York. New
York State Mus. Sci. Serv. Bull. 424. v + 133 pp. 1974. [Transectional illustrations
of Heteranthera dubia (p/. 38; stem, peduncle), Pontederia cordata (pl. 39; stem).]
Ouive, E. W. Contributions to the histology of the Pontederiaceae. Bot. Gaz. 19: 178-
184. pl. 17. 1894. [Long crystals of calcium oxalate in Eichhornia crassipes and
Pontederia cordata, those in Heteranthera limosa evidently much shorter. ]
Ono, T. Embryologische Studien an einigen Pontederiaceen. Sci. Rep. T6hoku Univ
Biol. 3: 405-415. 1928. [Schematic drawing shows micropylar (and not chalazal, as
reported by Davis) megaspore developing into megagametophyte
Ornourr, R. The breeding system of Pontederia cordata L. Bull. Torrey Bot. Club 93:
407-416. 1966. [Floral of other P f breeding
systems in Eichhorn and Pontederia. ]
Perry, F. Water gardening. ed. 3. 2 unnumbered + xvii + 338 pp. 62 bee London.
1961. [Numerous references to Eichhornia, Heteranthera, and Ponteder
Proctor, G. R. Flora of the Cayman Islands. xii + 834 pp. London. 1984. “Pontede-
riaceae (EL ichhornia crassipes), 228, 229; good gies:
RADFORD, A. E., H. E. AHLEs, & C. R. BELL. Manual of the vascular flora of the Carolinas.
Ixi + 1183 pp. Chapel Hill, North Carolina. 1968. [Pontederiaceae, 272, 273; Eich-
hornia crassipes, Pontederia cordata (including P. sles Nutt.), Heteranthera
dubia, H. reniformis oo cidass to northeastern North Car
Rao, T.S., & R. R. Rao. Pollen morphology of ponies Tiel. Pollen Spores 3:
45, 46. 1961. [Illustrations of peli sakes on (also light micrographs) and
Monochoria vaginalis; pollen said to be one- or two-sulculat
RicHarps, J. H., & S.C. H. BARretr. The pee nara basis of tristyly i in Eichhornia
paniculata (Pontederiaceae). Am. Jour. Bot. 71: 1347-1363. 1984. [Action of genes
ae development of floral morph first apparent sali eked in differ-
entiation ongation followed
by apie floral- tube, and style elongation, all of which a be controlled by
hormones produced in anthers.
Rickett, H. W. American wildflowers. 252 pp. New York. 1964. [Pontederiaceae, 58,
59: color photographs of Fichhornia crassipes (pl. 35, p. 63) and Pontederia cordata
(pl. 36, p. 66).]
—. Wildflowers of the United States. Vol. 2. The southeastern states. Part 1. x +
1987] ROSATTI, PONTEDERIACEAE 47
322 pp. New York. 1966. [Pontederiaceae, 89-91; color photographs of Eichhornia
crassipes, Pontederia cordata, and P. . lanceolata ee 29, p. 91); line drawings of
Heteranthera dubia, H. limosa, and H. reniformis, p. 90.]
ROTHERT, W. Die Krystallzellen der Pontederiaceen. Bot Zeit. 58: 75-106. p/. 4. 1900.
[Numerous illustrations of calcium oxalate crystals, including raphides, in Fich-
hornia.|
SCHONLAND, S. The apical meristem in the roots of Pontederiaceae. Ann. Bot. 1: 179-
182. 1887. [Eichhornia azurea, E. crassipes, Pontederia cordata
ScHutz, A. G. Las Pontederiaceas de la Argentina. Darwiniana 6: 45-82. pls. J-5. 1942.
[Eichhornia (4 spp.), Heteranthera (4 spp.), Pontederia (2 spp.), Reussia (1 sp.);
illustrations, photographs
ScHWARTz, O. Anatomische, morphologische und systematische Untersuchungen iiber
die Pontederiaceen. Beih. Bot. Centralbl. 42: 263-320. 1926.
. Zur Systematik und Geographie der Pontederiaceen. Studien zu einer Mono-
graphie der Familie. Bot. Jahrb. 61(Beibl. 139): 28-50. 1927. [Protologues and
justifications for tribes and sections later employed in Die natiirlichen Pflanzenfami-
lien.
—. Pontederiaceae. Nat. Pflanzenfam. ed. 2. 15a: 181-188. 1930.
ScuttHoRre C. D. The biology of aquatic vascular plants. xviii + 610 pp. London.
967. [References to species of Pontederia indicate that plants are usually sterile if
Be in deep water (68), that the roots are the only organs in which xylem has
both vessels and tracheids (169), and that vegetative parts are of great importance
as food for pigs and muskrats (453).
Simpson, M. G. Systematics and aoe ultrastructure of the Pontederiaceae. (Abstr.)
Am. Jour. Bot. 71(5, part 2): 18 84.
. Pollen ultrastructure of the oe ae evidence for exine homology with
the Haemodoraceae. Grana (in press). [Exine sculpturing oe architecture said to
indicate close relationship between the two families (pers. c
SinGH, V. Vascular anatomy of the flower in some species of the ae ae Proc.
Indian Acad. Sci. B. 56: 339-353. 1962. [Raphides and tannin-filled cells scattered
in parenchyma of perianth, stamens, ovary wall, ovules, and central axis in Eich-
hornia crassipes (raphides but not tannins mentioned for Monochoria);, presence of
inverted bundles in perianth and leaf lamina indicates petiolar nature of both in E.
crassipes. |
SMALL, J. K Flora of the Florida Keys. xii + 162 pp. New York. 1913. [Pontederiaceae,
29, ae
of the southeastern flora. xxii + 1554 pp. Chapel Hill, North Carolina.
1933. eae 265-268: Eichhornia crassipes (Piaropus crassipes) thought
to be native in interior peninsular Florida.
SmituH, R. W. Endosperm of Pontederiaceae. Bot. Gaz. 45: 338, 339. pis. J-4. 1908.
[Illustrations of megagametophyte development in Pontederia, cells deteriorate but
nuclei of antipodals persist in Eichhornia and Pontederia; see, however, COKER,
Ww. R. Smirtu.]
SmitH, W. R. A contribution to the life history of the Pontederiaceae. Bot. Gaz. 25:
324-337. pls. 19, 20. 1898. [Descriptions and eee : embryology of Eich-
hornia crassipes, Heteranthera graminis (probably = dubia), and Pontederia
cordata; antipodals in Eichhornia and Pontederia said o ephemeral, implied to
be so in Heteranthera, see COKER, R. W. SMITH.]
So_ms-LauBAcH, H. Pontederiaceae. Monogr. Phanerog. 4: 501-535. 1883a.
—. Uber das Vorkommen cleistogamer Bliiten in der Familie der Pontederaceae.
Bot. Jahrb. 4: 100, 101. 1883b. [Several genera discussed.]
Soueécges, R. Embryogénie des Pontédériacées. Développement de l’embryon chez le
Pontederia cordata L. Compt. Rend. Acad. Sci. Paris 242: 2080-2083. 1956. [Illus-
trations of developmental sequence.]
STANDLEY, P. C. & J. A. STEYERMARK. Pontederiaceae. Jn; Fl. Guatemala. Fieldiana
48 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Bot. 24(3): 42-52. 1952. [Eichhornia (3 spp.), Heteranthera (2 spp.), Pontederia (3
Spp.
STEBBINS, iG: L., & G. S. KHusH. Variation in the organization of the stomatal complex
in the leaf epidermis of monocotyledons and its bearing on their phylogeny. Am.
Jour. Bot. 48: 51-59. 1961. [Illustration of stomatal complex of Pontederia te
fig. 8; Pontederiaceae with two subsidiary cells in all species studied.]
STEYERMARK, J. A. Flora of Missouri. Ixxxiii + 1725 pp. Ames, Iowa. 1962. [Ponte-
deriaceae, 401-404.
TAKHTAJAN, A. L. Outline of the classification of flowering plants (Magnoliophyta). Bot.
Rev. 46: 225-359. 1980.
THIERET, J. W. Aquatic and marsh plants of Louisiana: a checklist. Louisiana Soc. Hort
Res. Jour. 13(1). 2 unnumbered + 45 pp. Univ. S.W. Louisiana, Lafayette. 1972.
[Eichhornia crassipes, Heteranthera dubia, H. limosa, H. reniformis, Pontederia
cordata vars. cordata and lanceolata.
THORNE, R. F. A phylogenetic classification of the Angiospermae. Evol. Biol. 9: 35-
106. 1976.
VALENTINE, D. H., ed. Pontederiaceae. Jn: T. G. TuTIn et al., eds., Fl. Europaea 5: 85,
86. 1980. [Eichhornia (E. crassipes), Heteranthera (H. reniformis), and Monochoria
(probably M. vaginalis) by D. A. WEBB; eee (P. cordata) by D. H. VALENTINE. ]
Voss, E.G. Michigan flora. Part I. Gymnosperms and monocots. xviii + 488 pp. 8 pis.
Bloomfield Hills, Michigan. 1972. eee 378, 379.]
VUILLEUMIER, B. S. The origin and Ae tia! development of heterostyly in the
angiosperms. Evolution 21: 210-226.
Warp, D. B. Checklist of the vascular ies of Florida. Part I. Psilopsida, Lycopsida,
Sphenopsida, Filicinae, Gymnospermae, Monocotyledoneae. Univ. Florida Agr.
Exper. Sta. Tech. Bull. 726. 72 pp. 1968. [Pontederiaceae, 50; Eichhornia crassipes,
Watts, W. A. The full-glacial vegetation of northwestern Georgia. Ecology 51: 17-33.
2 foldout diagrams. 1970. [In reference to Wisconsin glaciation, Pontederia (sp.?)
pollen from pollen zones Q1 (probably full-glacial) and aa (probably late-glacial)
at both Langan Pond and Bob Black Pond, Bartow Co.]
Wir, H.C. D. pe. Aquarium plants. (English translation by J. A. SCcHUURMAN.) Fron-
tisp. + 255 pp. London. 1964. [Heteranthera, 207, 208; H. dubia thought to occur
most often in alkaline water; H. reniformis thought to tolerate brackish water.]
ia R. E., Jr., & R. W. CHERY. Pontederiaceae. /n:
R. _ SCHERY, eds. , Fl. Pan . Ann. Missouri Bot. Gard. 31: 151- 157. 1944.
Seika (2 spp. ), ee ee (3 spp.), Pontederia (2 spp.).]
KEY TO THE GENERA OF PONTEDERIACEAE IN THE
SOUTHEASTERN UNITED STATES
General characters: Rooted or floating herbs, submersed, emersed, or sometimes on
wet ground, stems sympodial, either stout and sometimes connected by stolons or elon
gate; leaves simple, alternate, sessile and ligulate or petiolate, venation parallel, a distinct
reduced petiole and/or blade, each flowering stem with a single leaf sometimes differing
from the others, flowers perfect (some species tristylous); perianth of 6 petaloid tepals in
2 series, the lobes imbricate, fused to various degrees basally, actinomorphic to zygo-
morphic (then with 2 lips of 3 lobes each); stamens usually 6 or 3 (and then sometimes
with 3 staminodes), the filaments adnate to perianth tube, inserted at various levels (often
in the same flower), the anthers usually with introrse, longitudinal dehiscence; carpels 3
united; style single; ovary superior with I or 3 fertile locules; nectaries septal or absent;
1987] ROSATTI, PONTEDERIACEAE 49
ovules solitary (in unilocular ovaries) or numerous in each locule, anatropous, bitegmic;
fruit a I-seeded utricle or a many-seeded capsule; seeds small, those in capsules with
longitudinal ridges.
A. Inflorescences usually several- to many-flowered; perianths zygomorphic; stamens 6
(at least i alin flowers), long axes of anthers and filaments not parallel;
aoe
. Ovaries ae fertile, many late locules; fruit a capsule; t usually ee
ON I a sien Grapes tocid le, odie enusere ds Aaa ee ee ts 1. Eic nia.
B. Ovaries with 1 ee . os locule (and 2 aborted locules); Hee a Sac
plants rooted inssubstrate. 2.15.4 ceeewa yes ds tie eee ie 3. Pontederia.
>
ieee usually |- few- flowered; perianths actinomorphic or jee
morphic; stamens 3 (at least in chasmogamous flowers), long axes of anthers and
filaments parallel; nectaries absent. .................0 00 --ee ee 2. Heteranthera.
Tribe EICHHORNIEAE Schwartz, Bot. Jahrb. 61(Beibl. 139): 32. 1927.
—
Eichhornia Kunth, Fichhornia, Genus Novum [Diss.]. 1842; Enumeratio
Pl. 4: 129. 1843, nom. cons.
Perennial [or annual], submersed, emersed, or floating herbs. Stems stout,
more or less vertical, often connected by stolons. Adult leaves exstipulate,
either sessile, linear, and thin (if submersed), or petiolate, with the petioles
longer than the blades and usually more or less inflated, the blades broadly
elliptic to orbicular. Inflorescence a spike or panicle [or single flowered], pe-
dunculate, the subtending bract of each with a highly reduced petiole and blade,
the single leaf on each flowering stem with a large, sheathing base, little or no
petiole, and a reduced blade. Flowers perfect, some species tristylous; perianth
mostly funnelform, zygomorphic, with 2 lips of 3 lobes each. Stamens 6, un-
equal in length; filaments curved; anthers oblong, auriculate and somewhat
movable on the filaments, much shorter than the filaments, the adaxial 3 either
included deep within the perianth tube or near its summit, lower than the
abaxial 3, which are either near the summit of the perianth tube or exserted.
Ovary with 3 locules, each with numerous ovules on an axile placenta; stigma
(depending on style length) included deep within the perianth tube, or near its
summit, or exserted. Fruit a many-seeded, membranaceous capsule with loc-
ulicidal dehiscence; seeds longitudinally ribbed. (Eichornia A. Rich., 1850,
orthographic variant; Piaropus Raf., 1837, nom. rejic.) Type species: E. azurea
(Sw.) Kunth (Pontederia azurea Sw.), typ. cons. (Named for Johann Albrecht
Friedrich Eichhorn, of Berlin, 1779-1856.)— WATER HYACINTH.
A genus of about seven species native to the American tropics and perhaps
subtropics, including one, Eichhornia natans Solms, that appears to be closely
related to (if not conspecific with) plants that may occur naturally in tropical
Africa and Madagascar, and another, E. crassipes (Mart.) Solms, that through
introductions has spread throughout the tropics and to adjacent warm-tem-
perate areas. Eichhornia can be distinguished from other genera of Pontede-
riaceae by a combination of floral characters including a mostly funnelform
perianth, six stamens, and an ovary with three fertile, many-ovulate locules.
Schwartz (1927, 1930) placed Eichhornia in the monogeneric tribe Eichhor-
nieae Schwartz and proposed two sections in the genus that were neither ad-
50 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
equately defined by him nor widely accepted by later workers (e.g., Alexander,
Castellanos, Schulz), although they may have some utility. Section PRoto-
EICHHORNIA Schwartz (paniculate inflorescences, plants rooted in the ground)
included FE. paniculata (Sprengel) Solms and £.. paradoxa (Mart.) Solms, while
sect. Eueichhornia Schwartz (= EICHHORNIA) (spicate inflorescences, plants free
floating) included EF. azurea (Sw.) Kunth, EF. natans, and E. crassipes. Addi-
tional names, and perhaps species, exist, and the genus is in need of taxonomic
attention on a worldwide basis
Two species of Fichhornia have been reported from the southeastern United
States, each an introduction, apparently from Brazil. Eichhornia paniculata
differs from EF. crassipes, 2n = 32, in the characters by which the two sections
are distinguished and in its complete lack of inflated petioles. It was at least
at one time naturalized in peninsular Florida from plants in cultivation (Alex-
ander; Muenscher, 1944), but I have seen no specimens from the area and the
species 1s not included in recent floristic accounts (e.g., Godfrey & Wooten,
Long & Lakela, and Ward). Eichhornia azurea, 2n = 32, also lacking inflated
petioles, 1s an introduction in southern Texas (Correll & Correll).
Perianths in Eichhornia, including those of our plants, are various intensities
of blue, violet-blue, or lilac, with those of E. crassipes often pale and rarely
even white; those of EF. paniculata are often darker in the lower three lobes.
The upper-middle perianth lobe in E. crassipes usually bears a deep violet-
blue area with a yellow spot inside, while that of EF. paniculata has an unbor-
dered, bilobed yellow spot (Alexander).
Eichhornia crassipes, the water hyacinth, is generally considered to be the
world’s most serious aquatic weed. An enormous amount of research has been
conducted in an effort to understand many aspects of its biology, with the
ultimate but perhaps unattainable goal of eradicating it from areas and habitats
in which it is not native. The literature on this species, which has been reviewed
by Sculthorpe and more recently by Pieterse, is correspondingly immense. The
Hyacinth Control Journal (now the Journal of Aquatic Plant Management),
the existence of which underscores the significance and extent of the problem,
contains only a portion of what has been published.
Problems caused by the water hyacinth, although multifarious, are all more
or less direct results of the tremendous, rapidly accumulated biomass generated
by the plants. Floating mats are frequently large enough to obstruct navigation
completely, to impede drainage to the point of flooding, to contribute in various
ways to eutrophication, and to cause wastage of impounded water by displace-
ment and transpiration. It was conservatively estimated that in Louisiana
damage and losses attributable to the foregoing probably exceeded five million
dollars per year in the 1940’s (Penfound & Earle). The water hyacinth has been
reported to have detrimental effects on rice paddies (Sculthorpe) and to provide
excellent conditions for mosquitoes and other disease-carrying organisms (Viet-
meyer). The floating mats are thought to accelerate greatly and perhaps alter
fresh-water succession (see Sculthorpe) and to prevent the occupancy of lakes,
ponds, and streams by various kinds of waterfowl (Vietmeyer).
Methods of controlling the water hyacinth have been both numerous and
1987] ROSATTI, PONTEDERIACEAE 51
varied. Removal of the plants by hand has been effective in small waterways
and rice fields, but this may be hazardous if disease-carrying organisms are
present and is impractical if the mats have attained even relatively small sizes.
Various devices (including lasers) have been constructed either to cut temporary
paths through the mats or to destroy them completely, but the costs involved
have been high. Numerous chemicals, most commonly 2,4-dichlorophenoxy-
acetic acid (2,4-D), have been employed, but effects on the environment have
usually been detrimental. Drainage of infested areas has been effective in killing
the plants, but this may ultimately prove to be unwise because it favors seed
production, which could enhance the adaptability of the species. Many control
methods result in the accumulation of dead and decaying plant material that
must be removed in order to prevent eutrophication.
Attempts at biological control have included the use of fungi, snails, mites,
insects, fish, and manatees. Significant control by the host-specific weevil Neo-
chetina eichhorniae Warner has been reported in Louisiana (Goyer & Stark)
and in Florida (Center & Durden); N. bruchi Hustache and the pyralid moth
Sameodes albiguttalis (Warren) have also been released in Florida (Center &
Durden). Center & Durden (p. 28) note that “‘recent successes with biological
control of water hyacinth. . .have now been reported worldwide.”
Accounts regarding the first appearance of Eichhornia crassipes in the United
States are somewhat varied (see Penfound & Earle). Despite some evidence
that it was cultivated shortly after the Civil War, it was, according to some,
first shown at an exposition in New Orleans in 1884. The plant attracted a
great deal of attention as a beautiful, easily grown ornamental. Because of its
popularity and vigorous growth, its escape from cultivation and subsequent
naturalization were probably inevitable. In Louisiana, and elsewhere in the
world, its introduction to nature was commonly effected by exasperated gar-
deners who, in attempting to rid cultivated pools and ponds of this initially
desirable but soon troublesome aquatic, threw living material into local water-
ways in the hope that it would be carried away. Unfortunately, the plants
thrived out of cultivation in areas where natural enemies were lacking. The
species was reported from Florida in 1890, was known from each of the coastal
states in the Southeast (its maximum and present range in our area) by the
early 1900s, and was first recorded in California in 1904 (Bock, 1968). Never-
theless, in North America it appears to have been and continues to be a serious
problem only in Louisiana, Mississippi, and Florida (Sculthorpe).
The water hyacinth, a native of the South American tropics, has been intro-
duced and is now naturalized throughout most tropical and subtropical areas
of the world, with an adventive range extending into such warm-temperate
areas as the southeastern United States, California, Japan, southeastern China,
northern Africa, Portugal, Uruguay, and South Africa (for distribution map,
see Barrett, 1977, or Sculthorpe, p. 462). Although it is called the “Florida
devil” in South Africa (Vietmeyer), and although its introduction throughout
the Old World seems to postdate its first occurrences in North America, it is
unclear whether the species spread secondarily from that continent or was
introduced outside of the Western Hemisphere directly by plants obtained from
a2 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
South America. It seems likely that both contributed (see also discussion of
style morph distribution, below). It was introduced into Malaysia in 1894
(Backer).
The plants can rapidly cover stagnant or slow-moving bodies of fresh water
because of their remarkable capacity for vegetative growth and reproduction.
A single plant reportedly developed in one season into a patch of about 600
m?* through the production of a radiating system of stolons and associated
rosettes (Aston; see also Batanouny & El-Fiky). The foliage is killed by frost
or generally cold conditions, but the stems may survive and resume growth
when temperatures rise. Unconfirmed reports indicate that although the plants
are sensitive to salt water, stems protected by sheathing leaf bases may survive
exposure long enough for dispersal along sea coasts (Vietmeyer); stems so
protected may also withstand periods out of water. Seeds remain viable for up
to 15 years and may aid in dispersal of the plants, as well as in their reestab-
lishment following extermination of the parental plants.
Although it has been difficult to assess the relative importance of reproduction
by seeds in the spread of Fichhornia crassipes, there is now little doubt that it
has been very much underestimated in the past. While Hitchcock and colleagues
reported very few seedlings in Louisiana despite extensive seed production,
tremendous numbers of young plants were discovered along the banks of the
White Nile in November, 1963, less than six years after the species was first
seen in the region (Pettet). Ironically, the massive establishment is thought to
have resulted from attempts to eradicate the species with 2,4-D. The seedlings
were most abundant on the decomposed material left by the killed mats of F.
crassipes and were absent from adjacent banks of natural, sandy soil. The free-
floating habit of £. crassipes often limits sexual reproduction, particularly in
the adventive range of the species, by enabling the plants to reach and then
occupy habitats that never become favorable for germination and seedling
establishment. In habitats with seasonally fluctuating water levels, which are
more commonly occupied in the native range of the species, sexual reproduction
may be very important, since seeds germinate and seedlings become established
in warm, shallow water during periods of extensive desiccative damage to
vegetative parts.
Barrett (1980a, 1980b) determined that clones of Kichhornia crassipes from
Louisiana, Florida, California, Mexico, South America, Africa, and India all
retained the potential for sexual reproduction and that observations to the
contrary were due to environmental and not genetic factors. Sexual reproduc-
tion in nature is evidently limited by inadequate pollination and unsuitable
conditions for seed germination and seedling growth and not by the inbreeding
depression, self-incompatibility, and accumulation of deleterious mutations
often characteristic of largely vegetative species.
The free-floating habit and vigorous asexual reproduction of Eichhornia
crassipes have been held responsible in one way or another for the reported
disruption of tristyly in the species. These features have often resulted in pop-
ulations that are either monomorphic (particularly in the adventive range of
the species) or dominated by a single floral form. In either case selection has
presumably favored the development of self-compatibility, high levels of which
1987] ROSATTI, PONTEDERIACEAE 33
have been detected in many populations (Barrett, 1977; Francois; Mulcahy).
Barrett (1979) studied a marshland population in Costa Rica consisting of both
mid- and long-styled forms and determined that within each, seed production
following illegitimate pollinations was only slightly less than that associated
with legitimate pollen deposition, indicating both self-compatibility and weak
and/or residual self-incompatibility. The results of progeny tests involving
seeds obtained from these plants revealed low levels of disassortative (unlike
genotype) crossing for each floral form. While this pattern of crossing is at least
in part due to pollinator behavior (foraging bees tended to visit most of the
flowers of an inflorescence before departing), it also indicates high levels of
self-compatibility.
The habit and growth characteristics mentioned above have further con-
tributed to the disruption of tristyly in Eichhornia crassipes by allowing the
plants to occupy extensive areas, particularly within the adventive range of the
species where pollinators are supposedly ill adapted and/or limiting (Barrett,
1977). Flowers within the native range of E. crassipes are usually visited by
insects large enough to partition the different pollen types effectively and thereby
to cross-pollinate the three floral forms (e.g., Ancyloscelis gigas, a species of
long-tongued bee, is the major pollinator in the lower Amazon). Flowers in
the adventive range, on the other hand, have apparently been attracting smaller
pollinators, so there has been selective pressure to bring the anthers and stigmas
closer together. Such floral modifications would also be favored if pollinating
vectors were numerically limiting because they would increase the chances of
self-pollination. Barrett (1979) reported that four percent of the mid-styled
flowers sampled from a Costa Rican population considered to be outside the
native range of the species were semihomostylous (upper set of anthers adjacent
to stigma; also reported by Francois) and that this condition was accompanied
by increases in pollen deposition. The development of semihomostyly and
related phenomena is probably responsible at least in part for the weakened
pollen trimorphism observed as another aspect of the breakdown of tristyly in
E. crassipes in that size and number of pollen grains are dependent on anther
level (Barrett, 1979).
Semihomostyly in Eichhornia crassipes 1s eenerally thought to have been
derived from tristyly because its is restricted and because it evidently
has not been detected in the native range of the species. Reports of the condition
throughout E. heterosperma E. J. Alex., E. natans, and E. diversifolia (Vahl)
Urban (see Barrett, 1979), as well as in races of FE. azurea (Barrett, 1978a),
prompted Barrett (1979) to conclude that it developed a number of times within
the genus. Its relative infrequency in E. crassipes appears to be the result of
limited sexual reproduction and consequently slow evolutionary rates within
the species (Barrett, 1979).
Investigations into the distribution of style-morphs among New World pop-
ulations of Eichhornia crassipes have suggested that the species is native to the
Amazon basin and perhaps to parts of the Paraguay and Parana river systems,
as well, instead of to the tropics and subtropics of the New World in general,
as has been widely thought (see primarily Barrett & Forno). Trimorphic pop-
ulations, which if of limited occurrence would be expected primarily in areas
54 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
ofancient occupancy (assuming that the species is indeed primitively tristylous),
have been reported only from the Amazon basin in Brazil (where the species
is thought to have originated) and from lagoons near the confluence of the
Paraguay and Parana rivers in Argentina (to which it is thought to have spread
by natural means). Populations in Paraguay, Uruguay, Guyana, Venezuela, and
Colombia, as well as throughout the Caribbean, Central America, and warm-
temperate North America, evidently lack the short-styled morph and are there-
fore considered to have resulted from introductions. (The lack of specialized
pollinators in Central America also implies that the species is introduced there.)
Distribution data on style-morphs may provide insights into the spread of
Eichhornia crassipes when considered in conjunction with the genetic basis of
tristyly in the species. Since the short-styled morph (S__M__ or S__mm) cannot
be segregated from crossings involving the mid- (ssM__) and/or long- styled
(ssmm) morphs, its presence in the adventive range of the species would have
required separate introduction(s). The fact that it is evidently absent from these
areas suggests that the spread of the species throughout the world has involved
only a very few and perhaps even a single introduction, unless the short-styled
morph for some reason either was not selected by man from nature or is ill
equipped to become introduced and established outside its native range. Oth-
erwise, one would expect at least some introductions to have involved the
short-styled morph. The predominance of the mid-styled form in the adventive
range and of the short-styled morph in the native range (Barrett & Forno)
would then be explained by simple genetics. That seed production of the short-
styled morph in the Lower Amazon was found to be 44-75 percent higher than
that of the other two style-morphs (Barrett, 1977, 1980a) 1 1s not only consistent
with the foregoing but may imply that it de-emphasizes vegetative re]
and is therefore less well adapted to establishment onteide its native range.
Considerable effort has been expended to find uses for Eichhornia crassipes
on the assumption that exploitation would constitute the most economically
sound form of control (see primarily Pieterse). Plants have been investigated
as animal fodder (silage, hay, pelletized feed), but their high water content has
made harvesting, storage, and processing difficult. The costs of using the water
hyacinth as fertilizer and mulch have also been prohibitive. The plants have
been utilized with some success as sources of plant hormones and other chem-
icals and have been fermented to produce methane. Because the roots of E.
crassipes are effective in absorbing nitrates, phosphates, and potassium, the
species has been used to purify water that has been polluted by fertilizers.
Fishermen in the Philippines and in Bangladesh maintain circular mats that
provide shade and shelter and therefore attract fish, and farmers in Bangladesh
and Burma transform mats into floating gardens by the application of fertile
bottom muck. Neither the water hyacinth if grown for a crop nor the plants
grown on the floating gardens require manufactured fertilizer, irrigation, or
land. Leaves of water hyacinth are used in Thailand to wrap cigars and are
utilized by the Chinese in wicker and basket work.
REFERENCES:
Under family references see ALEXANDER; ASTON; BACKER; BARRETT (1978a, 1979):
CASTELLANOS; CoRRELL & CORRELL; GODFREY & WOOTEN; LONG & LAKELA: MUENSCHER
1987] ROSATTI, PONTEDERIACEAE 5)
(1944); ScHULZ; SCHWARTZ (1927, 1930); SCULTHORPE; and Warb. See also the Journal
of Aquatic Plant Management (formerly the Hyacinth Control Journal) and Aquatic
Arnott, H.J. A scanning electron microscope study of raphides in £7 hhornia crassipes.
(Abstr.) Bot. Soc. Amer. Misc. Ser. Publ. 158: 8. 1980. 0. [Dstbuton, structure, and
Seen of calcium oxalate crystals and the cyt vacuoles in which they
cur.
eee I, & H. C. GancuLee. Spermatogenesis in Eichhornia crassipes Solms. Jour.
Indian Bot. Soc. 16: 289-296. 1937. [Pollen binucleate.]
Barrett, S. C. H. Tristyly in Eichhornia crassipes (Mart.) Solms (water hyacinth).
Biotropica 9: 230-238. 1977. [Distribution map and helpful photographs of floral
for
eee al reproduction in Eichhornia crassipes (water hyacinth) I. Fertility of clones
from diverse regions. Jour. Appl. Ecol. 17: 101-112. 1980a. II. Seed production in
natural populations. bid. in 124. 1980b.
cological genetics of breakdown in tristyly. Pp. 267-275 in J. Haeck & J. W.
WoIERROR: eds., Structure and functioning of plant populations 2. Phenotypic
and genotypic variation in plant populations. Amsterdam, Oxford, and New York.
1985a. [Reviews his own work with Eichhornia, primarily E. paniculata; includes
previously cited but unpublished data.
ral trimorphism and monomorphism in continental and island populations
of Eichhornia paniculata ees Solms (Pontederiaceae). Biol. Jour. Linn. Soc.
25: 41-60. 1985b.*
ae Style morph distribution in New World populations of Eich-
hornia crassipes (Mart.) Solms-Laubach (water hyacinth). Aquatic Bot. 13: 29 9-306.
1982.
Barton, L. V., & J. E. Horcuxiss. Germination of seeds of Eichhornia crassipes Solms.
Contr. Boy ce Thompson Inst. 16: 215-220. 1951. [Combination of high tempera-
tures and light needed for complete germination of dormant seeds; some seeds able
to survive one month in ice, others two months in temperatures up to 15°C; when
stored for six to 12 months, more seeds remained viable at 20-30° than at 5° or
BaTANouny, K. J., & A. M. EL-Fixy. The water hyacinth (Eichhornia crassipes Solms)
in the Nile system. Aquatic Bot. 1: 243-252. 1975. [A single rosette produced 43
daughter rosettes in 50 days; estimated production for 200 days over 3.4 million.]
na REN biology. 186 pp. Unpubl. Ph.D. dissertation, Univ. California, Berke-
ley. 1966.* (Diss. Abstracts B. 28(1): 61. 1967.) [Includes sae A literature. ]
he water hyacinth in California. Madrofio 19: 281, 282. 8. [Northern limit
in California, and perhaps the world, about ten miles northw a oe Sacramento. ]
_ The distribution of the water hyacinth, Eichhornia crassipes (Mart.) Solms.
(Abstr.) XI Int. Bot. Congr. Abstr. 1969: 17. 1969a
—. Productivity of the water hyacinth E Haun crassipes (Mart.) Solms. aed
50: 460-464. 1969b. [Despite high death rates due nter cold, some plants
survived and resumed growth in the spring, allowing | i species at least to maintain
its distribution in California; rosette ae ie pp
rates attained in more tropical climates.]
Borescu, K. Die Gestalt der Blattstiele ae Eichhornia crassipes (Mart.) Solms in ihrer
Abhidngigkeit von verschiedenen Faktoren. Flora 104: 296-308. pl. 9. 1912. [Inflated
petioles induced by full light, low oe and floating habit; elongated petioles
associated with opposite conditio
Britton, N. L. Piaropus azureus. Addisonia 2: 67, 68. pl. 74. 1917. [Color plate.]
CENTER, T. D., & W. URDEN. Variation in waterhyacinth/weevil interactions re-
sulting from temporal differences in weed control efforts. Jour. Aquatic Pl. Manage.
24: 28-38. 1986. [Biological control by a host-specific weevil, Neochetina eichhorn-
56 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
iae, in Canal-M, Palm Beach County, Florida, in 1983-1984. Neochetina bruchi and
the pyralid moth Sameodes albiguttalis also present and probably effective. Citation
of several other recent successes in biological control throughout the world.
SPENCER. The phenology and growth of water hyacinth (Eichhornia
Hees ee Solms) in a eutrophic north-central Florida lake. Aquatic Bot. 10:
1-32. 1981. [Data on numerous aspects (e.g., le blade area, biomass distribution,
oo. crop) recorded for comparison with thos control insects
(not identified) established and widespread. ]
Curtiss, A. H. The water hyacinth in Florida. Plant World 3: 38-40, 1900. [Recounts
the disappearance from the St. Johns River system of millions of plants, covering
tens of thousands of acres, around 1896.]
Francois, J. Observations sur l’hétérostylie chez Eichhornia . (Mart.) Solms.
Acad. Roy. Sci. Outre-Mer Bruxelles Bull. Séances 3: 501-519. 1964.* [Interprets
semihomostyly to be evolutionary precursor to tristyly; see, however, BARRETT,
1979.
Goyer, R. A., & J. D. STARK. The impact of Neochetina eichhorniae on waterhyacinth
in southern Louisiana. Jour. Aquatic Pl. Manage. 22: 57-61. 1984. [Significant
control by N. eichhorniae within 14 months at a site in Assumption Parish. ]
Harper, R. M. The water-hyacinth in Georgia. Plant World 6: 164, 165. 1903. [Lux-
uriant growth discovered in pool near Valdosta; chances of escape considered small. ]
. Habenaria repens and Piaropus crassipes in Leon County, Ace ee ie
267-270. 1916. [Piaropus = Eichhornia; introduction thought t
Hivcaceck, A. E., P. W. ZIMMERMAN, H. KirkPATRICK, JR., & T. T. EARLE. Growth
and reproduction of water hyacinth and alligator weed and their control by means
of 2,4-D. Contr. Boyce Thompson Inst. 16: 91-130. 1950.
Jounson, A. M. The mid-styled form of Piaropus paniculatus. Bull. Torrey Bot. Club
51: 25-28. 1924. [= Eichhornia
Littte, E. C. S. The control of water weeds. Weed Res. 8: 79-105. 1968. [Includes E.
crassipes. |
. Handbook of ee 2 aquatic plants. A review of world literature. FAO
Fish. Tech. Paper 187. vii 76 pp. Rome. 1979.*
Mutcany, D. L. The et biology of Eichhornia crassipes (Pontederiaceae).
Bull. Torrey Bot. Club 102: 18-21. 1975. [Although many findings were later sub-
stantiated by BARRETT, some (e.g., that short-styled form does not exist: that floral
form is determined by two nee at a single locus) were not.
maries in Spanish and French. ) vii + 174 pp. Washington, D. C. 1976. [Numerous
references to Eichhornia crassipes, a weed in 52 nations.
PENFOUND, W. T., & T. T. Earve. The biology of the water hyacinth. Ecol. Monogr.
18: 447-472. 1948. [Distribution, nature and extent of damage, morphological/
anatomical details, phenology, role in succession, autecology (including effects of
seenaney vegetative and sexual reproduction (including pollination and seed
mination), and control methods; stages of germination and seedling development
hee d.]
Pettet, A. Seedlings of Eichhornia crassipes: a possible complication to control mea-
sures in ~ Sudan. Nature 201: 516, 517. 1964.
PIETERSE, A. The water hyacinth (Eichhornia crassipes),; a review. fae Trop.
Agric. res ae 42. 1978. [666 citations covering biology, control, and u
RicHarps, J. H. Developmental potential of axillary buds of water hyacinth, Fichho rnia
crassipes Solms (Pontederiaceae). Am. Jour. Bot. 69: 615-622. 1982. [Plants grown
in distilled water produced more inflorescences than those in nutrient solutions;
axillary buds of the former developed into “renewal shoots” (which continue the
main axis), those of the latter formed stolons (which produce new plants). |
1987] ROSATTI, PONTEDERIACEAE oi}
[ae
Heteroblastic development in the water hyacinth E
Bot. Gaz. 144: 247-259. 1983. [Scanning electron and light fe Saale seedling-
to-adult leaf transition illustrated.]
TAG EL SEED, M., & M. Operp. Sexual reproduction of Eichhornia crassipes (Matt.)
Solms in the Nile. Weed Res. 15: 7-12. 1975. [Mid-styled form predominant, short-
and long-styled forms absent or very rare; low seed set thought to be due to high
temperatures and low humidity, but see BARRETT, 1980b.]
VIETMEYER, N. D. The beautiful blue devil. Natural History 84: 64-73. 1975. [Despite
several inaccuracies, an interesting account with several good photographs. ]
Tribe HETERANTHEREAE Schwartz, Bot. Jahrb. 61(Beibl. 139): 35. 1927.
2. Heteranthera Ruiz & Pavon, Fl. Peruv. Chil. Prodr. 9. 1794, nom. cons.
Perennial or annual, submersed, emersed, or floating herbs. Stems stout and
more or less vertical to elongate and more or less horizontal. Adult leaves
stipulate or exstipulate, either sessile, linear (strap shaped) and thin, or petiolate,
with the petioles longer than the blades and not inflated, the blades reniform,
cordate, or lanceolate. Inflorescence a spike or single flowered, sessile or pe-
dunculate, the subtending bract lacking a petiole and blade, the single leaf on
each flowering-stem identical to all other leaves. Flowers perfect; perianth
salverform, with 6 lobes, actinomorphic or subactinomorphic (1 lobe different
in shape and/or spaces between lobes unequal). Stamens 3 (sometimes | in
cleistogamous flowers), equal in length or the lateral 2 shorter, inserted on
adjacent adaxial tepals; filaments straight or curved; anthers oblong or ovate,
sometimes auriculate and somewhat movable on the filaments, sometimes
nearly equal in length to the filaments, exserted, in subgen. Zosterella becoming
circinately coiled after anthesis. Ovary with 1 locule, the ovules numerous in
2 or more rows on each of 3 more or less completely intrusive placentae; stigma
usually exserted. Fruit a many-seeded membranaceous capsule with loculicidal
dehiscence; seeds longitudinally ribbed. (Schollera Schreber, 1791, not Roth,
1788; Heterandra Beauv., 1799; Leptanthus Michx., 1803, nom. superfl. [in-
cludes type of Heterandra Beauv.]; Zosterella Small, 1913, Eurystemon E. J.
Alex., 1937.) Type species: H. reniformis Ruiz & Pavon, Fl. Peruv. Chil. 1:
43. 1798. (Name from Greek heteros, different, and antheros, anther, in reference
to the unequal anthers of most species, including the type.)— MUD-PLANTAIN,
WATER STAR-GRASS, BUFFALO-GRASS
A small genus of about 12 species native to tropical and temperate regions
of the New World and Africa. Heteranthera is distinguished from other Pon-
paar by a suite of floral characters, including salverform perianths with
ix equal or nearly equal lobes, one- or imperfectly three-loculate ovaries with
numerous ovules, and three stamens.
Infrageneric classifications of Heteranthera have been varied. Persoon (1805)
evidently was the first to subdivide the genus (as Leptanthus Michx.), estab-
lishing two subgenera (see Brizicky) based on androecial morphology: Heter-
anthera (including the types of both Heteranthera Ruiz & Pavon and Heter-
andra Beauv.), with dimorphic stamens (“‘Filam. longitudine inaequalia, antherae
biformes”); and Leptanthus, with stamens of only one form (“Antherae uni-
58 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
formes lineares, filamenta aequalia’’). Solms-Laubach later (1883a) established
two sections: Leptanthus Solms (including the type of Heteranthera), with
petiolate leaves; and Schollera Solms, with strap-shaped (ligulate) leaves.
Schwartz (1927) recognized three sections on entirely different grounds. Section
Protoheteranthera (= sect. HETERANTHERA) was characterized by three- to many-
flowered inflorescences with no cleistogamous flowers; sect. Heterantheropsis,
by one- or two-flowered inflorescences with no cleistogamous flowers; and sect.
Euheteranthera, by many-flowered inflorescences with one or more cleistog-
amous flowers.
Small (in Small & Carter) segregated the monotypic genus Zosterella from
Heteranthera on the basis of monomorphic (vs. dimorphic) stamens and linear
(vs. broad) leaf blades. Although such a treatment generally agrees with that
of Persoon, neither has been widely accepted. Recent studies by Horn (1985a),
however, suggest that division of Heteranthera along these lines may be most
tenable. Neither leaf morphology, on which the classification of Solms-Laubach
was based, nor the presence or absence of cleistogamous flowers, by which
Schwartz’s sections were partially delimited, has proven to be taxonomically
significant. Horn (1984a) reported that all species in the group initially produce
strap-shaped (linear) leaves and that the mature leaf form is habitat dependent.
Horn (1985a) also determined that all species produce cleistogamous flowers,
usually in response to development under water, and that such structures are
for the most part morphologically identical to chasmogamous flowers (see also
Thieret). Underwater development commonly results in reduced numbers of
flowers per inflorescence as well, unless the species is one that normally pro-
duces only one- or two-flowered inflorescences.
Horn (1985a) recently completed a revision of Heteranthera sensu lato that
employed a number of biosystematic methods (e.g., flavonoid chemistry, cy-
tology, pollen and seed morphology, vegetative anatomy, developmental bi-
ology), as well as numerical (cluster and principal component) and cladistic
analyses of populations and species, respectively. Although a fairly convincing
case for the existence of two groups was presented, I do not agree that the data
support their recognition at the generic level (viz., Heteranthera and Zosterella).
Horn’s decision to do so may have resulted from a failure to incorporate out-
group comparisons in the analyses: the characters by which Heteranthera and
Zosterella were reported to differ (e.g., internode length on flowering stem,
length of time flowers stay open, perianth pubescence, androecial morphology
[monomorphic or dimorphic stamens], filament inflation, anther shape and
coiling, seed size) seem much less significant than those by which other genera
in the Pontederiaceae differ (e.g., fusion of perianth parts, number of stamens,
attachment and dehiscence of anthers, number of locules per ovary and of
ovules per locule, and fruit type). In addition, the stamens of H. limosa (Sw.)
Willd. and H. peduncularis Bentham are only slightly dimorphic (indeed, Per-
soon included Leptanthus ovalis Michx. [= H. limosa] in subg. Leptanthus,
Horn, however, has correctly placed it with the other species having dimorphic
stamens), further lessening the distinction between the two groups. I am, there-
fore, recognizing as subgenera the two groups treated as genera by Small and
1987] ROSATTI, PONTEDERIACEAE 59
Horn. The species concepts of Horn appear to be sound, and much of the
following is based on his revision. (Unless otherwise indicated, material at-
tributed to Horn is taken from his dissertation.)
Subgenus HETERANTHERA (annuals with dimorphic stamens and noncoiling
anthers) comprises the eleven species placed by Horn in Heteranthera sensu
stricto; all but one (H. callifolia Reichenb. ex Kunth, of sub-Saharan Africa)
are native to the New World. Two groups were identified in subg. HETERANTHE-
RA by Horn’s cladistic analysis, although they were not given names (see,
however, Horn, 1986b). One group of species, all 2 = 14, is represented in the
Southeast by H. /imosa and probably by H. rotundifolia (Kunth) Griseb. The
other group, in which x = 8, has among its members H. multiflora (Griseb.)
Horn, 2n = 32, and H. reniformis Ruiz & Pavon, 2n = 48, both found in our
area. All species of the subgenus in our area produce petiolate leaves
Heteranthera limosa and H. rotundifolia, each with single-flowered inflores-
cences, are identical in flavonoid chemistry, chromosome number, and pollen
and seed morphology. Plants of H. /imosa commonly form rosettes and have
ovate to elliptic leaf blades, actinomorphic perianths, and nearly monomorphic
stamens, while those of H. rotundifolia do not form rosettes and have at least
some round leaf blades, subactinomorphic perianths (one lobe cordate at the
base), and clearly dimorphic stamens (the lateral filaments curved). Plants of
H. limosa usually occur in shallow water, commonly at the edges of ponds and
in roadside ditches, and are submersed as seedlings. Rosette-forming individ-
uals and others with elongate, horizontal stems occur in the Southeast and may
represent two biologically meaningful taxa, according to Correll & Correll,
although Horn considered the latter condition to be induced by growth in water
10 cm or more deep. The distribution of H. /imosa extends from California
and the central United States (including Tennessee, Mississippi, Arkansas, and
Louisiana) to central South America. Plants of H. rotundifolia grow in small
odies of water or on mudflats. With the exception that the species has not
been reported either from California or from our area, it has a distribution
almost identical to that of H. limosa. Although Steyermark did not report H.
rotundifolia from Missouri (he apparently did not consider it to be distinct
from H. limosa), Horn indicated that it occurs throughout the state and along
the Missouri side of the Arkansas border.
Although panels Schreber i isa later pono and is herons illegitimate at the generic level,
i name for owever,
Zosterella Small is also ace ae since it is the more > familiar name for these plants, it is
appropriate to make the following new combination at the level of subgenus
Heteranthera subg. Zosterella (J. K. Small) Rosatti, comb. et stat. nov.
Zosterella J. K. Small in J. K. Small & J. J. Carter, Fl. Lancaster County [Pennsylvania], 68. 1913.
Type species: Z. dubia (Jacq.) J. K. Small aes dubia Jacq.).
Leptanthus Michaux (1803) is a GEMS name, since Michaux cited Heterandra Palisot de
Beauvois (Trans. Am. Philos. Soc. 4: 17 99) (as ‘‘Heteranthera’’), for which the type species is
Heterandra reniformis Beauv., 1799, not 5 Hexranter reniformis Ruiz & Pavon, 1798, although
both names apply eresantealy to the same speci
60 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Heteranthera multiflora and H. reniformis both have spicate inflorescences,
and they are difficult to distinguish vegetatively. However, in H. multiflora the
flowers are purple, and the inflorescence is more than twice as long as the
subtending bract, while in H. reniformis the perianths are white and the spike
is usually about as long as the bract. In the United States H. reniformis occurs
from Connecticut and southern New York southward to southeastern Virginia,
western North Carolina, northern Georgia, and western Florida, and westward
to Louisiana, southern and western Missouri, and southern Illinois, with dis-
junct localities along the Rio Grande in Texas. It is also found in Mexico and
Central America, Cuba, Hispaniola, and Jamaica. In South America it is known
from Venezuela and Colombia, southward to northern Argentina, Paraguay,
and southern and eastern Brazil. Heteranthera multiflora has a similar but
more disrupted distribution. In the United States it is known from New Jersey
south to northeastern North Carolina; from southwestern Illinois, Missouri,
southeastern Nebraska, Kansas, Oklahoma, Arkansas, and southwesternmost
Tennessee; and from outlying stations in Mississippi and southernmost Texas.
Disjunct localities have been found in northern Venezuela, northern Argentina,
Paraguay, and southern and eastern Brazil. Although plants of both species can
either float or become rooted in shallow water or moist ground, those of H.
multiflora reportedly are able to occupy deeper water than those of H.. reniformis
because of their superior ability to produce elongate stems. Heteranthera pe-
duncularis, primarily of high elevations in Mexico but also reported from
southeastern Arizona and Guatemala, is very similar to H. multiflora and H.
reniformis but can evidently be distinguished from them by its glabrous or
glabrate (vs. pubescent) lateral staminal filaments.
Subgenus ZOSTERELLA (perennials with monomorphic stamens and coiling
anthers) comprises one, or perhaps two, species, both present in the Southeast.
The leaves are linear in both and resemble those of Potamogeton species, except
that they lack a distinct midrib. Heteranthera Liebmannii (Buch. ex Magnus)
Shinners (Zosterella longituba E. J. Alex.) has been recognized by some (e.g.,
Alexander; Correll & Correll) as being distinct from H. dubia (Jacq.) MacM.
(Zosterella dubia (Jacq.) Small) because of differences in flower size and seed
morphology. The perianth tubes of H. Liebmannii are usually much longer
than those of H. dubia (5-12 vs. 1.5-7 cm), and the seeds of the former are
nearly globose, black-brown, and 14- to 16-ribbed, while those of the latter are
ellipsoid, yellow-brown, and 10- to 12-ribbed. Horn has reported, however,
that from north to south there is a general increase in perianth-tube length and
that, although there is a genetic component, shorter perianth tubes were pro-
duced on cooler mornings among experimental plants. He also found that seed
color was related to development and that the number of ribs per seed varied
within populations.
Heteranthera dubia occurs at various depths and tolerates a relatively wide
range of temperatures (Steyermark) in still to swift, usually alkaline water
(Hellquist & Crow; Muenscher, 1944; De Wit). The species is known from
southern Quebec to North Dakota, south to Texas and Florida, and from more
scattered localities in Washington, Oregon, California, Arizona, Mexico, Cen-
1987] ROSATTI, PONTEDERIACEAE 61
tral America, and the Caribbean region. Heteranthera Liebmannii is found on
mud or in relatively still water from Alabama to Mexico and the Caribbean
(i.e., it has a more southern distribution than H. dubia) and is reportedly more
abundant than H. dubia in Texas (Correll & Correll).
Horn (1983) determined that mature seeds of Heteranthera dubia sink upon
being released in autumn and germinate the following spring. Plants flower in
the first year and may overwinter in foto beneath the ice, although growth does
not occur below 8°C. In shallow and/or swift water the plants may produce
much shorter stems and internodes, forming denser, more circular patches
(Steyermark) that may provide food and shelter for fish (Correll & Correll).
Plants growing on mud develop short, stiff leaves and stems and have been
recognized under various names (see Horn, 1983). Although such variants have
been considered to be environmentally induced and therefore unworthy of
formal taxonomic recognition (Horn, 1983; Steyermark), it is interesting and
possibly significant that they are more likely to flower than those in more
typical, aquatic conditions (Fassett). While emersed plants flower to some
extent, most submersed ones are sterile or develop only flowers that are hidden
in the leaf axils and do not open (Voss, under family references). Thieret
reported that such flowers are structurally identical to chasmogamous ones and
showed that they were induced when buds did not reach the surface or when
they were pulled under water by the current.
Flower color, which is variable in Heteranthera, has been described in detail
by Horn (1985a). Among species of subg. HETERANTHERA in our area, the basic
perianth color is purple, lavender, pale blue, or white (yellow, or rarely blue
or white in the extraregional H. Seubertiana Solms; blue or white in the ex-
traregional H. zosterifolia Mart.), while the upper middle lobe is variously
marked with dark purple, brown, green, and/or yellow. The central and lateral
stamens, as well as the filament and anther of any one stamen, usually differ
in color; either filaments or anthers are purple, blue, yellow, or white. Styles
and (evidently) stigmas are either purple or white (the style is yellow and the
stigma blue in H. zosterifolia). With the exception of purple stigmatic hairs,
all externally visible flower parts of H. dubia (subg. ZOSTERELLA) are yellow or
pale yellow.
There is some evidence that the stamen dimorphism (both in color and size)
found in species of subg. HETERANTHERA is related to pollination biology.
According to studies of H. reniformis by Lovell, pollen from the pale blue or
greenish anther of the long central stamen is deposited on a visiting bee while
it gathers pollen (the flowers lack nectaries) from the more conspicuous yellow
anthers of the shorter lateral stamens. Such observations, including that of a
green color for the central anther, evidently have not been corroborated by
other workers.
The only economic significance of Heteranthera involves the occurrence of
some of its members as weeds in rice fields: H. reniformis and H. limosa in
the United States (Barrett, 1978b) and H. reniformis in northern Italy (Webb,
in Valentine). The seeds of various species, including H. dubia, are eaten by
wildfowl (McAtee; see also Fassett). Both H. dubia and H. reniformis are
62 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
considered to be rare and endangered in various northeastern states (Hellquist
& Crow).
REFERENCES:
Under family references see ALEXANDER; BARRETT (1978b); me & CORRELL;
FASsETT; HELLQUIST & Crow; Horn (1984a, 1985a); HUNTER; LOVELL; MCATEE; MUEN-
SCHER (1944); ScHWaRTz (1927); SoLMs-LAUBACH (1883a); eee ee VALENTINE;
Voss; and DE Wit
AcosTINI, G. El género Heteranthera (Pontederiaceae) en Venezuela. Acta Bot. Venez.
9: 295-301. 1974. [Five species, including H. /imosa and H. reniformis.]
Brizicky, G. K. Subgeneric and sectional names: their starting points and early sources.
Taxon 18: 643-660. 1969. [Generic subdivisions in PERSOON’s Synopsis Plantarum
are subgenera. ]
East, E. M. The distribution of self-fertility in the flowering plants. Proc. Am. Philos.
Soc. 82: 449-518. 1940. [Dimorphism and heterostyly in Heteranthera.|
Horn, C. N. Life history of Heteranthera dubia (Jacq.) MacM. (Pontederiaceae) with
respect to seasonal and environmental ae on fponpholoe,. 104 pp. Unpubl.
Master’s thesis, Ohio State Univ., Columbus. 1980.
The annual growth cycle of Heteranthera pee in Ohio. Michigan Bot. 23: 29-
34, 1983. [Based on Master’s thesis; mudflat form shown by reciprocal transplants
to be induced wholly by environment. ]
Variation in the adaptations to the aquati t during seedling growth
in the genus Heteranthera (Pontederiaceae). (Abstr.) Am. Jour. Bot. 71(5, part 2):
172. 1984b. [Vegetative differences between Heteranthera and Zosterella appear to
be due to adaptations to different habitats.
. Zosterella vs. Heteranthera, a little used genus in the Pontederiaceae. (Abstr.)
ASB Bull. 31: 62. 1984c. [Recognition as separate genera favored by flower and seed
characters; vegetative aspects of little taxonomic value.
Morphology and distribution of Heteranthera (sensu lato; Pontederiaceae) in
the southeastern United States. (Abstr.) ASB Bull. 32: 46. 1985b. [H. dubia, H.
limosa, H. reniformis, H. rotundifolia.]
Adaptation to the aquatic environment by species of Heteranthera. (Abstr.)
Ibid. 33: 76. 1986a. [Experiments suggested that in response to growth in deeper
water, petioles and more elongate keane stems. }
—_. T ypifications and ¢ a new combination in Heteranthera (Pontederiaceae). Phy-
tologia 59: 290. 1986b. [H. eae eae Horn; sects. Schollera Solms in
C. and Leptanthus Solms in ed.]
Manus J. E. A study of the plone as: a seeds of Heteranthera limosa. 91
p. Unpubl. Ph.D. dissertation, Louisiana State Univ. and Agricultural and Me-
chanical College, Baton Rouge. 1969.*
Persoon, C. H. Leptanthus. Syn. Pl. 1: 56. 1805. [Leptanthus Michx.; the earlier Het-
eranthera Ruiz & Pavon included as a subgenus.
SMALL, J. K., & J. J. CARTER. Flora of Lancaster County [Pennsylvania]. New Yor
336 pp. 1913. [Pontederiaceae, 68, 69; Zosterella Small, Heteranthera, Pontederia.]
THIERET, J. W. Observations on some aquatic plants in northwestern Minnesota. Mich-
igan Bot. 10: 117-124. 1971. [Induction of “‘pseudocleistogamous” flowers in H.
dubia, 117, 118.
VARALDA, G., G. FORNERIS, & F. MONTACCHINI. New findings and interesting confir-
mations of species in the flora of Basso, Vercellese and Oltrepo, Alessandrino, central
east Piedmont, Italy. Allionia 26: 123- 130. 1983.* [H. limosa, H. reniformis.]
Voss, E. G. A vegetative key to the genera of submersed and floating aquatic vascular
.]
plants of Michigan. Michigan Bot. 6: 35-50. 1967. [Includes H. dubia
1987] ROSATTI, PONTEDERIACEAE 63
Tribe PONTEDERIEAE [Schwartz, Bot. Jahrb. 61(Beibl. 139): 39. 1927]
3. Pontederia Linnaeus, Sp. Pl. 1: 288. 1753; Gen. Pl. ed. 5. 140. 1754.
Perennial, emersed herbs. Stems stout and more or less horizontal. Adult
leaves exstipulate; petiolate, petioles usually much longer than blades, not
inflated; blades sagittate, cordate, ovate, or elliptic. Inflorescence a spikelike
panicle,* pedunculate, the subtending bract sometimes mucronate, the single
leaf on each flowering-stem with a large, sheathing base and a petiole much
shorter than the blade. Flowers perfect, all species tristylous except the homo-
stylous P. parviflora; perianth mostly funnelform, zygomorphic, with 6 lobes
in 2 lips of equal [or unequal] lobe number. Stamens 6, unequal in length;
filaments straight or curved; anthers oblong, auriculate, much shorter than the
filaments and somewhat movable on them, the adaxial 3 either included deep
within the perianth tube or near its summit, lower than the abaxial 3, which
are either near the summit of the perianth tube or exserted. Ovary with 2
abortive locules and | fertile one with a solitary ovule pendulous from a
terminal placenta; stigma (depending on style length) included deep within the
perianth tube, or near its summit, or exserted. Fruit a 1-seeded utricle enclosed
in the accrescent, roughened, ridged, and terminally coiled base of the perianth
tube, tipped by the coiled base of the style; seeds not ribbed. (Including Reussia
Endl., 1836, nom. cons., and Unisema Raf., 1808, ““Umsema.”’) TYPE SPECIES:
P. cordata L.; see Britton & Brown, Illus. Fl. No. U. S. & Canada, ed. 2. 1
462. 1913, and discussion below. (Named for Giulio Pontedera, 1688-1757,
professor of botany in Padua, Italy; see Critica Botanica, p. 94. 1737 [p. 77 in
English transl. by A. Hort, 1938].)—PICKEREL-WEED, BLACK-POTATO, WAMPEE,
WILD-GENTIAN.
A small New World genus of five species (Lowden), Pontederia is charac-
terized by a two-lipped perianth, a one-locular ovary (through the abortion of
two locules) containing a single pendulous ovule, and six stamens. The genus
is, for the most part, tropical to subtropical in its distribution. The plants grow
primarily in fresh inland water and in brackish rivers and marshes near the
sea.
Lowden’s revision of Pontederia incorporated evidence from chemistry (phe-
nolics), cytology, and morphology and also included considerations of nomen-
clatural history, dispersal mechanisms, breeding systems, and evolutionary
development. He reviewed the controversy surrounding interpretation of the
Linnaean genus Pontederia and concluded that of the three species listed in
the first edition of Species Plantarum, only P. cordata L. belonged and must
therefore be considered the type. Pontederia ovata L., with one stamen, was
clearly out of place in a genus that was included in the Linnaean class Hexandria,
and the species has since been removed to the Marantaceae. The third species,
P. hastata L. (actinomorphic perianths of mostly free parts, six stamens in
‘The flowers are sessile and are e uped in ile clust long the main axis of the inflorescence;
flowers along the axis, as well as wit luster, are in various stages of development, suggesting
that the soteeee represent meduced branches or ea systems. At least partial resupination of most
flowers d by their uniform orientation at anthesis. (Also see Leggett, 1875
64 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
SH
FicurE |. Pontederia. a-k, P. cordata: a, leaf blade and portion of petiole behind
flowering stem (terminal part) with leaf and bract subtending inflorescence, 4: b, flower
of long- styled form, with style and 3 mid-length stamens exserted, x 3: c, flower of short-
styled form, in semidiagrammatic longitudinal section (e.g., hairs not shown), showing
2 of 3 adaxial, mid-length stamens and 2 of 3 abaxial, long stamens, x 3; d, flower of
adaxial, short stamens and 2 of 5 abaxial, mid-length stamens, x 3; f, glandular hairs
of staminal filaments, x 50; vary, in longitudinal section, showing position of 1
aborted peties . lef and fertile locule with its single pendulous, anatropous ovule,
> h, ransverse section (at level of dashed line in “g’’), showing 2 aborted,
adaxial penn Per fertile locule with its single ovule, x 16; i, terminal part of flowering
1987] ROSATTI, PONTEDERIACEAE 65
groups of five and one based on length, and many seeds per fruit), has been
placed by most botanists in the Old World genus Monochoria Presl.°
Lowden decided to treat Reussia Endl. as a subgenus of Pontederia because
the supposed morphological differences between the two groups are weak and/
or evidently unclear. According to him, the perianth in Pontederia clearly has
two lips of three lobes each, while that in Reussia has an upper lip of four lobes
and a lower of two by some accounts, but an upper of five and a lower of one
by others. In addition, the genera are similar in other aspects of morphology,
including an ovary with one fertile locule and a single pendulous ovule. Lowden
reported haploid chromosome counts of n = 8 in subg. PONTEDERIA (ridges of
the persistent, accrescent perianth base encasing the fruit smooth or toothed;
owering shoot erect) for P. cordata var. cordata, P. parviflora E. J. Alex., and
P. sagittata Pres], and n = 16 in subg. Reussia (Endl.) Lowden (ridges of
perianth base spinulose; Raceane shoots prostrate) for P. rotundifolia L. f.
Chemical data provided by Lowden are consistent with the inclusion of
Reussia as a subgenus in Pontederia on the basis of coefficients of similarity®
he calculated for all pairs of the included taxa of Pontederia (P. cordata vars.
cordata, lancifolia (Muhl.) Torrey, and ovalis (Mart. in Roemer & Schultes)
Solms in DC., P. parviflora, and P. sagittata, of subg. PONTEDERIA; P. rotun-
difolia, of subg. Reussta), as well as on those I calculated for all pairings
involving Heteranthera limosa, Eichhornia crassipes, and the foregoing taxa
of Pontederia. Mean values for coefficients of similarity (zero indicating no
resemblance, one indicating identity) were lower between genera of Pontede-
riaceae (Pontederia-Eichhornia, 0.58, Pontederia-Heteranthera, 0.49, Eich-
hornia-Heteranthera, 0.50) than between subgenera of Pontederia (subg. Pon-
tederia-subg. Reussia, 0.69), although greater between subgenera than between
included taxa of subg. Pontederia (0.62
Lowden speculated that Pontederia originated in the American tropics from
ies aquatic ancestors with many-flowered spikes and flowers with zy-
gomorphic perianths of basally connate parts, six stamens, and a single pen-
*Rafinesque (Med. Repos. N. Y. II. 5: 532. 1808) placed Pontederi data, with a single seed per
ntained, however, that the term had been used to describe the accrescent base of the
perianth tube surrounding the fruit and did not therefore indicate a many-seeded fruit. Fernald
(Rhodora 27: Lie 1. 1925) pointed out that in dedicating the genus to Pontedera, Linnaeus primarily
had plants fro rth America in mind, and that in the fifth edition of Genera Plantarum he added
to Pontederia a 2 opi eaiceican plant with one-seeded seas thus strengthening the idea that his
concept of Pont ape mae sae ware per fru
li ae
*Between any two t
divided by the sum i ce number and the hunber of phenolics neal in ely one or the other.
stem during fruit maturation, which occurs under water, x ¥; j, accrescent, terminally
coiled base of perianth tube enclosing utricle, x 3; k, l-seeded utricle with persistent,
coiled base of style, x
66 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
dulous ovule. He proposed that sometime during the Tertiary the genus spread
from Central to North America, where it initially occurred farther west than
it does at present. Fossils attributable to Pontederia cordata have been re-
covered from the Green River Formation in Wyoming, now considered to be
early or mid-Eocene (Bradley, Knowlton).
Pontederia is represented in the Southeast by three commonly accepted but
taxonomically questionable varieties of P. cordata (see below). The species is
distinguished from others in subg. PONTEDERIA by a combination of characters,
including tristyly and teeth on the ridges of the persistent, accrescent perianth
bases. The plants are largely restricted to stream banks and pond edges where
bare ground (required for seed germination) is exposed by fluctuating water
levels; few individuals are found in “thigh marsh plant communities” (see
Whigham & Simpson).
Variety cordata (leaves sagittate, cordate, reniform, or hastate; mature floral
tube glabrous or sparsely glandular) occurs throughout the eastern United States
and adjacent Canada but is most abundant in the Great Lakes region, in the
Northeast, and on the Gulf and Atlantic coastal plains. It is also found in
southern Brazil and adjacent areas, as well as in Belize, where Lowden reported
specimens that suggested hybridization with P. sagittata. The distribution of
var. lancifolia (P. lancifolia Muhl., 1813; P. angustifolia Pursh, 1814; P. lan-
ceolata Nutt., 1818) (leaves narrowly to broadly lanceolate, mature floral tube
glandular) matches that of var. cordata, with the exception that it appears to
be rare in the Great Lakes region and otherwise less common than var. cordata
in North America outside of southern Georgia and Florida; Lowden reported
it from two localities in Cuba as well. Perry observed that var. /ancifolia is less
hardy than var. cordata, which perhaps explains its more southern distribution.
Godfrey & Wooten reported that vars. cordata and /ancifolia are not easily
distinguished in Florida and southern Georgia, where each occurs in abundance,
and my own observations suggest that the same is true elsewhere in the South-
east. Both varieties are popular among gardeners and have become naturalized
in parts of the Old World (Aston; Casper & Krausch; Clapham et al.; Valentine).
According to Lowden, var. ovalis is restricted to South America and differs
from broad-leaved specimens of var. /ancifolia in its densely pubescent upper
peduncles. Nevertheless, Mather M-277 (Gu), from Marion County, Florida,
was determined by Lowden to belong to var. ovalis. My observations indicate
that the upper peduncles of many specimens of var. /ancifolia from our area
are as densely pubescent as those of this specimen.
The two subgenera of Pontederia differ in the relative importance of vege-
tative and sexual reproduction (Lowden). In subg. Reussia, members of which
have few-flowered inflorescences and long, trailing stems, reproduction through
fragmentation of adventitiously rooted stems has a greater immediate value
than reproduction by seeds, especially in populations composed of a single
floral form (in which all pollinations would be illegitimate and thwarted by
physiological incompatibility systems). In contrast, sexual reproduction may
be of greater importance in subg. PONTEDERIA because inflorescences are many
flowered and the stems are more erect, above ground, and short
All species of Pontederia are tristylous, with the exception of P. parviflora
1987] ROSATTI, PONTEDERIACEAE 67
(subg. PONTEDERIA), in which homostyly (semihomostyly according to Barrett,
1979) is thought to have been derived from the tristylous condition (Lowden).
Ornduff studied the breeding system of P. cordata in a number of populations
along the Atlantic Coastal Plain in the Southeast. Except for one population
in North Carolina in which only short- and mid-styled flowers occurred, all
three floral morphs were represented in each population. Populations varied,
however, in the relative proportions of each morph, presumably because of a
combination of founder effects and vegetative reproduction (see, however, Price
& Barrett, 1982).
Although data regarding the genetic basis for tristyly in the diploid Pontederia
cordata are not yet completely available, Barrett, Price, & Shore assumed it
was the same as that observed in the diploid Eichhornia paniculata, in which
two alleles are present at each of two loci, one of which is epistatic to the other.
Essentially the same is true of E. crassipes, except that this species is a tetraploid
(see also Barrett, 1985a [under Eichhornia], Charlesworth; see, however, Barrett
& Anderson).
In Pontederia cordata, as in the majority of other tristylous plants investi-
gated, legitimate pollinations are most effective in producing seed. Illegitimate
pollinations are less productive both because they are less frequent and because
of the existence of a physiological incompatibility system. Ornduff provided
data from artificial pollinations indicating that in P. cordata the incompatibility
is due to “carpellary factors” (i.e., is of the sporophytic type) and is strongest
in the short-styled form, slightly weaker in the long-styled, and clearly weakest
in the mid-styled (see also Barrett ef a/.). Barrett & Anderson summarized data
from P. cordata vars. cordata and lancifolia, P. rotundifolia, and P. sagitatta
showing that in each the level of self-compatibility, as determined by percentage
of seed set in flowers pollinated with the most compatible pollen (i.e., that from
short, medium, or long stamens), is clearly and consistently greatest in the
mid-styled form, with the exception that seed set in the long-styled form of P.
rotundifolia is approximately equal to that of its mid-styled form. These data
also suggest that the relationships between self-compatibility levels in the short-
and long-styled forms are rather inconsistent among the four taxa. Barrett &
Anderson proposed several hypotheses to explain their observations.
Price & Barrett (1982) investigated tristyly in 74 North American populations
of Pontederia cordata, including 45 from the Southeast, and for the most part
substantiated the findings of Ornduff. They also determined, however, that the
mid-level (medium) stamens of short-styled flowers produced about twice as
many pollen grains as those of long-styled ones. Although the basis of this
difference could not be established, Barrett, Price, & Shore later suggested that
it could result from differences in the time of anther development, since the
mid-level stamens of the short-styled morph are the lower set, while those of
the long-styled morph are the upper set (i.e., the lower set of anthers develops
first, so these are therefore larger and more productive of pollen). Price &
Barrett (1982) also suggested that pollen from short-styled flowers fertilizes
more ovules of the mid-styled morph than does pollen from the long-styled
morph (although Barrett, Price, & Shore later reported that field studies pro-
vided only limited evidence that this was so) and that this difference may
68 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
influence the composition of natural populations in favor of the short-styled
form.
On the basis of data gathered from the same 74 populations mentioned
above, Barrett, Price, & Shore reported that of 69 trimorphic populations, 76.8
percent were significantly anisoplethic (although morph frequencies varied
among populations, presumably because most had not yet reached equilibri-
um). Most frequently encountered were a predominance of the short-styled
morph and a deficiency of the long-styled one, regardless of variety (var. cordata
or var. /ancifolia), habitat type (permanent or temporary), locality (Ontario
and Wisconsin or the Carolinas, Georgia, Florida, and Louisiana), population
size (more or fewer than 500 inflorescences), location within a population
(divided into 10 x 2 m sections), or time (populations observed for five years).
Because of the large sample of populations employed, they considered it un-
likely that historical factors (e.g., dispersal, disturbance, establishment) alone
could be responsible for the anisoplethy observed in Pontederia cordata; they
proposed instead the existence of some selective advantage for the short-styled
morph and a corresponding disadvantage for the long-styled one. It is inter-
esting that Price & Barrett (1984) reported that legitimate pollinations were
most frequent in the long-styled morph, followed in order by the mid- and
short-styled morphs, possibly due at least in part to differences in amount of
surface area available for pollen deposition (e.g., the pollinator’s proboscis tip,
which normally delivers pollen to short styles, is smaller than its head, which
delivers to mid-length styles). Nevertheless, Price & Barrett (1982) reported
no Statistically significant differences among floral morphs in flowering phe-
nology, fruit weight, germination percentage, number of inflorescences per
individual, or flowers (all or chasmogamous only) and seeds per inflorescence.
It is notable that while the situation in P. sagittata appears to be almost identical
to that in P. cordata, it is considerably different in Eichhornia (see discussion
of that genus).
Perianths in species of Pontederia (including P. cordata as represented in the
Southeast) are purple, blue-purple, blue, pale blue, or white, and the anthers
are blue. The extraregional P. subovata (Seub. in Mart.) Lowden differs from
this pattern in sometimes having blue-green perianths, while P. parviflora (the
only homostylous species of Pontederia, see above) has greenish white to white
perianths and black to brown anthers (Lowden). The upper-middle perianth
lobe in species of Pontederia bears a single bilobed yellow spot (Lowden) or
two separate yellow spots (Lovell).
The flowers of Pontederia cordata attract a number of insect visitors, pri-
marily bees of the genera Bombus, Melissodes, and Xylocopa (Price & Barrett,
1982, 1984). The emergence of Dufourea novaeangliae (Robertson), a small
solitary bee, coincides remarkably well with the onset of flowering in P. cordata,
and the insect is not known to visit any of the many other species concurrently
available (Lovell, Percival; see, however, Hurd). According to Hazen, the nu-
merous insects that visit P. cordata do so primarily for nectar, which is produced
by three septal nectaries, but some hymenopterans also collect pollen. Price &
Barrett (1982) determined that the frequency of visits to P. cordata by bum-
1987] ROSATTI, PONTEDERIACEAE 69
blebees (Bombus spp.) in a Canadian (Ontario) population was independent of
floral form.
Evidence provided by Price & Barrett (1984) suggests that the frequency of
legitimate pollinations in populations of Pontederia cordata may be dependent
on the type of pollinators involved and may therefore vary geographically. In
northern North America, species of Bombus, which have broad preferences
and are therefore probably not highly co-adapted to the breeding system of P.
cordata, are the most important pollinators. In the South, on the other hand,
a diverse set of more specific (long-tongued) pollinators is involved, perhaps
most importantly species of Melissodes. These observations may help to explain
why significant levels of legitimate pollination (i.e., levels significantly greater
than those predicted by a model that assumes random pollination) appear to
become less frequent with increasing latitude in the species as a whole. In
Florida, populations of all three morphs experienced significant levels of le-
gitimate pollination, in the C arolinas only some did, and in Ontario none did.
The fruits and associated perianth bases of Pontederia are buoyant because
of the presence of aerenchyma in the latter and normally float for more than
15 days, according to Schulz. Transport by water is considered to be the primary
means of dispersal. Dissemination by ducks and other animals is less important
and probably involves only relatively short distances (see Sculthorpe). Ponte-
deria cordata has been recorded as a food source for the southern black or
mottled duck (Anas fulvigula), and the seeds have been found in the stomachs
of wood ducks (Aix sponsa) (Ridley). Lowden observed that the spinulose
perianth bases encasing the fruits of P. rotundifolia become attached to livestock
in El Salvador and Costa Rica, but in subg. PONTEDERIA (including our plants)
such surfaces are smooth or only toothed, and the fruits are probably less
effectively dispersed in this way.
Pontederia cordata is widely grown as an aquatic ornamental, and it some-
times escapes cultivation. It is reportedly naturalized in Britain (Clapham et
al.) and southern Europe (Valentine). In South America and perhaps elsewhere
it frequently occurs as a weed in rice fields (Barrett, 1978b).
REFERENCES:
Under family references see ARBER (1920), ASTON; BARRETT (1978b, 1979); BARRETT,
Price, & SHORE; CASPER & KRAUSCH; CHARLESWORTH; CLAPHAM, TUTIN, & WARBURG;
GoprreyY & Wooten; HUNTER; LOVELL; ORNDUFF; PERRY; SCHULZ; SCULTHORPE; and
VALENTINE. Under Eichhornia see BARRETT (1985a).
AnperSON, J. M., & S. C. H. Barrett. Pollen tube growth in tristylous Pontederia
cordata L. (Pontederiaceae). Canad. Jour. Bot. 64: 2602-2607. 1986. [Pollen readily
germinated on stigmas in all pollen/stigma combinations; in most, growth of legit-
compatibility.]
BARRETT, S.C. H. The breeding system of Pontederia rotundifolia L., a tristylous species.
New Phytol. 78: 209-220. 1977. [Systems of Brazilian and Costa Rican populations
showed strong resemblances to that of P. cordata.
70 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
. M. Anperson. Variation in expression of trimorphic incompatibility in
Pontederia cordata L. (Pontederiaceae). Theor. Appl. Genet. 70: 355-362. 1985.
[Developmental model proposed to explain high levels of self-compatibility in mid-
styled morph; expression of incompatibility associated with style length, may be
determined by pleiotropic effects of major genes for tristyly; variable expression of
incompatibility within floral morphs suggests polygenic system.
BRADLEY, W. H. Paleolimnology. Pp. 621-652 in D. G. Frey, ed., Limnology in North
America. Madison, Wisconsin. 1963. [Fossils from the Green River Formation in
Wyoming (early or middle Eocene) attributed to Pontederia: see also KNOWLTON. |
Fassett, N. C. Three aquatics from southern Maine. Rhodora 39: 273, 274. 1937. [P.
cordata f. taenia Fassett, with leaf blades lacking or barely apparent. ]
Grover, D. E., & S. C. H. Barrett. Pollen loads in tristylous Pontederia cordata
populations from the southern U.S.A. Am. Jour. Bot. 73: 1601-1612. 1986. [All
three floral morphs in a Louisiana population exhibited significant levels of legiti-
mate pollination; overall, such levels were most frequently observed in the long-
styled morph. ]
HAuMAN-Merck, L. Sur un cas de géotropisme hydrocarpique chez Pontederia rotun-
difolia L. Rec. Inst. Bot. Léo Errera 9: 28-32. 1913.* [Erect inflorescences bend
downward 180° to ripen fruits underwater; see ARBER (1920), 239, 240, 375.]
Hazen, T. E. The trimorphism and insect visitors of Pontederia. Mem. Torrey Bot.
Hetsey, R. M., & A. W. H. Damman. Biomass and production of Pontederia cordata
and Potamogeton epihydrus in three Connecticut rivers. Am. Jour. Bot. 69: 855—
864. 1982. [In P. cordata maximum values attained 100-150 days after beginning
of spring growth, which depends on large biomass of overwintered rhizomes and
roots. ]
Hurp, P. D., Jr. Superfamily Apoidea. Pp. 1741-2209 in K. V. KRoMBEIN et al, Catalog
of Hymenoptera in America north of Mexico. Vol. 2. Washington, D. C. 1979.
[Dufourea novaeangliae said to visit flowers of Pontederia (presumably P. cordata)
and Fagopyrum, 1936; visitors to flowers of P. cordata, 1853, 1944, 2143, 2165.]
Keppy, P. A. Shoreline vegetation in Axe Lake, Ontario: effects of exposure on zonation
patterns. Ecology 64: 331-344. 1983. [Range of water depths tolerated by P. cordata
greatest on the most shaded shores, the species absent from the most exposed shores. |
KNOWLTON, F. H. Revision of the flora of the Green River Formation, with descriptions
of new species. U. S. Geol. Surv. Prof. Pap. 131: 133-182. pls. 36-40. 1923. [Leaf
fossils most closely resembling P. cordata described as Pontederites, see also BRADLEY. ]
LainG, H. E. Respiration of the rhizomes of Nuphar advenum and other water plants.
Am. Jour. Bot. 27: 574-581. 1940. [Rhizomes of P. cordata able to respire anaerobi-
cally for seven days without noticeable injury.
Leccett, W. H. Pontederia cordata, L. Bull. Torrey Bot. Club 6: 62, 63. 1875. [Tristyly;
spikes compound, “spikelets” mostly three-flowered.] Ibid. 170, 171. 1877. [Tristy-
ly.]
Lowpen, R. M. Revision of the genus Pontederia L. Rhodora 75: 426-487. 1973.
[Phenolic compounds isolated but not identified.]
Otis, C. H. The transpiration of emersed water plants: its measurement and its rela-
tionships. Bot. Gaz. 58: 457-494. 1914. [In P. cordata stomata occur on the petioles
and both sides of the leaf blades (although more numerous abaxially); transpiration
rates high. ]
PercivAL, M.S. Floral biology. xv + 243 pp. Oxford, England (and several other cities).
1965. [P. cordata, 155, 160.]
1987] ROSATTI, PONTEDERIACEAE 71
Price, 8. D., & S. C. H. Barrett. Tristyly in Pontederia cordata (Pontederiaceae).
Canad. Jour. Bot. 60: 897-905. 1982.
_ The function and adaptive significance of tristyly in Pontederia cordata
L. (Pontederiaceae). Biol. Jour. Linn. Soc. 21: 315-329. 1984.
Rip.ey, H. N. The dispersal of plants throughout the world. Frontisp. (pl. 16) + xx +
744 pp. 22 pls. 1930. [Pontederia, 194, 231, 491, 493. Mistakenly refers to accrescent
perianth base as pericarp, 194.]
WuicHaM, D. F., & R. L. Simpson. Germination and dormancy studies of Pontederia
cordata L. Bull. Torrey Bot. Club 109: 524-528. 1982. [Rootstocks did not require
cold treatment and could begin to grow whenever temperatures were above freezing;
seeds required eight weeks of moist, cold stratification before germination; germi-
nation rates were highest when temperatures reached 30°C for part of the day,
although minimum temperatures were as low as 5°C.]
Note added in proof. Since this treatment was completed, a paper of consid-
erable significance has appeared in the literature. Various cladistic analyses
reported by Eckenwalder & Barrett (under family references; annotation based
on a manuscript copy of the abstract) suggested that the Pontederiaceae and
Philydraceae are sister groups and that the former is divisible into two groups
of two genera each. Pontederia (including Reussia as a subgenus) and Eich-
hornia were depicted as one clade, while Heteranthera (including Eurystemon,
Hydrothrix, Scholleropsis, and Zosterella) and Monochoria comprised the oth-
er. The family, the two clades, three of the four genera, and both subgenera
were considered to be monophyletic, while Eichhornia was said to be paraphy-
letic. The cladograms generally indicated that tristyly is not the primitive
breeding system in the Pontederiaceae and that it did not arise more than once
in the family. Although it was shown to be a synapomorphy of the Pontederia-
Eichhornia clade, its evolutionary relationship to homostyly in £ ichhornia was
not resolved. The possibility that the dimorphic stamens of the Heteranthera-
Monochoria clade were not derived from a tristylous condition was also sug-
gested. A base chromosome number of n = 8 for the family was favored, from
which n = 7 and n= 15 would have been repeatedly derived. The cytological
diversity in the family was thought to have resulted from both aneuploidy and
polyploidy.—T. J. R.
KAUL, LITHOCARPUS 73
REPRODUCTIVE STRUCTURE OF LITHOCARPUS
SENSU LATO (FAGACEAE):
CYMULES AND FRUITS
ROBERT B. KAUL!
Seventy-three species were examined for structural and developmental de-
tails of the cymules and fruits. The cymules bear one to seven or more flowers
and are subtended by one to nine or more bracteoles. Generally, the number
of flowers and bracteoles in the pistillate cymules is the same or less than in
are torn or disintegrate, or fall away as the cupule matures, leaving the cupule
essentially naked. In species with scaly cupules at maturity, the scales enlarge
and sometimes also become adpressed, thickened, or elongated. Some cupules
the same cymule that have been elevated by the maturing cupule of the fertile
flower, but in some cases they could be developed from latent primordia that
are axillary to the cupular scales.
Nearly all that is known of the reproductive structure of Lithocarpus Blume
comes from studies done for taxonomic purposes. Most notable are the con-
tributions of Camus (1948, 1952-1954) and Soepadmo (1968, 1970, 1972),
which contain numerous illustrations and some discussion of reproductive
structure and its possible phylogeny. Hjelmqvist (1948) detailed floral and
cymular structure in several species and provided some phylogenetic assess-
ments. Nevertheless, only a few species have been investigated for reproductive
detail, and no comprehensive overview of the genus is available. Here I report
on morphological, developmental, and evolutionary aspects of partial inflo-
rescences (cymules) and fruits in 73 species that represent eight of the 14
subgenera proposed by Camus (1952-1954). I give particular attention to the
organization of the cymules and cupules. Details of floral structure will be
presented elsewhere.
Lithocarpus, with perhaps 300 species when taken in its broadest sense, is
second in the Fagaceae only to Quercus L. in number of species. The genus
ranges from northeastern India across central China to Korea and southern
Japan, south to southeastern Asia, the Philippines, and the East Indies as far
east as New Guinea. There is one American species, L. densiflora, which occurs
in the coastal mountains from Douglas County, Oregon, south to Ventura
'School of Biological Sciences, University of Nebraska, Lincoln, Nebraska 68588-0118.
© President and Fellows of Harvard College, 1
Journal of the Arnold Arboretum 68: 73-104. ee 1987.
74 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
County, California, and at scattered locations in the Sierra Nevada of California
(Little, 1971). The range of the genus 1s almost exactly congruent with that of
the third largest genus of Fagaceae, Castanopsis (D. Don) Spach, but it is much
less than that of Quercus.
Lithocarpus is found on a variety of soil types from sea level to about 4000
m, but it 1s most abundant at middle elevations, where it is sometimes one of
Some taxonomists (e.g., Barnett, 1940, 1944; Camus, 1952-1954; Soepadmo,
1972) have recognized Lithocarpus in the broad sense, but others (e.g., Li,
1963; Lin & Liu, 1965; Liao, 1969) have preferred to restrict that name to
some species and to place others in segregate genera. Those favoring the broad
interpretation justify their position by noting that there are intermediate species
between the groups. Further taxonomic complications arise from the fact that
certain species are intermediate in many respects between Lithocarpus s.l. and
both Castanopsis and Querc
Camus (1952-1954) ecu med 14 subgenera in Lithocarpus, most of them
with fewer than 15 species. Because her subgeneric classification covers the
entire genus in its broadest sense, it is the basis of reference for the work
presented here. Soepadmo (1970, 1972), in his treatment of Lithocarpus for
the Flora Malesiana, described some new species and reduced or did not accept
some of Camus’s species; of the 136 species recognized by Camus for Malesia,
he accepted only 64 as good species but did not assign them to subgenera. His
nomenclature is used for the southeastern Asian species discussed here.
The classification of Lithocarpus is based mostly upon cupule and fruit char-
acters (Barnett, 1940, 1942, 1944; Camus, 1952-1954; Li, 1963; Lin & Liu,
1965; Liao, 1969; Soepadmo, 1970, 1972), as it is in other Fagaceae. Gross
inflorescence and flower characters are useful in separating genera (Soepadmo,
1970; Kaul & Abbe, 1984) but not in distinguishing species
Camus (1952-1954) believed Lithocarpus to be one of fhe most primitive
members of the family. She cited seven reproductive characters as primitive
(but was not clear about the reasons for those assessments): the abortive ovules
apical in the nut (known elsewhere only in Quercus subg. Cyclobalanopsis and
one section of subg. Quercus); the scar of the nut large in some species; the
cupule asymmetric in some species; the tomentum that lines the cupule dense
(known elsewhere only in Quercus subg. Cyclobalanopsis and some sections of
subg. Quercus), the cupule poorly developed at anthesis, as is also the case in
Quercus; the cupule fused for much or all of its length to the nut in some
species; and the partitions of the nut absent or poorly developed in some species.
Schottky (1912) and Hjelmqvist (1948) believed Lithocarpus to be the most
pri eae genus of the family, and they suggested that it gave rise—or is a sister
group—to Quercus s.s. and Cyclobalanopsis (Quercus s.l. subg. Quercus and
an Cyclobalanopsis, respectively). Forman (1966; see also Elias, 1971), how-
ever, postulated separate origins of Quercus and Lithocarpus from hypothetical
ancestors and thus implied morphological parallelisms of the two; Trigono-
balanus Forman was seen as having some intermediate characteristics.
Camus (1952-1954, p. 1188) also noted the ‘‘affinités indéniables” of Litho-
carpus subg. Cyclobalanus with Quercus subg. Cyclobalanopsis. Both have more
than three styles per flower in many instances, annular cupules, apical abortive
1987] KAUL, LITHOCARPUS i)
ovules, rudimentary perianthopodia in some instances, and entire, evergreen
leaves. The stigmas, styles, and stamens of each subgenus are typical of their
genera, however, and their characteristics are not shared by the two subgenera.
It is mostly because of these distinct floral characteristics that Barnett (1940),
Camus (1952-1954), and Soepadmo (1968, 1970, 1972) maintained Lithocar-
pus distinct from Quercus despite the similarities in fruits and cupules. I have
shown all these and other differences between the two genera elsewhere (Kaul,
19352722)
The cupules of Lit/ pus and Quercus are often indistinguishable, but those
of Lithocarpus have a greater variety of shapes and ornamentation. Further,
although there is a rather sharp distinction between the lamellate cupules of
Quercus subg. Cyclobalanopsis and the scaly ones of See Rees in some
species of Lithocarpus there are intermediate cupular patt
Further of L Sie arise when some
of the species that strongly suggest Castanopsis sect. Pseudopasania are ex-
amined. These were placed in Lithocarpus subg. Pseudocastanopsis by Camus
(1952-1954) and resemble Castanopsis because of cupular and foliar similar-
ities (i.e., the scales in three groups, the castanopsoid hairs on the abaxial leaf
surface, and the cupules of L. fissa opening by three valves). Soepadmo (1970)
noted several differences between the two genera: Castanopsis has the inner
bark surface smooth, the wood rays only uniseriate, and the cupules solitary
(but enclosing one to three nuts). The cupule has a definite number of growing
points separated by vertical rows of scales, and its vascular system shows a
dichasial pattern. Lithocarpus has the inner bark surface longitudinally ridged,
the wood rays both uni- and multiseriate, the cupules solitary or clustered and
each enclosing a single nut, and the cupular vascular system not dichasial. The
cupule has a continuous, circular growing edge, and there are no sutures.
Barnett (1940) believed that Lithocarpus and Castanopsis are very close and
that their separation is perhaps more artificial than natural. Nevertheless, she
believed their fruit structure distinct enough to treat the two as genera. She
noted that in species of Lithocarpus with spiny cupules (e.g., L. garrettiana, L.
lappacea, L. longispina, L. recurvata), the spines are certainly recurved scales.
The spines and tubercles of Castanopsis, however, do not appear to be the
original cupular scales but develop later, often in the axils of the original scales.
She included in Lithocarpus those species with oblique cupular lamellae, wheth-
er tuberculate or not, in which the fruit is oblique (e.g., L. blumeana, L. en-
cleisacarpa). She placed C. acuminatissima in Castanopsis, however, because
it has oblique cupules with irregular whorls of short spines or tubercles and
because it has some castanopsoid anatomical characters.
In Lithocarpus each pistillate flower has its own cupule (as is the case in
Quercus), but sometimes the cupules are grouped and even fused. In extreme
cases of fusion, the combined cupules appear almost as a single cupule enclosing
several nuts. Soepadmo (1970) showed that in organization of the vascular
system of the cupule, Lithocarpus is the same as Quercus but markedly different
from Castanopsis. Where adjacent cupules are fused, the unified wall that
separates the flowers retains the separate vasculature of each cupule.
Forman (1966) interpreted the one-flowered cupule of Lithocarpus as being
derived from a three-flowered cymule whose valves fused to form one cupule
76 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
around each flower; the one-flowered cupule of Quercus became so by loss of
some valves and the lateral flowers. Thus the one-flowered cupules of both
genera were seen as convergently evolved. This interpretation was illustrated
by Ehas (1971).
Camus (1952-1954) and Soepadmo (1970) noted the variety of patterns of
cupular fusion to the nut. In some subgenera the mature cupule entirely encloses
the nut and is totally fused to it for its entire length (subg. Oerstedia, some
sections of subg. Lithocarpus); in others the cupule entirely covers the nut but
is only partially fused to it (subgenera Lithocarpus (sect. Costatae), Pachybal-
anus, Synaedrys) or is not fused except for the basal scar (subg. Pseudosynae-
drys, and some species of subgenera Pasania and Pseudocastanopsis). In the
unique subgenus Cory/opasania the cupule not only encloses the nut but also
is much prolonged beyond it into a narrow tube; the cupule is only basally
fused to the nut. In many taxa (subgenera Cyrtobalanus and Gymnobalanus,
as well as many species of subgenera Cyclobalanus and Pasania) the cupule
covers just part of the nut and is not fused to it but the basal scar is large.
Camus believed that the greater degree of fusion is the more primitive condition
in the genus.
In some species of Lithocarpus the cotyledons are free, but in others they
are fused. The latter condition is found in some species of Quercus, too, and
Nixon (1985) considered it to be the derived condition in that genus. The
endocarp is tomentose in many species, as it is in some members of Quercus.
MATERIALS AND METHODS
I have examined more than 1000 specimens that my colleagues and I col-
lected in Asiatic and southwestern Pacific island forests. We took special care
to collect developmental as well as mature material. Most of the specimens
were identified by E. Soepadmo, the most recent monographer of southeastern
Asiatic Lithocarpus (Soepadmo, 1970, 1972) and by other taxonomists residing
in the areas of provenance of the specimens.
Most of the specimens were stored in FAA, quinoline-sulfate solution, or
glycerin-alcohol. All are documented by dried voucher specimens in my col-
lection, for which various sets of duplicates are deposited in A, BH, G, K, L, MIN,
SING, and US.
OBSERVATIONS
GROSS STRUCTURE OF THE INFLORESCENCES
The overall structure of the inflorescences of Lithocarpus has been dealt with
in some detail (Kaul & Abbe, 1984; Kaul, 1986). The genus was shown to have
the most elaborate gross inflorescence structure among Lithocarpus, Casta-
nopsis, Castanea, and Quercus. It was suggested that this elaborate structure
is the least specialized condition—one that gave rise to more advanced inflo-
rescences by loss of branching and separation of staminate from pistillate
flowers first within the spike and ultimately, in Quercus, into separate spikes.
1987] KAUL, LITHOCARPUS 7
Spikes bearing usually sessile cymules are variously aggregated into repro-
ductive branches that are caducous or persistent. In a few species some spikes,
especially the staminate ones but occasionally the pistillate as well, are branched
at a cymule (Kaul, 1986). The spikes are variously entirely staminate, entirely
pistillate, androgynous, or androgynecandrous, and more than one pattern often
occurs on a given tree. Furthermore, some cymules contain various combi-
nations of staminate, pistillate, or perfect flowers (see Kaul & Abbe, 1984, fig.
4). Those cymules at the transition point on a spike between staminate and
pistillate cymules more often have both flower sexes or perfect flowers than
do more proximal or distal cymules. Within a spike, the pistillate flowers are
more likely to occur proximally than distally, but the spikes bearing pistillate
flowers are more abundant distally in the total spike-bearing shoot system. In
a few instances the staminate and pistillate cymules are mixed for short dis-
tances along the spike. These phenomena are illustrated in the papers cited
above, while details of cymule and fruit structure are emphasized here. There
is much infraspecific variability in reproductive structure both locally and
throughout the ranges of the species, and variant morphological patterns are
likely to be found in specimens of the species illustrated here that are collected
from other parts of their ranges.
CYMULES IN LITHOCARPUS
The groups of flowers spaced along a spike are often called cymules, dichasia,
or partial inflorescences. ‘““Cymule” is used here generally for the presumably
condensed pleiochasia and dichasia that characterize Lithocarpus and other
Fagaceae.
In the specimens examined for this study, the number of flowers in a cymule
ranged from one to seven (or more in a few instances), but one, three, and five
were the usual numbers (TABLE). (Downward departures from the typical num-
bers are common in a few cymules at the extreme proximal and distal ends of
a spike in most species; such exceptions are not included in the data presented
here.) Often the staminate and pistillate cymules on a specimen contain the
same number of flowers (this was true for 29 of the 73 species shown in the
TABLE), and where the number of flowers is variable and rather high in the
staminate cymules it is also that way in the pistillate cymules (e.g., Lithocarpus
elegans and L. harmandii, TABLE). However, the number of flowers in a pis-
tillate cymule never exceeds that in the staminate cymules on the same plant
and, in fact, is frequently lower (see TABLE). There is some variability in cymule
flower number from tree to tree and even from branch to branch within some
species (e.g., L. celebica, L. dealbata, L. fenestrata, L. harlandii, L. lucida, L.
reinwardtii, and L. sootepensis).
In most cases all the flowers of a staminate cymule are fully formed at
anthesis. Only occasionally do clearly abortive flowers appear, as in Lithocarpus
buddii, where the central (uppermost) flower is fully developed but the two
lateral ones are abortive. Likewise, all the flowers of the pistillate cymules are
usually nonabortive at anthesis, but many of them abort later due to apparent
lack of pollination or fertilization. The abortive pistillate flowers are often
readily observed attached to or just below the cupule of a fully formed nut.
78 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Cymule characteristics of Lithocarpus.
Pistillate cymules Staminate cymules
SUBGENUS No. of No. of No. of No. of
species flowers evident flowers bracteoles
bracteoles
CYCLOBALANUS A. Camus
ta Barnett 3 5+ 3 5
; tia ta Hatusima ex Soep 3 3 3
clement A. 1,3 3 133 3
conferta Soep. 1 3 1 3
conocarpa Rehder ] l - -
cyclophora A. Camus = - 3 5
daphnoidea A. Camus 1 3+ - -
eichleri A. Camu 1 3+ - -
encleisacarpa A. Camus 1 3 3 5
ewyckii Rehder ] 3 153 3
korthalsii (Endl.) Soep 3 5+ 3 5+
lampadaria 5-7 7 5 7
lucida Rehde 1-3 1 3 3
lutea Soep. 1 3+ 3 3
macphailii A. Camus 1-3 5+ 3 5
mariae Soep. 1 3+ ] 3
meijeri Soep. 1 1,3 1,3 3,5
neorobinsonii A. Camus 1 3 ] 3
nieuwenhuisii A. Camus 1 3 - -
pattaniensis Barnett 1-3 5-7 1 3-7
hilippinensis A. Camus - - 153 133
rassa (Miq.) Rehder 1 3 1 3
reinwardtii A. Camus L533 35) 1,3 |
sericobalan Warb 3 or =
suffruticosa (Ridley) Soep l 3 3
GYMNOBALANUS A. Camus
havilandii Barnett l 3 l 3
kingiana amus 1 3 1 3
konishii hde 1 3 3 5)
ieue vie eer: 1 3 1 3
LIEBMANNIA A. Camus
hendersoniana A. Camus 3-5 5+ 3-5 5+
LITHOCARPUS Markgraf
beccariana A. Camus 1 ] l 1
maingayi Rehder = - 3 =
perakensis Soep 1 3 - -
turbinata (Stapf) Forman i 3 1 3
PACHYBALANUS A. Cam
amygdalifolia Hayata 1-3 3-7 3 5
nantoensis Haya 1 1 l 5
truncata Rehder & Wilson 5 ca. 7 5 ca. 7
PASANIA A. Camus
uddii (Merr.) A. Camus 3 3 3 3,7
eae ne Rehde 1 3 3+ 3
celebica Rehder 1,3 3+ 3 3
cooerts ae Rehder 1 3 1,3 3
ee ii Saga ex Hooker f.)
1 1 1
dasystachys eae ) Rehder 153 35D 3 3
dealbata A. iD 355) 9+
ee oe 1 ] 3,5 Sit
edulis Nakai 1 - | eae | 1,3
1987] KAUL, LITHOCARPUS 79
Cymule characteristics of Lithocarpus (continued).
Pistillate cymules Staminate cymules
SUBGENUS << OF No. of No. of No. of
eci flowers evident flowers bracteoles
bracteoles
ae Ee Hatusima ex Soep. 3-5 3 3-5 5+
lephan 1 3 1 3
See eee l 3 1 5
fenestrata Rehder 3 3 | 9+
formosana Hay 3} 3+ 3 5
garrettiana A. Camus 3 3+ 3 l
hancei Rehder 3 3+ 3 3
harlandii Rehder 1,3 3 3 B55
harmandii A as 3-5 3? 4-7+
kawakamii Haya 3-5 3+ -
papillifer een ex Soep. l 1 2,3 -
polystachya Rehder 1-3 3+ 3 35D
rufovillosa Rehder l 3 L335 3
sabulicola A. Camus 1 3+ 1 i]
scortechinii A. Camus 1 3+ 3 7+
soleriana Rehder 1 1 3 7+
sootepensis A. Camus 153) 3 3 3
spicata Rehder & Wilson 3 3 3 5+
sundaica Rehder & Wilson l 3+ 1,3 LS
ternaticupula Hayata 3 3+ 3 3
thomsonii Rehder 3 3+ 3 5
wallichiana Rehder 3 3 3 3
wrayi A. Camus 1 3+ 3 3
PSEUDOCASTANOPSIS Hickel & A. Camus
i amus 1 1 1235 4
SYNAEDRYS A. Camus
cor Rehde l 3 3 3
ko Taine “Haya l 3 i 3
pulchra Markgra 1 3 1 3
In multi-flowered cymules the sequence of anthesis begins with the central
(uppermost) flower and progresses to the subjacent pair and then to the lowest
pairs (see, for example, FiGures 29, 30). In three-flowered cymules the central
(upper) flower opens first and the subjacent pair soon afterward.
The bracteoles that subtend the cymules vary within subgenera and species
and sometimes between staminate and pistillate cymules in the same inflores-
cence (see TABLE). The number of bracteoles sometimes equals but more often
exceeds the number of flowers in the cymule, but it is rarely less (see TABLE).
In both pistillate and staminate cymules there is a single, usually larger,
primary bracteole centered below the cymule (see, for example, FiGures 16,
17). Subsequent bracteoles are ot smaller, sometimes progressively so, and
are usually paired across the c
The subtending bracteoles a o pistillate cymules sometimes grade into the
cupular bracteoles (hereinafter called ‘“‘scales”), but for the most part they are
80 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Bal
10
Ficures |-13. 1-4, Lithocarpus turbinata: 1, spike tip at anthesis, staminate cymules
above, alae aid perfect below (bracteoles in black and mene stippled in most
gu
aborti er; 4, immature cupule. 5, L. beccariana: spike tip at anthesis, stami
(upper) and pistillate (lower) cymules with I flower. 6-8, L. hendersoniana: 6, spike tip
at anthesis h3 to 5 flowers; 7, pistillate
cymule after anthesis, somewhat raised upon peduncle: 8, ane cupules, | abortive.
1987] KAUL, LITHOCARPUS 81
distinguished by their size (as the illustrations show), their greater thickness,
and occasionally their coloration. In some cases the uppermost bracteoles are
connate and form an entire or serrate border above the pistillate cymule, but
usually all the bracteoles are free. While some of the bracteoles are deciduous
or break off as the cupule expands after fertilization, the primary, and often
other, bracteoles persist below the matured cupule. At least the primary brac-
teole is usually readily apparent at cupular maturity, although it is often greatly
exceeded by the cupule and its scales.
The bracteoles of the staminate cymules are more easily seen because they
are not crowded by cupular scales. They are more often connate than are those
of the pistillate cymules, even within a species, and sometimes the connation
is so extreme that an accurate count is impossible (see, for example, FIGURES
89, 94). In rare instances the partially connate upper bracteoles enclose smaller
bracteoles that suggest a rudimentary cupule enclosing the staminate flowers
(see FiGuRE 62, uppermost cymule).
In staminate and pistillate cymules that have more than four bracteoles, it
is usually possible to enumerate the bracteoles and bracteole pairs at least to
the quaternary level or, if there is no connation, beyond. In many pistillate
cymules, however, the intergradation of subtending cymule bracteoles with the
cupular scales often makes such distinctions arbitrary beyond the primary or
secondary bracteoles. Even in the earliest developmental stages of a few species
that have been studied, it is not always possible to distinguish the first cupular
scales from the subtending bracteoles.
PISTILLATE CYMULE ORGANIZATION AND CUPULAR STRUCTURE
Among and within subgenera, there are great differences in the relative con-
tributions of the cupular scales to the mature cupules, which vary more than
those of Quercus. The flowers, cymules, and immature and mature cupules are
shown in FiGures 1-111 for 38 species from seven subgenera. Camus (1948),
in Volume 3 of her Ad/as, illustrated many species but did not include details
of cymule bracteoles or cupule development. Her plates are cited below to
complement my illustrations.
SUBGENUS LITHOCARPUS. In both species that were studied developmentally
(Lithocarpus beccariana, L. turbinata), the fruits are large, elongate, and figlike;
the cupule encloses the nut almost entirely (FiGuRE 4; Camus, 1948, pl. 355).
In L. turbinata there are three obvious cymule bracteoles, above which the
cupular scales are prominent at anthesis (FIGURE 1). These scales are pushed
upward as the cupular lamellae extend, and some of the lamellae become
excentric and disrupted in the process (FIGURES 2-4); the scales become widely
9, 10, L. cornea: 9, upper portion of spike at anthesis, showing staminate and pistillate,
1-flowered cymules; 10, mature cupule with 2 abortive cupules fused to it. 11-13, L.
pulchra: 11, spike tip at anthesis, showing staminate and perfect flowers; 12, lateral view
of pistillate cymule some time after anthesis, showing 2 f 3 bracteoles; 13, immature
cupule, showing scale-bearing tubercles and all 3 bracteoles. Figures 1, 2, 4, 10, 13, x 2;
Figure 3, x 0.3; Figures 5-9, 11, 12, x 4
82 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
separated, but many of them persist on the mature cupule (FIGURE 3). At
maturity the primary bracteole also usually persists, but the secondary brac-
teoles do not; however, some scales remain near the base of the cupule. Litho-
carpus beccariana has but one bracteole below each pistillate cymule, and above
it are the cupular scales (FIGURE 5). At maturity the cupule is virtually scaleless,
and the scale-bearing lamellae are greatly extended (Camus, 1948, pl. 355).
FiGure 3 shows an apparently adventitious, abortive flower and cupule borne
well up on the mature, nut-enclosing cupule.
SUBGENUS LIEBMANNIA. The three-or-more-flowered pistillate cymules of
Lithocarpus hendersoniana have an obvious primary bracteole below them,
and a low ring of overlapping bracteoles above that forms a pointed cowl at
the distal end of the cymule (Ficure 6). After pollination the cymule becomes
pedunculate, and the primary bracteole is elevated on the peduncle (FIGURE
7). The cupular lamellae are continuous at first but later become interrupted
(FiGure 8), perhaps because of the rupturing stresses of diametric growth. The
mature nut is included in the cupule.
SUBGENUS SYNAEDRYS. The cupular scales of Lithocarpus cornea are prominent
at anthesis, and the three subtending bracteoles are clearly distinguished (FIGURE
9). Enormous expansion of the cupular lamellae is accompanied by great growth
in the scales, which become appressed and fused to the lamellae (FiGURE 10).
The mature cupule covers most of the nut, except for a broad polar area. FIGURE
10 shows two abortive flowers and cupules attached at the base of the cupule.
In Lithocarpus pulchra the three bracteoles of the one-flowered cymule are
evident at anthesis (FiGures 11, 12) and in fruit (FiGuRE 13), but an additional
ring of bracteoles that surrounds the cupular scales quickly loses its identity as
the cupule enlarges. The scales of the mature cupule are widely separated, each
of them raised upon a mound of cupular tissue (FicureE 13; Camus, 1948, pi.
370).
SUBGENUS PACHYBALANUS. In both Lithocarpus amygdalifolia and L. truncata
at least seven bracteoles subtend the multi-flowered cymules (Ficures 14, 16);
in the former species the one-flowered cymules have but three (FicureE 14).
There are other bracteoles within the multi-flowered cymules. The cupular
scales are hidden at anthesis by all these bracteoles, but they quickly become
evident afterward. The mature cupule encloses much of the nut and is adorned
with large, widely spaced cupular scales (Camus, 1948, pl. 377). The cymules
of L. nantoensis have just one bracteole, the primary, and above it is a ring of
presumably fused bracteoles that entirely encircles the cupule (FiGurRE 15).
SUBGENUS GYMNOBALANUS. Three distinct bracteoles subtend each one-flow-
ered cymule of Lithocarpus havilandii at anthesis (FIGURE 19), and they usually
persist below the mature cupule (Fiures 20, 2 1). The numerous cupular scales
are prominent at anthesis (FIGURE 19) but are mostly adnate to the cupule at
maturity (FIGURE 21), at which time they are not obviously arranged in con-
centric rings. The nut is enclosed by the cupule when immature but is mostly
exposed at maturity (FIGUREs 20, 21).
The one-flowered cymules of Lithocarpus konishii and L. lauterbachii have
1987] KAUL, LITHOCARPUS 83
\
gd
3
ed
U 22
Ficures 14-24. 14, Lithocarpus amygdalifolia: spike tip at anthesis, staminate and
pistillate cymules 1- to 3-flowered, flowers removed from uppermost staminate cymule
to reveal 5 bracteoles. 15, L. nantoensis: spike tip at anthesis, staminate and pistillate
i f
pistillate cymules. 24, L. lauterbachii: near-terminal segment of spike at anthesis, all
cymules 1-flowered. Figures 14, 20, 21, x 2; all others, x 4.
84 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
fs
Na =,
ps
Ley x
FiGures 25-33. 25-28, Lithocarpus lucida: 25, segment of staminate spike with 2
3-flowered cymules, upper | with flowers removed to reveal 3 bracteoles; 26, segment
of pistillate spike with | 1-flowered and 2 3-flowered cymules after anthesis; 27, 2
1-flowered cymules with immature fruits, cupular scales evident; 28, immature fruits,
older than those of Figure 27, cupular lamellae now devoid of scales, abortive fruit visible
at lower end of lower cupule. 29-33, L. lampadaria: 29, segment of staminate spike,
1987] KAUL, LITHOCARPUS 85
a primary bracteole and, above it, a ring of free but overlapping bracteoles
around the cupule (FiGuRES 23, 24). In both species the mature cupule covers
less than half of the broad, low nut, and it is heavily invested with overlapping
cupular scales (Camus, 1948, p/. 385).
SUBGENUS CYCLOBALANUS. The mature cupule is often devoid of cupular scales
(FiGurREs 28, 33, 35, 41, 44, 55-57, 66), or it may have weakly developed scales
that are widely separated (Ficures 47, 51). In all the species of this subgenus
illustrated, the early developmental stages clearly show the presence of cupular
scales (FIGURES 27, 31, 32, 34, 38, 42, 43, 45, 46, 49, 50, 52, 59, 65). Many
scales are deciduous or become distorted and exceeded by the massive growth
of the cupule, and the mature cupule is then naked or nearly so. The cupular
lamellae are more or less concentric in many species, but in a few they are not
distinguishable at maturity (FiGurREs 33, 35). In these the mature cupule consists
of random or vaguely concentric scaleless enations. In some multi-flowered
cymules the lowest few lamellae embrace all the flowers (FiGuRES 26, 33, 39),
but each flower eventually develops its own cupule (Ficures 28, 33, 39). Other
multi-flowered cymules lack such collectively embracing lamellae, and the
flower cupules are distinct from the earliest stages (FIGURES 49, 50, 64).
The pistillate cymules of Lithocarpus lucida (FIGURES 26-28) are one- or
three-flowered. All three flowers do not ordinarily mature in the latter case
(FiGuRE 28), nor do some of the one-flowered cymules. There is but one dis-
cernible subtending bracteole below each cymule, whether it 1s one- or three-
flowered. Above it is a ring of tissue that perhaps represents fused bracteoles
and that forms the first lamella of the cupule embracing all the flowers. The
next structures to appear are partial lamellae that collectively embrace all the
flowers (FiGureEs 26, 27). It is not until well after pollination that the truly
concentric, cupular lamellae arise in acropetal sequence. The scales are readily
visible at these early stages. As the cupules near maturity, the scales have fallen
or have become split and stretched beyond recognition, the cupule then appears
to be scaleless (FIGURE 28). The massive growth of the cupular lamellae causes
distortions among the contiguous cupules so that at least the first-formed (low-
est) lamellae are often distinctly excentric. Abortive flowers become partially
or completely buried in the maturing cupule (e.g., the central flower in the
upper cymule and the lateral flowers in the lower cymule of FiGure 28). At
maturity the cupule covers less than half of the nut (Camus, 1948, pl. 386).
In Lithocarpus reinwardtii the cymules are also one- or three-flowered (FIGURES
37-40). The one-flowered cymules are subtended by three distinct bracteoles,
above which the scale-bearing tric] ll ppear in acropetal sequence.
The last few lamellae to form are weakly developed and show no external
evidence of scales (FiGuRES 40, 41). The mature cupule is scaleless, although
showing 5-flowered cymules, upper | with flowers removed to reveal 7 bracteoles; 30,
segment of spike showing mixture of staminate and pistillate cymules, all multi-flowered;
31, 32, maturing pistillate cymules after anthesis, some flowers and their cupules abortive;
33, mature fruit with 2 basal, abortive flowers in cupules. Figures 28, 33, x 2; all others,
x 4,
86 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
FiGures 34-41. 34, 35, egress aggregata: 34, segment of pistillate spike at
thesis, showing pedunculate 3-flowered cymules and their bracteoles; 35, segment of
Pistllat spike bearing mature fruits | abortive cupules. 36-41, L. reinwardtii: 36,
howing |- and 3-flowered cymules; 37, 1-flowered
pistillate cymules at anthesis: 38-40, maturing cupules with evident scales: 41, mature
ead (nut removed), showing essentially scaleless lamellae. Figure 41, x 2; all others,
1987] KAUL, LITHOCARPUS 87
some evidence of scales can be seen in the lowest few lamellae (FiGurE 41).
The three-flowered cymules, which are less common in my specimens than the
one-flowered, are subtended by at least five distinct bracteoles (FIGURE 39),
and there are other structures at the base of the cymule that may also represent
subtending bracteoles. The first two lamellae to form surround all three flowers,
but later lamellae embrace only one. Further details are shown by Camus (1948,
pl. 397
The one-flowered cymules of Lithocarpus bullata and L. ewyckii are also
subtended by three obvious, distinct bracteoles (FIGURES 42-47) that persist
below the mature cupule. The first lamella to form above the bracteoles of L.
bullata bears a few scales (FiGuRES 42, 43) that persist to maturity of the cupule.
l llae have more scales, many of which persist but become widely
separated as the diameter of the cupule increases (Figure 44). The uppermost
lamellae are scaleless from their earliest stages. The first lamellae of L. ewyckii
are more irregular than those of L. bullata, but they, too, are scaly. The later
lamellae are regular and concentric and retain many of their scales into ma-
turity, at which time the scales are widely spaced, sometimes reflexed, and
often broken (FIGURE 47).
The cymules of Lithocarpus macphailii are distinctly pedunculate at anthesis
(Ficures 49, 50), but the peduncle does not lengthen very much as the cupule
matures. There are three basal bracteoles (shown in lateral and ventral views
in Ficures 49 and 50, respectively). Another series of distinct bracteoles is
evident at the distal end of the peduncle, just below the individual flowers
(these are shown in black for emphasis in FIGURES 48-51). These, too, persist
into maturity of the cupule (FiGuRE 51), and they are readily distinguished by
their location, thickness, and color from the other bracteoles below the flowers.
Each flower develops its own cupule, but there is a loose ring of distinct or
partially fused bracteoles that embraces all the flowers below their cupules
(Figures 49, 50). As the cupular lamellae expand, the scales become widely
separated but (as in the other species of this and many other subgenera) do not
enlarge (FiGuRE 51). At full maturity, only a small upper portion of the nut is
visible (Camus, 1948, pi. 407).
Most of the cymules of Lithocarpus encleisacarpa are one-flowered, and each
is subtended by three bracteoles (FiGuRESs 52-55). At anthesis the cymules are
sessile, but they become pedunculate by elongation of the first few lamellae of
the cupule (FiGurEs 52-56), succeeding lamellae increase in diameter more
than in length, and the mature cupule is turbinate. The cupular scales are evident
at anthesis (FiGuRE 52) but are barely apparent when the cupule matures
(Ficures 55-57). As the nut enlarges, the cupule ruptures, usually along three
irregular arcs that cut through some of the upper lamellae (FIGURES 56, 57;
Camus, 1948, p/. 406).
The pistillate cymules of Lithocarpus neorobinsonti have one primary brac-
teole at the base (FiurEs 59, 61); above this is an irregular lamella that may
represent other, fused bracteoles. The somewhat irregular lamellae (even the
uppermost, poorly developed ones) of the cupule retain their scales to maturity.
The upper part of the cupule ruptures irregularly as the nut enlarges, with the
tears extending only into the region of weak development of the lamellae
(Figures 60, 61; Camus, 1948, p/. 4/0).
Succeeding
88 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Ficures 42-51. 42-44, Lithocarpus bullata: 42, 43, lateral and basal views of |-flowered
pistillate cymule somewhat beyond anthesis, showing bracteoles and young cupule; 44,
1987] KAUL, LITHOCARPUS 89
Ficures 52-61. 52-57, Lithocarpus encleisacarpa: 52-54, |-flowered pistillate cy-
mules at anthesis and in early fruit, 3 bracteoles evident below each cymule, cupular
scales evident at anthesis (FiGURE 52) but becoming remote and ruptured as cupule
matures; 55, mature cupule with see scaleless lamellae; 56, 57, dehiscing cupule in
eel and polar views. 58-61, L. neorobinsonii: 58, segment of staminate spike at
anthesis, cymules 1-flowered and with 3 bracteoles; 59, pistillate, 1-flowered cymule after
anthesis, cupular scales evident; 60, nearly mature cupule with scales now remote and
lamellae weakly developed; 61, mature cupule, upper portion dehisced irregularly and
revealing nut. Figures 52-54, 58, x 4; all others, x 6.
3-flowered cymules at anthesis, lateral and basal views (cymules pedunculate from an-
thesis); 51, nearly mature cupule with persistent bracteoles and remote cupular scales.
Figures 47, 51, x 2; all others, x 4
90 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
FiGurES 62-71. 62-66, pene? pattaniensis: 62, segment of staminate spike,
showing |-flowered cymules, each w or more bracteoles, upper 2 cymules with flower
3-flowered pistillate cymules at anthesis; 65, 1-flowered pistillate cymule some time after
anthesis, showing beginnings of lamellar growth of cupule, scales evident; 66, nearly
mature cupule with lamellae prominent, scales now remote and r she dicate bracteoles
evident. 67-70, L. rufovillosa: 67-69, maturing 1|-flowered pistillat with 3 brac-
teoles, scales prominent near anthesis (FIGURE 67) but lamellae prominent in fruit (FIGURES
1987] KAUL, LITHOCARPUS 91
Above the obvious primary bracteole of the one- and three-flowered cymules
of Lithocarpus pattaniensis are other, basally fused bracteoles that encircle the
flower(s) (FIGURES 63-65). There are usually four of these in the one-flowered
cymules but more in the three-flowered ones. Some of these bracteoles persist
into maturity of the cupule (FiGuRE 66). The cupular scales are evident at
anthesis (FIGURES 63, 64) but are tiny and often ruptured on the massive
lamellae of the mature cupule (FIGURE 66). The scales are adjacent at anthesis
but become separated during cupular expansion (FIGURES 65, 66). When the
cupule is fully mature, it reveals a small portion of the nut (Camus, 1948, p/.
517)
The mature cupule of Lithocarpus aggregata does not show the obvious
lamellae of the above-described species. Instead, it bears vaguely defined rows
of enations that carry little or no evidence of cupular scales (FiGuRE 35).
However, cupular scales and lamellae are clearly evident in earlier develop-
mental stages (Figures 34, 35). At anthesis the three-flowered cymules are
pedunculate, and the peduncle is evident through maturity of the fruit (FIGURE
35). There are three bracteoles under each cymule, and above them are two
more, each near a lateral flower; there is no bracteole immediately below the
central flower (FIGURE 34). These persist into fruit. There is one lamella (or
sometimes two) encircling the peduncle, but above it the lamellae embrace
single flowers (FiGuRE 35). After several obvious lamellae have formed, the
succeeding ones are, from their inception, indistinct; it is they that form the
irregular rows of enations in the upper part of the cupule.
The pistillate cymules of Lithocarpus lampadaria often have five flowers,
but more or fewer are common. Below each cymule is a single, distinct primary
bracteole, above which is a series of six or so free but overlapping paired
bracteoles (FIGURE 30). The primary bracteole and some of the others persist
into fruit, but they are often completely distorted by the massive growth of the
cupules and the resulting juxtaposition of the abortive flowers (FIGURES 32,
33). Of the hundreds of fruiting cymules examined, none bore more than three
fully developed nuts, and most had none, one, or two.
SUBGENUS PASANIA. The cymule bracteole patterns of this subgenus resemble
those of the other subgenera, but the cupular ornamentation 1s very diverse.
There are three bracteoles subtending the one-flowered pistillate cymule of
Lithocarpus rufovillosa (FIGURE 68), and they persist into fruit. The cupular
scales are evident at anthesis (FIGURE 67), and soon thereafter their alignment
in rows is apparent (FiGuRE 68). The massive growth of the lamellae separates
the scales, many of which fall, leaving the cupule barely scaly at maturity
(FiGures 69, 70). In fact, many of the lower lamellae are scaleless (FIGURES
69, 70).
The distinctively pedunculate one- and three-flowered cymules of Lithocar-
69, 70); 70, segment of mature fruit, cupule covering about half of nut, scales retained
only on upper lamellae. 71, L. sootepensis: segment of pistillate spike very soon after
anthesis, cymules 1- and 3-flowered and with 3 bracteoles, peduncle evident at anthesis
and eventually carrying mature fruits. Figures 66, 70, x 2; all others, x 4
92 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Za
a
~
4
ao A
S >
5 AILS
FiGures 72-83. 72-74, Lithocarpus wrayi: 72, segment of stami t anthesis,
showing |- and 3- Acweica cymules, uppermost | with flowers removed to reveal 3
bracteoles; 73, spike tip, with distal, 1-flowered staminate cymules and 1- and 2-flowered
bracteol rm; 76, 1-flowered pistillate cymule at anthesis, 3 bracteoles shown; 77,
pistillate cymule after anthesis, showing extensive growth of cupular scales. 78, 79, L.
1987] KAUL, LITHOCARPUS 03
pus sootepensis each have a basal primary bracteole and a pair of secondary
bracteoles that become elevated on the elongating peduncle (FiGure 71). The
cupular scales are well developed soon after anthesis and are evident in the
mature cupule (Camus, 1948, p/. 4/6).
Three bracteoles subtend the one- or several-flowered cymules of Lithocarpus
wrayi (FIGURE 73), and at least the primary bracteole can be seen below the
mature cupule. The scales of the cupule are large at anthesis, and they remain
prominent on the cupule in fruit, eventually becoming reflexed (Camus, 1948,
pl. 441). The cupule does not show lamellae, although the persisting, subulate
scales are aligned in concentric rows (FIGURE 74
A primary bracteole and a pair of secondary ones subtend the pistillate
cymules of Lithocarpus hancei (FiGuRE 84). Most of the cymules have three
flowers, but some of the more distal ones are one-flowered (FiGuRE 84). A ring
of connate bracteoles surrounds the flowers, and within that (but not visible
in FiGureE 84) are the young cupules. As the nut and cupule begin to grow, the
cupular scales emerge (FiGuRE 86, lower, abortive cymule); at maturity the
relatively small cupule shows irregular rings of annular enations, most of which
bear a tiny cupular scale (FIGURE 86, mature nut and cupule; Camus, 1948,
pl. 415).
The numerous scales of Lithocarpus papillifer (FIGURE 87), so evident at
anthesis, remain small and adpressed on the mature cupule. There is but one
obvious bracteole below each one-flowered cymule.
The long cupular scales of Lithocarpus garrettiana are evident from anthesis
onward (FiGures 79, 90-92), elongating considerably during cupular growth
so as to be trichomelike at maturity. In the lower part of the mature cupule,
the scales are in concentric rows (FIGURE 92); those higher up are usually
crowded, and their arrangement in rows is not evident. The drying, dehiscing
cupule splits open along three radial arcs (FiGURE 91) that extend halfway or
less down the cupule, the upper part of the cupule sometimes breaking away
in a crudely circumscissile dehiscence (FIGURE 92). There is some variation in
dehiscence pattern of the cupule; only the usual one is illustrated in FiGuREs
91 and 92 (cf. Camus, 1948, p/. 434).
As in many species of subg. Cyclobalanus, the mature cupule of Lithocarpus
soleriana has concentric lamellae bearing vestiges of cupular scales (FIGURE
96; Camus, 1948, p/. 467). The primary bracteole subtends the one-flowered
cymule and is surmounted by a ring of partially connate bracteoles that enclose
the cupule; the cupular scales are evident at anthesis (FIGURE 95). With ex-
garrettiana: 78, segment of stami , Showing 3-flowered cymules with | bracteole;
79, segment of pistillate spike at anthesis, showing 3- and 4- flowere d cymule es, each
bracteoles (not in black). 81, L. harlandii: segment of staminate spike at anthesis with
1- and 3- powered cymules, uppermost l with ee removed to reveal bracteoles. 82,
nil, hesis, showing | -flowered cymules,
uppermost 1 with flower removed to reveal 3 i ; 83, segment of pistillate spike
at anthesis showing 1|-flowered cymules with 3 bracteoles. Figures 74,77, x 2; all others,
x 4,
94 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Ficures 84-92. 84-86, Lithocarpus hancei: 84, spike tip at anthesis, showing few
distal, staminate, |-flowered cymules, each with 3 bracteoles, pistillate cymules 3-flow-
pistillate spike, 1 cymule with only abortive flowers and cupules, mature cupule with
scale-bearing enations. 87, L. papillifer: segment of pistillate spike soon after anthesis,
1-flowered cymules each with | obvious bracteole, numerous styles on each flower. 88,
staminate spike at anthesis, most cymules 5-flowered, lowest | with 3 flowers removed
to reveal complex bracteole pattern. 90-92, L. garrettiana: 90, flower in cupule soon
after anthesis, cupular scales already very long; 91, mature cupule invested with elongate,
recurved scales and showing 3 lines of dehiscence from upper pole; 92, mature cupule,
dehisced upper portion fallen away. Figures 86, 91, 92, x 2; all others, x 12
1987] KAUL, LITHOCARPUS 95
pansion of the cupule as maturity nears, the scales are separated and often
ruptured, but most of them persist.
The pistillate cymules of Lithocarpus harmandii are among the most complex
in the genus. They have one to seven flowers and are surrounded by a mass
of bracteoles (FiGUREs 97, 98). Below each cymule is a single obvious primary
bracteole, above which is a complex ring of barely connate bracteoles. At
anthesis the cupular scales are not evident because they are hidden by the ring
of bracteoles (FIGURE 97), but soon thereafter they become prominent (FIGURE
98). Each flower develops its own cupule, but only one or two—very rarely
three — mature into fruit. The abortive flowers continue to grow for some time
and develop obvious but small cupules (FiGuRE 100). The cupule surrounding
a mature nut is a mass of more or less concentrically arranged enations, most
of which bear a tiny cupular scale (FiGuRE 100; Camus, 1948, pls. 470, 471).
The pistillate cymules of Lithocarpus elegans are also complex. They usually
hold three to five flowers, with a few having one or six or more (FiGURE 101).
A primary bracteole and a lateral pair of secondary ones are attached to the
elevated buttress that bears the flowers (Figures 101, 103). There are no other
obvious bracteoles in the cymule at anthesis, but there are faint ridges on the
buttress that suggest a ring of reduced bracteoles (not visible in FiGure 101).
There are no readily discernible cupular scales at anthesis, but they appear
soon thereafter. Their arrangement in concentric rows is then evident. The
rings of scales are very tightly compressed, and the scales are appressed but
readily visible in the mature cupule (FiGuRE 103; Camus, 1948, p/. 481). As
the nut matures, the partially enclosing cupule ruptures along four or five arcs
(FicureE 103).
One primary bracteole and a pair of lateral bracteoles, one below each lateral
flower, are characteristic of the three-flowered cymules of Lithocarpus wallichi-
ana. There is also a ring of partially connate bracteoles that partially surrounds
the cymule (FiGuRE 105). The cupular scales are not entirely concealed by these
bracteoles at anthesis, and they later | t (Ficure 106). Although
it is not obvious in FiGureE 106, the scales a are aligned in concentric rows. At
maturity of the cupule, the scale-bearing concentric lamellae are evident; they
have persistent, separated, torn scales (FIGURE 107; Camus, 1948, p/. 503).
The abortive flowers and cupules are shown in Figure 107. Any one of the
three flowers in a cymule can mature into a fruit. The upper cymule in FIGURE
107 shows the matured cupule of the central flower (the nut is removed to
show the scar) subtended by two abortive lateral flowers; the lower cymule has
one abortive and one fertile lateral flower and an abortive central flower.
Occasionally, more than one flower matures a nut
Although the cupular scales of Lithocarpus scortechinii are hidden by the
bracteoles at anthesis (FIGURE 76), they quickly become prominent (FIGURE
77); by cupular maturity they are long and reflexed (Camus, 1948, pi. 442).
The mature cupule covers much less than half of the nut. There are one primary
and two distinct lateral bracteoles below the one-flowered cymule, and a ring
of barely connate bracteoles above that (FiGURE 76). When the cupular scales
enlarge, the ring of bracteoles is not readily distinguishable from the scales
(FIGURE 77).
96 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Ficures 93-100. 93, Lith | segment nea th
4 cymules staminate, with 3 to 5 flowers and 5 to 7 Gace ot all visible fa
pistillate cymules with | flower and | bracteole. 94-96, L. soleriana: 94, segment of
staminate spike with I- and 3-flowered cymules, 2 with flowers seein to show nu-
merous bracteoles; 95, segment of pistillate spike at anthesis, showing | -flowered cymules,
each with | bracteole, cupular scales prominent; 96, mature cupule covering about half
of nut, lamellae somewhat scaly. 97-100, L. harmandii: 97, 98, segments of pistillate
1987] KAUL, LITHOCARPUS o)
As in Lithocarpus scortechinii and other species, the mature cupule of the
only American member of the genus, L. densiflora, covers little of the nut and
is thickly invested with rather long, often recurved scales (Camus, 1948, pi.
444). The cymules are one-flowered and are subtended by a large primary
bracteole (FIGURE 93); above this is a ring of slightly overlapping bracteoles
that enclose the cupular scales, which are evident at anthesis (FIGURE
The one-flowered pistillate cymules of Lithocarpus sabulicola have three
bracteoles, one primary and two secondary, and there is a ring of strongly
connate bracteoles that surrounds the remainder of the cymule. The ring does
not entirely conceal the cupular scales at anthesis (FiGURE 83). At maturity the
nut projects well beyond the scaly cupule (Camus, 1948, pl. 464).
The cymule bracteoles of Lithocarpus dealbata are not clearly distinguishable
from the cupular scales. Although the primary bracteole is easily observed, the
secondary ones are less so (FIGURE 88). Beyond them is a series of structures
that are not clearly bracteoles or scales. The cupule encloses most of the nut
at maturity, and it is invested with concentric rows of widely spaced, appressed,
slightly elongate scales (Camus, 1948, pis. 450, 451).
SUBGENUS PsEUDOCASTANOPSIS. The cymules of Lithocarpus fissa subsp. fissa
are one-flowered, and each has one bracteole (FiGureE 109). At anthesis the
cupular scales and lamellae are obscured, but they are evident at maturity, at
which time the cupule dehisces and the nut emerges (Figures 110, 111).
THE STAMINATE CYMULES
The staminate cymules are borne on staminate spikes, as well as above and
below the pistillate cymules on mixed-sex spikes (Kaul & Abbe, 1984). On the
latter spikes there are sometimes a few cymules that bear both staminate and
pistillate flowers at the area of transition from entirely pistillate to entirely
staminate cymules. The flowers in that area may be perfect, while those away
from it are imperfect. Such transitional conditions are especially evident in
species with multi-flowered cymules
The staminate cymules are subtended by one or more bracteoles whose
number and arrangementare the same as or different from those of the pistillate
cymules of the same species. Often there are more bracteoles subtending the
staminate than the pistillate cymules (see TABLE).
Ficures | and 11 show the one-flowered staminate cymules of Lithocarpus
turbinata and L. pulchra on the rachis beyond the pistillate cymules. Eac
staminate cymule has one long primary and two shorter secondary bracteoles,
a condition often found in one-flowered staminate cymules in other subgenera.
However, by contrast, the one-flowered cymules of L. beccariana (FIGURE 5)
have only a primary bracteole. The situation is more complex in L. hender-
spikes at and shortly after anthesis, , Fespectively, eymuls 3- to 7-flowered, each with |
prominent bracteole, wel flowers of uppermost cymules
of Figure 97; 99, segment of staminate spike just before anthesis, cymules 7-flowered,
each 3-bracteolate; 100, mature cupule and nut, each scale borne on enation. Figures
96, 99, 100, x 2; all others, x 4.
98 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Ficures 101-111. 101-103, Lithocarpus elegans: 101, segment near tip of androgy-
nous spike at anthesis, upper 4 cymules staminate, uppermost with all 3 flowers removed
to show oo Distilate cymules 3- to > flowered, each with 3 bracteoles and raised
upon buttres , 5-flowered cymules subtended
y numerous oe upper 2 cymules with flowers removed; 103, mature, fruit-
nearing anthesis, each cymule with 3 flowers and 3 bracteoles; 105, segment of pistillate
spike at anthesis, each cymule with 3 flowers and 3 bracteoles, cupular scales evident;
106, pistillate cymule after anthesis; 107, mature cymules, each with 2 abortive flowers
nd cupules, mature cupule covering about half of nut and barely scaly. 108-111, L
1987] KAUL, LITHOCARPUS 29
soniana (Ficure 6): above the primary bracteole there is a series of low, basally
connate bracteoles that encircle the entire cymule, just as they do in the pistillate
cymules of that species. In all these species the bracteole pattern is the same
in staminate and pisullate oe
The one- and three-flow te cymules of Lithocarpus amygdalifolia
and L. nantoensis (Ficures 14, 15) have five bracteoles, with the primary
always the largest. The five-flowered staminate cymules of L. truncata usually
have seven bracteoles (FIGURE 17). The quaternary ae extends completely
over the distal end of the cymule (not visible in figur
In the three species illustrated from subg. Coan the staminate
cymules are one- and three-flowered and three-bracteolate, and three-flowered
and five-bracteolate (FiGuRES 18, 22, 24)
In the large subgenus Cyc/obalanus the bracteoles of the staminate cymules
range from three to seven or more per cymule. A complex example is shown
for Lithocarpus lampadaria in Figure 29. The staminate cymules are mostly
five-flowered, and there is an elongate primary bracteole below each one. Above
it, in pairs, are six additional bracteoles, some overlapping and some not
(FIGURE 29, top). Four of the flowers of the cymule have one or more bracteoles
beside them, but the distal flower does not. The bracteole pattern of the
pistillate cupule is likewise complex (FIGURE 3
Simpler bracteole patterns exist in Lithocarpus reinwardtii, where both the
one- and the three-flowered staminate cymules have three bracteoles (FIGURE
36), as do some of the pistillate cymules (FiGurE 37). In L. macphailii the
three-flowered staminate cymules have five bracteoles, and the pistillate cy-
mules have that many or more (FiGuREs 48-51). The simplest case is that 0
L. neorobinsonii (FiGuRE 58), in which the staminate and pistillate cymules
both have one flower and three bracteoles.
The bracteole pattern is somewhat complex in Lithocarpus pattaniensis be-
cause, although the cymules are always one-flowered, there are three or some-
times more bracteoles present, even on the same specimen (FIGURE 62). When
the single flower is removed from the bracteoles, as in the upper two cymules
in FiGuRE 62, it can be seen that the secondary bracteoles are slightly confluent
above the cymule, where they form a point that suggests another, reduced
bracteole. Furthermore, within that encircling series of bracteoles there is some-
times a second set of four (two to six) tiny ones that suggest a rudimentary
cupule (FIGURE 62, uppermost cymule).
The largest subgenus, Pasania, also has a great range of bracteole patterns
in the staminate cymules. Some three-flowered cymules have but one bracteole
(e.g., in Lithocarpus garrettiana, FiGuRE 78), and some have three bracteoles
(e.g., in L. lucida, Figure 25; L. wrayi, Figure 72; L. hancei, Ficure 85; and
L. wallichiana, Figure 104). Some cymules with five or more flowers also have
fissa: 108, 109, segments of staminate and pistillate spikes at anthesis, each cymule with
1 flower, staminate with 3 or more bracteoles, pistillate with 1; 110, cupule nearing
maturity and showing early signs of dehiscence; 111, mature, dehisced cupule revealing
part of nut, lamellae prominent and barely scaly. Figures 103, 107, 110, 111, x 2; all
others, x
100 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
only three bracteoles (e.g., in L. harmandti, Figure 99), but so do some one-
flowered cymules (e.g., in L. wrayi, Figure 73; L. sabulicola, Figure 82, L.
hancei, FiGuRE 84
omplex bracteole patterns in the staminate cymules of subg. Pasania are
illustrated here by seven species. Lithocarpus scortechinii (FiGURE 75), L. har-
landii (Ficure 81), L. dealbata (Ficure 89), L. soleriana (FiGuRE 94), and L.
elegans (Ficures 101, 102) illustrate a common arrangement: an identifiable
primary bracteole and usually an identifiable pair of secondary ones. Beyond
these three bracteoles is a series of smaller, sometimes irregular ones that are
not always obviously paired. At the distal end of the cymule, the bracteoles
are reduced and apparently fused; they usually surmount the cymule. Such
complexity occurs in these species in one-, three-, and multi-flowered cymules,
as shown in the figures. The three- and five-flowered staminate cymules of L.
densiflora have five and seven bracteoles, respectively.
e€ most complex staminate bracteole pattern among the species studied is
that of Lithocarpus fenestrata. In addition to having a series of complex brac-
teoles similar to those of the species discussed in the preceding paragraph, each
flower is subtended by a whorl of small bracteoles that suggests a rudimentary
cupule (FiGuRE 80, upper two cymules, the small bracteoles not darkened).
Lithocarpus fissa, of subg. Pseudocastanopsis, has one-flowered staminate
cymules, each with four subtending bracteoles, the fourth one located at the
distal end of the cymule (FiGure 108)
DISCUSSION
Some aspects of the bracteole patterns and the floral arrangement support
the interpretation that the groups of flowers provisionally called cymules are
actually that. Evidence is provided by the sequence of opening of the flowers
in both staminate and pistillate cymules. In every instance the distal flower
opens first, with the subjacent pair next, and the lowest pair last (i.e., the
sequence 1s strictly basipetal within the cymule). Where more than five flowers
are present in a cymule, the sequence of opening beyond the fifth flower is also
generally basipetal, but the pattern is less obvious.
The primary bracteole and the paired secondary, tertiary, and subsequent
bracteoles, as well as the absence of a bracteole immediately below the central
flower, all suggest a condensed cyme. When the cymule has a single flower,
sometimes one and sometimes three or more bracteoles subtend it. Where the
number of bracteoles exceeds the ‘number oF flowers subtended, it is possible
that each excess bracteol lost flower
or branch of a complex, now-condensed branching system
The bracteoles subtending the pistillate cymules are anaoubieay homolo-
gous with the cupular scales above them. The bracteoles merely represent the
lowest bracteoles of the condensed branching system, while the scales are the
bracteoles of the branches whose phylogenetic condensation formed the cupule.
Some evolutionary increase in scale number could have occurred after steril-
ization of bracteoles and while the cupule was evolving.
1987] KAUL, LITHOCARPUS 101
Fey and Endress (1983) interpreted the fagaceous cupule as a complex, cy-
mose branching system with shortened, united axes and with persistent brac-
teoles that form the cupular scales. They showed that, at least in earlier on-
togenetic stages, the scales are regularly arranged in a pattern suggesting that
of branched cymes. The subtending bracteoles discussed in this paper are then
merely the lowest bracteoles of the much-reduced cymose system (cf. Fey &
Endress, 1983, fig. 2/). In many pistillate cymules the subtending bracteoles
intergrade with the cupular scales, as would be expected with this interpretation.
In every instance where the ontogeny has been observed, the cupular scales
are present at anthesis (but are sometimes obscured by the bracteoles). They
may persist and even enlarge with the cupule, fully investing it at maturity, as
in many species of subg. Pasania. In extreme cases (e.g., Lithocarpus garret-
tiana, FiGuRE 91) the scales elongate greatly and the cupule becomes coarsely
hirsute. They may also persist without enlarging, so that the mature cupule has
obvious but small and often widely spaced scales, as in many species of sub-
genera Lithocarpus, Synaedrys, and Gymnobalanus and in some species of subg.
Cyclobalanus. The extreme condition is seen especially in the last subgenus,
where in many species the scales are lost during ontogeny because they either
fall from the cupule or become ruptured during cupular expansion. Such mature
cupules essentially lack scales, consisting of massively enlarged axial tissue of
the cupule. The morphological nature of this axial tissue is yet to be defined,
however.
Special conditions exist in subg. Synaedrys and in a few species of other
subgenera. For example, in Lithocarpus cornea of subg. Synaedrys (FiGureE 10),
the scales enlarge with the cupule and become totally adnate to it so that at
maturity the cupule is mostly covered by them. In L. pulchra of the same
subgenus (FiGuRE 13), the scales or scale tips become elevated on tubercles,
which completely cover the cupule. The morphological nature of these tubercles
is unknown.
Soepadmo (1970) studied the vascular anatomy of the cupule of Lithocarpus
and found the same vascular organization as that in the Quercus cupule (Kaul,
1985, fig. 36). In pistillate cymules that mature more than one fruit, the cupules
usually become connate laterally. When this occurs, the vascular systems of
the individual cupules remain distinct in the fused, “interseminal” cupular
walls. The more or less regular patterns of dichotomous branching of the cupular
vascular bundles, ultimately serving each scale with a vascular trace, could be
interpreted as evidence of the cymose history of the cupule (Kaul, 1985), but
the extreme condensation in the cupule and the lack of intermediate forms
make any interpretation of vascular evidence tentative.
The function of the cupule is probably protection, first of the flower and
later of the fruit, and in this aspect its evolutionary history resembles that
postulated for the inferior ovary. However, the ovary of Lithocarpus is inferior
and the ovary wall at anthesis is not especially thick, although it becomes so
with maturity. Additional, often formidable, protection is possibly provided
by the cupule from anthesis onward, not only by the scales but also by the
large amounts of tannins, crystals, and sclereids present.
In all species the cupule provides complete coverage of the immature nut,
102 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
but in many the maturing nut emerges from the cupule, by which time its own
pericarp is very strong.
As in Quercus, effective dissemination of the fruits of Lithocarpus requires
animals (but see Boucher, 1981). Monkeys, squirrels, and similar mammals
are known to be especially important in burying the nuts (Camus, 1952-1954;
pers. obs.), which have hypogeal germination. Some nuts are, of course, eaten
by those animals, but many are buried and not exhumed.
The real or apparent dehiscence of some cupules recalls the more obvious
dehiscence of the cupules of Castanea and Castanopsis. The pattern is regular
in some species of Lithocarpus (e.g., L. encleisacarpa, L. garrettiana) but ir-
regular in others. Correspondence of dehiscence lines to sutures between valves
is unknown for Lithocarpus but is understood for some other fagaceous genera.
Some mature cupules of Lithocarpus bear abortive pistillate flowers at various
sites (see, for example, Ficures 3, 8, 10). Often it is clear that these abortive
flowers are merely other flowers of the cymule that have been elevated some-
what by the overwhelming growth of the cupule of the fertilized flower (FIGURES
8, 10, 35). In other instances such abortive flowers have probably actually
formed upon the cupule itself from latent floral primordia of the ancestral,
now-condensed, cymose branching system that produced the cupule (see FIGURE
3). Fey and Endress (1983) stated that apparently adventitious staminate flow-
ers upon the cupule of Fagus sylvatica L., as reported by Cole (1923), are not
unexpected if each cupular valve is interpreted as a modified branching system.
That concept also seems valid for the presence of pistillate flowers on the upper
regions of mature cupules.
In such a large genus as Lithocarpus, there has undoubtedly been substantial
adaptive radiation, parallelism, and convergence leading to a plethora of pat-
terns of reproductive structure. There is very little published information that
relates reproductive structure in the genus to habitat or pollination specializa-
tions, making interpretations of structure/function relationships difficult.
The homology of staminate with pistillate cymules, as suggested by Kaul
and Kaul (1981), is corroborated by the evidence presented here. Not only do
those cymules have similar bracteole patterns in general, but they also occupy
interchangeable sites in some spikes. In a few staminate cymules, such as those
of Lithocarpus fenestrata and L. pattaniensis, there is a set of bracteoles interior
to the main ones. These are probably additional residual bracteoles of a con-
densed branching system and may represent a rudimentary system of cupular
scales in the staminate cymules, perhaps fully homologous with the cupular
scales of the pistillate cymules. In some cymules the flowers are both staminate
and pistillate, or perfect, or perfect and imperfect (sometimes all of these on a
single spike), indicating that separation of the sexes is not complete at flower
and cymule levels. In Quercus, by contrast, the functional sexes are strictly
separated into different spikes (except in obviously aberrant specimens), but
the pistillate flowers often have well-developed staminodia, especially in the
tropical species (Kaul, 1985). Neither Quercus nor Lithocarpus is dioecious.
1987] KAUL, LITHOCARPUS 103
ACKNOWLEDGMENTS
This study was funded by National Science Foundation grants DEB-7921641
and DEB-8206937. I am indebted to the numerous persons and institutions
cited elsewhere (Kaul & Abbe, 1984) for their assistance in the field and the
laboratory.
LITERATURE CITED
BARNETT, E.C. 1940. A suyey of the genus Quercus and related genera of the Fagaceae
in Asia with a d account as Siamese species of these genera. Unpubl.
D. Sc. thesis, University of Aber
1942. The Fagaceae of Thailand and their geographical distribution. Trans.
Bot. Soc. Edinburgh 33: 327-343.
Keys to the species groups of Quercus, Lithocarpus, and Castanopsis of
eastern Asia, with notes on their distribution. [bid. 34: 159-
Boucner, D. H. 1981. Seed predation by mammals and forest dominance by Quercus
oleoides, a tropical lowland oak. Oecologia 49: 409-414.
Camus, A. 1948. Les chénes. Monographie des genres Quercus et Lithocarpus. Atlas,
vol. 3. Encycl. Econ. Sylvic. 7: 152-165.
1952-1954. Les chénes. /bid. 8: 511-1196.
Cote, L. W. 1923. ae ae phenomena in the inflorescences of Fagus silvatica.
Ann. Bot. (London) 37: 147-150.
Eutas, T.S. 1971. oa genera - Fagaceae in the southeastern United States. J. Arnold
Arbor. 52: 159-19
Fey, B. S., & P. K. ces 1983. Development and morphological interpretation of
the cupule in Fagaceae. Flora 173: 451-468.
Forman, L. L. 1966. On the evolution of cupules in the Fagaceae. Kew Bull. 18: 385-
419
Hyevtmovist, H. 1948. Siucies on the floral morphology and phylogeny of the Amen-
tiferae. Bot. Not. Suppl. 2: 1-171.
KauL, R. B. 1985. Ree cdnctive morphology of Quercus. Amer. J. Bot. 72: 1962-
1977
1986. Evolution and reproductive biology of inflorescences in Lithocarpus,
Castanopsis, Castanea, and Quercus (Fagaceae). Ann. Missouri Bot. Gard. 73: 284—-
296.
& E.C. Aspe. 1984. Inflorescence architecture and evolution in the Fagaceae.
J. Arnold Arbor. 65: 375-401.
& M.N. KAuL. 1981. Homologies between staminate and pistillate inflores-
cences in the Fagaceae. XIII Int. Bot. Congr., Sydney. Abstr. 283
Li, H.-L. 1963. Woody flora of Taiwan. Livingston Publ. Co., Narbeth, Pennsylvania.
Liao, J.-C. 1969. Morphological studies on the flowers and fruits of the genus Litho-
carpus in Taiwan. Mem. Agric., Natl. Taiwan Univ. 10: 1-32.
Lin, W.-F., & T. Liu. 1965. Studies on the classification of Fagaceae in Taiwan. Bull.
Taiwan Forestry Res. Inst. 110: 1-5
Littte, E. L. 1971. Atlas of United States trees. Vol. 1. U.S.D.A. Misc. Publ. 1146.
Govt. Printing Office, Washington, D. C.
Nixon, K. 1985. Cotyledon characters of Mexican white oaks: distribution and phy-
logenetic significance of fused cotyledons. Amer. J. Bot. 72: 964.
Scuotrky, E. 1912. Die Eichen des Paaabete ite Ostasiens und ihre pflanzengeo-
graphische Bedeutung. Bot. Jahrb. Syst. 47: 617-7
104 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
SoePADMO, E. 1968. A revision of the genus Quercus L. subgen. Cyclobalanopsis (Oer-
ste e Schneider in Malesia. Gard. Bull. Singapore 22: 355-427.
970. Florae Malesianae a XLIX. Malesian species of Lithocarpus
BI. anes Reinwardtia 8: 197-
. 1972. Fagaceae. Fl. Males. I. io. 265. 403.
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 105
CONTRIBUTIONS TO A FLORA OF ANGUILLA AND
ADJACENT ISLETS
RICHARD A. HOWARD AND ELIZABETH A. KELLOGG!
The small island of Anguilla is north of St. Martin and with it comprises
the westernmost of the “limestone Caribbees,” separated from the British
Virgin Island group of Anegada, Tortola, Jost Van Dyke, and Virgin Gorda
by the Anegada passage (166 km wide and 604 m deep). A brief checklist of
the vegetation of Anguilla was published by Boldingh (1909), listing 150 species
as his collections, sight records, or literature references; subsequent additions
are few. Pére Le Gallo visited Anguilla in 1955 and 1956 and aeare anew
flora of the island, which was available for our study but has never been
published. Our visit in 1985 produced 125 collections with additional sight
records for a total flora of 443 species. Previously three taxa had been consid-
ered endemic to Anguilla, but all are now known from other islands. Rondeletia
anguillensis is described as new and is considered endemic to Anguilla. Com-
parisons are made with the Virgin Islands to the west of Anguilla.
The small island of Anguilla? lies about 10 km (6 mi) north of St. Martin;
together the two islands form the western extension of the Leeward Island
complex, known as the “limestone Caribbees” (Harris, 1965). Anguilla is sep-
arated from the British Virgin Island group of Anegada, Tortola, Jost Van
Dyke, and Virgin Gorda by the Anegada passage, 166 km (100 mi) wide and
over 604 m (2000 ft) deep. Shoals extend north to Sombrero. Close to Anguilla
are Anguillita, Dog Island, Prickly Pear Cays, Seal Island, Scrub Island, and
Little Scrub Island. Road Bay offers the only large and partially protected harbor
for fishing boats and visiting yachts.
Anguilla is at latitude 18°13'12”N and longitude 65°4'22”W. It is approxi-
mately 28 km (16 mi) long and 8 km (4 mi) wide at its broadest point, with
an area of 90 sq. km (35 sq. mi) (see Map 1). The highest point is Crocus Hill,
with an elevation of 59 m (192 ft). The center of the island is mildly depressed
to form a basin, in which the principal town of The Valley is located. The
island’s population is about 7000. According to Southey (1827), the island of
Anguilla, then called Snake Island, was colonized by the British about 1650
and remained a part of the British Commonwealth. In 1967 Anguilla separated
from the independent state of St. Kitts-Nevis and Barbuda.
‘Arnold Arboretum, Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138.
Wilson, for example, visited the Anguilla Cays but not Anguilla, and his collections have boca cited
incorrectly in Flora Neotropica monographs.
© President and Fellows of Harvard College,
Journal of the Arnold Arboretum 68: 105-131. ae. 1987.
106 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
ISLAND
HARBOUR
SHOAL Bay
AVANNAH Bay
THe FOUNTAIN S
Crocus Bay
OPN
THE VALLEY
Crocus Hitt
AIRPORT
West ENnp
7 a RENDEZVOUS BLowInG Point
q Bay
Map 1. Island of Anguilla.
The geology of the island was most recently studied by Christman (1953).
From the sea, the island appears almost flat (see Ficure 1)—a raised platform
of coralline limestone. The beds of limestone and marl are underlain with
andesite tuffs equivalent to and contemporaneous with the Pointe-Blanche
Formation on St. Martin. Vaughan (1926) has described it as the lowest Mio-
cene type in the Caribbean. Scattered old volcanic boulders are found near
Crocus Bay, near Old Road Bay, and on Dog Island (Le Gallo, unpubl. ms.).
Weathered limestone pavement is evident in many places, devoid of soil cover
and pitted by broad, shallow solution hollows or penetrated by tubular channels
where most plants are rooted. The existing soil on the limestone is terra rossa,
an alkaline, reddish brown clay of low fertility. Elsewhere, a blackish, highly
alkaline clay called rendzina has accumulated in poorly drained depressions.
The limestone forms sea cliffs on the north coast estimated to approach 30 m
(100 ft). Several karstic sinkholes, the most famous being ‘“‘The Fountain,” are
present near Shoal Bay. Uplifted coastal limestone benches are few and rela-
tively low; they were seen only on the south coast. Coastal embayments have
been cut off by sand bars (see FiGure 2) and exist as salt ponds (see FIGURE
3) that are only occasionally activated. Inland lakes are shallow and brackish.
Drinking water is obtained from roof catchments, although a public water
supply from several shallow wells produces mildly brackish water. Average
annual rainfall is 1026 mm, with the peak months being May and August
through November. The figures for average monthly rainfall in mm for the
years 1931-1981 (with data for 1982 in parentheses) are as follows:
January 62 (35) April 64 (29)
February 39 = (173) May 102. (121)
March a7 (32) June 65 (105)
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 107
IGUR View looking west from Shoal Bay hotel development. Note plant of
Scaevola plumieri established on sandy beach (left).
July 79 (89) October 130 =(184)
August 102 (39) November 130 (39)
September 130 (55) December 83 =(108)
VEGETATION
Harris (1965) has termed the vegetation of Anguilla an evergreen woodland.
By Beard’s (1955) classification, it would be called an evergreen bus hland, more
popularly known in the area as thorn scrub. Although Beard believed that the
Anguillan thorn scrub is the natural vegetation of the island, Harris suggested
that it represents a subclimax created by biotic processes of impoverishment
and selection of xerophytic, sclerophyllous species. Harris (1965, p. 137) ad-
mitted, however, that in “Anguilla, the communities of native plants are more
complex, and aliens much less abundant.” He emphasized (p. 137) a “mod-
erately large population dependent mainly on shifting cultivation, together with
considerable development of plantations which resulted in the complete or
partial clearance of the whole island.” Our observations led to a somewhat
different conclusion. It appeared to us that agriculture is at best tenuous on the
shallow soils; the extensive exposed limestone pavement, with plants rooting
in solution holes, indicates that the thorn-scrub vegetation has always been
dominant.
Ona special trip to aid West Indian agriculture, Morris (1891) recommended
that the thorn-scrub areas be cleared as a work-relief project and that such fiber
108 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
FiGure 2. Aerial view of west end of the island near Rendezvous Bay.
sources as Agave and Furcraea be planted, but this was never developed. A
government-sponsored planting of A/oé vera has long been abandoned. His-
torical records show that the cultivation of sugar cane and cotton and the
planting of mahogany were unsuccessful in the low-rainfall climate. Subsistence
agriculture today consists of small home gardens and an ea larger plot
of cassava, pigeon peas, sweet potatoes, okra, and pumpki
Boldingh (1909a), in the only existing list of plants of Anguilla, reported (p.
2) “a vegetation that consisted chiefly of prickly plants resembling in superficial
appearance the Croton vegetation of the Dutch Antilles. I did not see any
tropical wood.” In 1985 a low shrub vegetation dominated most of the un-
cultivated areas and no active charcoal pits were observed, suggesting a paucity
of appropriate charcoal wood or the complete acceptance of kerosene and
electricity for cooking. Cattle were certainly fewer than in the past, and goats
and sheep were mostly tethered in appropriate feeding locations and had little
effect on most of the thorn scrub.
The neem, Azadirachta indica, has been introduced relatively recently and
is perhaps the most common shade tree. Large specimens of Mangifera indica,
Meliococcus bijugatus, Swietenia mahagoni, Tamarindus indica, and Zizy-
phus mauritiana exist around habitations. Occasional trees of Ficus citrifolia,
Guapira fragrans, Pisonia subcordata, and Tabebuia pallida are the largest
native species.
In coastal areas and around salt ponds, the dominant woody plants are
Argusia gnaphalodes, Avicennia germinans, Coccoloba uvifera, Conocarpus
erecta, Erithalis fruticosa, Hippomane mancinella, Laguncularia racemosa,
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 109
iGURE 3. Road Bay harbor looking north, with salt pond development to right.
Canella, Capparis, and Exostema species growing on distant point.
Scaevola plumieri, and Suriana maritima. Very few individuals of Casuarina
equisetifolia, Chrysobalanus icaco, or Thespesia populnea were encountered.
Although Harris (1965) indicated ‘“‘mangrove swamps” at Little Harbour and
Sea Feathers Bay, we found Rhizophora mangle only in Road Bay pond, where
there were a few isolated individuals.
Locally dominant shrubs included Antirhea acutata, Bourreria succulenta,
Byrsonima lucida, Canella winterana, Castela erecta, Coccoloba krugii, Como-
cladia dodonaea, Croton flavens, Eugenia axillaris, E. foetida, Exostema cari-
baeum, Gyminda latifolia, Jacquinia arborea, J. berterii, Malpighia emargi-
nata, Phyllanthus epiphyllanthus, see unguis-cati, Plumeria alba,
Randia aculeata, and Reynosia uncinata. These may be in mixed populations,
and occasionally a single large specimen may dominate an area. Coccoloba
krugii and C. uvifera are known to hybridize on other islands (Howard, 1957).
On Puerto Rico, St. Thomas, St. Croix, and Virgin Gorda the hybrids resembled
C. uvifera. Three distinct plants on Anguilla were called to our attention by
Andrew Parker and shown to us by Oliver Hodge. They were isolated indi-
viduals with leaves more like those of a very large C. krugii. One plant had
been coppiced; its leaves were intermediate in shape but with the texture of C.
krugii and the abundant pubescence of C. uvifera. One plant had fruits com-
parable to those of C. krugii, while the other two had sterile fruits resembling
those of C. uvifera.
ost abundant spiny plants on Anguilla were Acacia macracantha,
Castela erecta, Clerodendrum aculeatum, Comocladia dodonaea, Pithecello-
110 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Comparisons of island size, altitude, and recorded flora.
NUMBER OF SPECIES
REA MAXIMUM ALT. Introduced/ Endemic/
ISLAND (sq. m1) (ft) Indigenous cultivated restricted*
egada 14 15 198 33
Anguilla 35 192 321 122 1 [3]
arbuda 62 47 229 32 {1
Jost Van Dyke 4 1054 73t 12
St. Bartholomew 10 800-1000 336 53 [4]
St. Martin 38 1119, 1391 392 l
To 24 1263, 1780 4844 152 2
Virgin Gorda 8 1539 372 I
*Bracketed numbers represent species now in synonymy.
+Trees only.
tDicotyledons only.
bium unguis-cati, Randia aculeata, Reynosia uncinata, Zanthoxylum flavum,
Z. punctatum, and Z. spinifex. The scramblers Caesalpinia crista and C. diver-
gens, with extremely spiny fruits, may be added to this list. Other scramblers
forming local entanglements include Boerhavia scandens, Cissus verticillatus,
Heteropteris purpureus, Merremia dissecta, Passiflora foetida, P. suberosa,
Plumbago scandens, Rhynchosia minima, R. reticulata, Stigmaphyllon diver-
sifolium, S. emarginatum, S. lingulatum, Tournefortia volubilis, and Urechites
lutea. Parasitic plants were Cassytha filiformis, Cuscuta americana, Dendro-
pemon caribaeus, and Phoradendron trinervium.
Existing floristic studies (Box, 1939; D’Arcy, 1967, 1975; Le Gallo, 1957:
Little, 1969; Little et a/., 1976; Monachino, 1941) of the small northern islands
are not comparable, and significant comparisons are difficult to make (see
TABLE).
The following taxa were originally described as endemic.
ANEGADA: Cynanchum anegadensis (Britton) Alain. Type: Britton & Fishlock
962 (ny). Current status: endemic. Fishlockia anegadensis (Britton) Britton
& Rose. Type: Britton & Fishlock 990 (Ny). Current status: the basionym,
Acacia anegadensis Britton, is preferred. Endemic.
ANGUILLA: Bouteloua vaneedeni Pilger. Type: Boldingh 3521B (s?). Current
Status: now known from St. Bartholomew and from Camaguey province,
Cuba. Myrtus anguillensis Urban. Type: Boldingh 3509B (B?) (= Psidium
longipes (Berg) McVaugh var. orbicularis (Berg) McVaugh). Current status:
now known from the eastern Bahamas, the Turks and Caicos islands, Ja-
maica, St. Bartholomew, Barbuda, and Antigua. Rondeletia anguillensis R.
Howard & E. Kellogg. Type: R. Howard & E. Kellogg 20105 (a). Current
status: endemic. Thrinax morrisii Wendl. Type: H. A. A. Nicholls s.n., 1890
(kK). Current status: known from Florida, Cuba, Haiti, Puerto Rico, the Ba-
hamas, and the Turks and Caicos islands.
BaRBUDA: Coccothrinax boxii Bailey. Type: Box 669 (BH) (= Coccothrinax
barbadensis (Lodd.) Bec). Current status: known from the Lesser Antilles,
Trinidad and Tobago.
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA Vi
St. BARTHOLOMEW: Peperomia barthelemyana Trel. Type: Questel 275 (not
located) (= Peperomia myrtifolia (Vahl) Dietr.). Current status: known from
St. Croix and the Lesser Antilles. Peperomia barthelemyana Trel. var. reducta
Trel. Type: Questel 361 (not located) (= Peperomia myrtifolia (Vahl) Dietr.).
Current status: known from St. Croix and the Lesser Antilles. Peperomia
questeliana Trel. Lectotype: Questel 2518 (Ny) (= Peperomia humilis Dietr.).
Current status: known from Florida, Central America, and the Greater and
Lesser Antilles. Peperomia myrtifolia (Vahl) Dietr. var. major Trel. Type:
Questel 803 (Ny) (= Peperomia myrtifolia (Vahl) Dietr.). Current status:
known from St. Croix and the Lesser Antilles.
St. Martin: Calyptranthes boldinghii Urban. Type: published as Boldingh
2370B (B?) but 3270B on label. Current status: endemic and known only
from the type collection.
Tortoia: Calyptranthes kiaerskovii Krug & Urban. Type: Eggers 3217 (B?).
Current status: original material sterile and identification uncertain; now also
reported from Virgin Gorda. Sida eggersii E. G. Baker. Type: Eggers 31 &3
(3m?, K?). Current status: known only from a single tree on Jost Van Dyke.
D’Arcy (1967) reported the species from Tortola and Culebra but did not
encounter it.
VirGIN Gorpa: Croton fishlockii Britton. Type: Fishlock 31] (ny). Current
status: endemic.
It can be estimated that the floras of the “limestone Caribbees”’ and adjacent
islands each consist of about 500 species. With the few exceptions of species
whose distribution is limited to adjacent islands, the species that dominate the
vegetation of any island can also be found on Puerto Rico and the drier areas
of Hispaniola, occasionally Cuba, and to a lesser extent Guadeloupe. The
islands with peaks of 1000 feet or more are likely to have a rain shadow that
affects the island and niches where zonation of the vegetation can be established.
The lower islands of Barbuda, Anguilla, and Anegada are more apt to receive
fortuitous rain showers. Barbuda and Anegada have been more extensively
cultivated or grazed in the past, and a larger percentage of the existing vegetation
is adventive and secondary. Anguilla stands out in the amount of limestone
avement area having what we concluded to be a natural and less disturbed
vegetational type.
BOTANISTS WHO HAVE VISITED ANGUILLA
L.-C. RICHARD, 1786. Urban (1902) reported that Richard had collected on
Anguilla during his voyage north from Cayenne in the spring of 1867. We have
seen no collections or citations of such specimens. Box (1939) located citations
for four type specimens from Antigua and for one specimen from Barbuda.
D. Morris, 1890. Morris visited the Lesser Antilles as an agricultural con-
sultant in 1890 and gathered 30 to 40 living plants ofa dwarf palm, later named
Thrinax morrisii by Wendland, during a visit to Anguilla on December 14
and 15.
AZ JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
H. A. A. NicHoLis, 1891. Nicholls, a medical doctor and specialist on yaws,
was on Barbuda in August, 1891, and wrote on November 11, 1891 (in Wend-
land, 1892), “I went again to Anguilla.” His collections, sent to Kew, are
unnumbered. They have not been encountered except for fruiting specimens
of Thrinax morrisii, one of which is now the lectotype, that he gathered to
permit the description to be completed and published.
W.R. Exuiott, 1891. Elliott had been a gardener on Jamaica and later Grenada.
The reason for his trip to Anguilla (1891) is not known, but his small collection
of 34 numbers was identified by Box (1940).
I. BoLpINGH, 1906. Boldingh was preparing a report of the vegetation of the
Dutch Antilles (1909b, 1913) and visited Anguilla on September 6 and 7, 1906.
His publication (1909a) remains the only paper on the flora of Anguilla and
is based on his collections numbered between 3449 and 3599.
G. G. Goopwin, 1926. Goodwin, accompanied by his wife, visited Anguilla
between April | and 9, 1926, as part of the Ottley Puerto Rican expedition of
the American Museum of Natural History, in search of recent and fossil mam-
mals. A single specimen numbered /2 was found in the herbarium of the New
York Botanical Garden. Botanical collections are not mentioned in the catalogs
and journals of the expedition.
P. WAGENAAR HUMMELINCK, 1949, 1973. Hummelinck visited Anguilla and
Dog Island June 16-20, 1949, and June 30-July 3, 1973. He collected two
species of Agave, as well as algal and faunal specimens. The algal collections
are listed in Vorman (1968). Hummelinck (1981) also published observations
on “land and fresh-water localities,” with photographs of Anguilla.
I. VeLEz, 1950. Prior to the publication of his Herbaceous Angiosperms of the
Lesser Antilles in 1957, Velez spent fiscal year 1949-1950 collecting between
the Virgin Islands and Grenada. He reported (p. 2) Anguilla to be among the
islands that “‘were thoroughly studied.” A single specimen, Velez 3749 (us)
(Thrinax morrisii), collected in January, 1950, was reported in the literature
encountered. Velez (1957, p. 2) reported that ‘“‘a complete set was deposited
in the Herbarium of the Inter American University of Puerto Rico. Duplicates
of most of them were sent to the Herbarium of the Imperial College of Tropical
Agriculture, Trinidad.” Velez’s citations of species distribution were taken from
the literature, were sight records, or were supported by specimens. They have
been troublesome to untangle. On a visit to the Inter American University,
one of us (R. A. H.) discovered that his herbarium, through neglect, had been
completely destroyed by insects. The set sent to Trinidad was later given to
Kew, where we have seen specimens from other islands. Lists of determinations,
preserved at Kew, are not complete but cite specimens numbered 3006 to 3/58
from the Virgin Islands, 3759 to 3287 from Grenada and the Grenadines, 3290
to 3337 from St. Lucia, and 3338 to 3386 from St. Vincent. A few specimens
have been found in Gu, ny, and us, but nothing from Anguilla.
C. Le GALLO, 1955, 1956. Le Gallo collected on Anguilla September 1-5, 1955,
and on the adjacent islets of Scrub and Dog March 3, 1956. His unpublished
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 113
and undated manuscript must have been written sometime in 1959. His plant
specimens as cited are numbered 2053 to 2071 and 2470 to 2521. Le Gallo is
not listed in Barnhart (1965), so it is of interest to record here the biographical
information we obtained first from Ms. Céline Arseneault, botanist-librarian
of the Montreal Botanic Garden, and subsequently from the tribute to Le Gallo
by Pére Maurice Barbotin (1976), of Guadeloupe.’
G. R. Proctor, 1958-1959. Proctor collected extensively in the Leeward Is-
lands; between December 30, 1958, and January 18, 1959, he gathered 250
numbers, 1/8518 to 18704 and 18731 to 18816, on Anguilla. Complete sets of
his specimens are at A and JJ.
D. R. Harris, 1960. Harris spent part of August, 1960, on Anguilla prior to
publication of his “Plants, Animals, and Man in the Outer Leeward Islands,
est Indies” in 1965. His collections of about 50 species were given to the
British Museum (Natural History). We have included all the species in our
listing, but since we have not seen specimens, they are not cite
R. W. Reap, 1974. Read, of the Smithsonian Institution, visited Anguilla on
June 7, 1974, to find and collect Thrinax morrisii. He made no other collections
there (pers. comm.).
R. A. Howarp AND E. A. KELLOGG, 1985. We collected on Anguilla February
5-9, 1985. Our specimens numbered 20043 to 20168 are deposited in the
herbarium of the Arnold Arboretum (A).
ACKNOWLEDGMENTS
Weare grateful for the help of Oliver Hodge, of the Public Health Department
on Anguilla, who accompanied us in the field and supplied local names and
uses of the plants we encountered. Andrew Parker, of Powys, Great Britain,
was recently stationed on Anguilla and called our attention to the hybrid pop-
‘Pére Casimir Le Gallo was born June 25, 1906, at Erdeven, diocése of Vannes, France. He took
his holy orders in the Congregation of St.-Esprit October 1, 1933. He was a professor at Collége St.-
Alexandre, Touraine, Quebec, from August 30, 1934, to January, 1935. He served as Vicar Apostolic
Seeking a warmer clime, he went to the West Indies in 1951 as curate of St. B Bartholomew, where he
also taught natural science at the seminary-college of Blanchet. In 1958 he f Vieux
Fort, although he continued weekly teaching at Blanchet. Eight years later the bishop me him
with several successive assignments in Sacré-Coeur, in Fatima, at the oule, and finally in Baie-
assignment in St. Pierre and Miquelon. He was not to occupy this position, however, for an injury
to his foot developed into gangrene and his leg was amputated. He spent = and painful months
in the hospital before his residence in the religious community at Wolxhe . He yearned for his
His primary botanical interest was in mosses, but he collected algae and lichens as weil as vascular
plants. Twenty-one papers by Le Gallo appeared in Le Naturaliste Canadien between 1945 and 1965,
priests who were botanists. Proctor collected with Le Gallo in Guadeloupe in 1959 and as a result
of that trip named Diplazium legalloi in his honor
114 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
ulations of Coccoloba; subsequently in correspondence he supplied additional
information we have incorporated. His interest in the local vegetation may
lead to an illustrated ethnobotanical publication. George R. Proctor, who has
collaborated with us in the production of the Flora of the Lesser Antilles,
supplied lists of his collections from Anguilla. Pére Le Gallo gave Proctor his
unpublished manuscript notes on Anguilla, which we have been privileged to
use. Céline Arseneault, of the Montreal Botanical Garden, and Jacques Porte-
cop, Université des Antilles, Guadeloupe, located biographical information on
Le Gallo for us. Finally, our travel and work on this flora was made possible
through the support of National Science Foundation grant BSR-8307701 and
a grant from the Atkins Fund of Harvard University, for which we are appre-
ciative.
LITERATURE CITED
BarsoTin, M. 1976. Le Pére Casimir Le Gallo, 25 Juin ten Juin 1976. Leafl.
Eglise Guad. 211: 8-11. Edit. Evéché Basse-Te erre, Gua pe.
BARNHART, J. H. 1965. Biographical notes upon eae cE 1-3. G. K. Hall &
Co., Boston.
BEARD, J. S. 1955. The classification of tropical American vegetation types. Ecology
36: 89-100.
eae I. 1909a. A yas to the knowledge of the flora of Anguilla. Recueil
v. Bot. Néerl. 6: 1-34.
1909. The ee ae Dutch West Indian islands of St. Eustatius, Saba, and
St. Martin. E. J. Brill,
1913. Flora vOOr “ Fe detains West-Indische eilanden. J. H. De Bussy,
Amsterdam.
Box, H. E. 1939. Flora of Antigua and Barbuda. Unpublished manuscript (British
Museum (Natural History)).
1940. Report upon a collection of plants from Anguilla, B.W.I. J. Bot. 78:
14— 16.
CHRISTMAN, R. A. 1953. Geology of Saint-Bartholomew, Saint-Martin and Anguilla,
Lesser ae Bull. Geol. Soc. Amer. 64: 65-93.
D'Arcy, W. G. 1967. Annotated check-list of the dicotyledons of Tortola, Virgin
Te Rhodora 69: 385-450.
1975. Anegada Island: vegetation and flora. Atoll Res. Bull. 188: 1-40.
Ev_utiotr, W. R. 1891. Botanical enterprises in the West Indies. Kew Bull. 1891: 103—
168
GouLp, F. W. 1979. Poaceae. Pp. 25-220 inR. A. pata oo Lesser Antilles.
Harris, D. R. 1965. Plants, animals, and man in the ee Leeward Islands. West
Indies. Univ. Calif. Publ. Geogr. 18: 1-164.
Howarp, R. A. 1957. Studies in the genus Coccoloba, IV. The species from Puerto
Rico and the Virgin Islands and from the Bahama Islands. J. Arnold Arbor. 38:
211-242.
HUMMELINCK, P. W. 1981. Studies on the fauna of Curacao and other Caribbean islands.
Publ. Found. Sci. Res. Surinam & Netherlands Antilles
Le GALLo, C. 1957. Myrtus orbicularis nee Burret, endemuaue des Petites Antilles
du nord. Bull. Soc. Bot. France 104: 158-
. Contribution a la florule d’Anguilla. Cea bien manuscript (Arnold Arbore-
tum
LITTLE, E. L., JR. 1969. Trees of Jost Van Dyke (British Virgin Islands). U. S. Forest
Serv. Res. Paper 1TF-9.
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 115
, R. O. Woopsury, & F. H. WApsworTH. 1976. Flora of Virgin Gorda (British
Virgin Islands). U. S. Forest Serv. Res. Paper 1TF-21.
Monacuino, J. 1941. A check-list of the spermatophytes of St. Bartholomew. Carib-
bean Forest. 2: 25-47, 49-66.
Morris, D. 1891. Report of a botanical mission to the West Indies. Kew Bull. 1891:
109-162
QuesteL, A. 1941. The flora of the island of St. Bartholomew. Imprimerie Catholique,
Guadeloupe.
SouTHEY, T. 1827. Chronological history of the West Indies. Longman, Rees, Orme,
Brown, & Green, London
Ursan, I. 1902. Symbolae Antillanae. Vol. 3. Borntraeger, Leipzi
VAUGHAN, T. W. 1926. Notes on the igneous rocks on the ee West Indies and
on the geology of the island Anguilla. J. Wash. Acad. Sci. 16: 345-358.
VeLez, I. 1957. Herbaceous angiosperms of the Lesser Antilles. Inter American Univ.,
Puerto Rico.
VorMAN, M. 1968. The marine algal vegetation of St. Martin, St. Eustatius and Saba.
Publ. Found. Sci. Res. Surinam & Netherlands Antilles
WENDLAND, H. 1892. Thrinax morrisii Wendl. Gard. Chron. TL. 11: 104.
APPENDIX. The known flora of Anguilla.
The collections cited sais ferred to by the ee abbreviations: B = Boldingh,
E = Elliott, GG = Goodwin, H = Hummelinck, HK ward and Kellogg, LG = Le
Gallo, P = Proctor. ee to Le Gallo’s anes a > Boldingh collections are at
Utrecht and were at Berlin. They were also sought in Ny, but very few of the cited
numbers could be found. Le Gallo’s (unpubl. ms.) collections were studied by Monachino
(ny) and by Miss G. J. A. Amshoff. Again, a search at ny located very few of the numbered
collections. Those seen, as well as those of Proctor and our own, are indicated by the
herbarium acronym, most frequently a. Common names are included only when they
were provided by local residents.
GYMNOSPERMAE
ARAUCARIACEAE
Araucaria heterophylla (Salisb.) Franco, Christmas plant. Cultivated. HK sight.
ANGIOSPERMAE
MoONOCOTYLEDONEAE
AGAVACEAE
Agave beauleriana Jacobi. Cultivated. HK —
Agave karatto Miller. Cultivated. HK sig
Agave scheuermaniana Trel. H 160, 161,
Agave sisalina Perrine, fiber pole, pita, a plant. Cultivated. HK sight; H ///, 112.
Sansevieria hyacinthoides (L.) Druce. Naturalized. HK sight; P 18760 (a).
sight.
ucca guatemalensis Baker, Spanish needle. Cultivated. HK sight.
AMARYLLIDACEAE
Crinum sp. Cultivated. HK sight.
Hymenocallis caribaea (L.) Herbert, spider lily. P 18630 (a).
Zephyranthes candida (Lindley) Herbert, crocus, snowdrop. Parker sight.
116 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
ARACEAE
Alocasia plumbea C. Koch. Cultivated. HK sight.
Colocasia esculenta (L.) Schott. Cultivated. HK sight.
Dieffenbachia seguine (Jacq.) Schott. Cultivated. AK sight.
Monstera acuminata C. Koch. Cultivated. HK sight.
Rhaphidophora aurea (Linden & André) Bi a Cultivated. HK sight.
Xanthosoma sagittatifolium (L.) Schott. Cultivated. HK s ight.
ASPARAGACEAE
Asparagus setaceus (Kunth) Jessop. Cultivated. HK sight.
Asparagus sprengeri Regel. Cultivated. HK sight.
BROMELIACEAE
Tillandsia recurvata L., wild pine. B s.n., P 18566 (A).
Tillandsia usneoides L., jumbie beds. Parker sight.
Tillandsia utriculata L., wild pine. P 18634 (a).
COMMELINACEAE
Aploleia monandra (Sw.) Moore. Cultivated. HK s ight.
Callisia fragrans (Lindley) Woodson. Cultivated. HK 20108 (A).
Commelina elegans Kunth. P 18752 (a).
Rhoeo spathacea (Sw.) Stearn. HK sight.
Tradescantia pallida (Rose) Hunt. Cultivated. HK sight.
CYMODOCEACEAE
Syringodium filiforme Kiitz. P 18626 (A).
CYPERACEAE
apes pauciflora (Liebm. : ae LG 2485; P 18804 (a).
Cyperus calcicola Britton. LG
Cyperus laevigatus L. P 187 a
Cyperus oxylepis Nees ex Steudel. P 18749 (a).
Cyperus rotundatus L. B s.n.; P 18772 (a).
Eleocharis geniculata (L.) Roemer & Schultes. P 18771 (A).
Eleocharis mutata (L.) Roemer & Schultes. P 18769 (A).
Fimbristylis cymosa R. Br. subsp. spathacea (Roth) T. Koyama. B 3527, as Fimbristylis
thacea Roth; HK 20151 (a); P 19659 (a
Fimbristylis ferruginea (L.) Vahl, pond grass. B 3495,
Fimbristylis ovata (Burman f.) Kern. B 3514, 3573, both as Fimbristylis monostachya
Hassk.; P 18666 (A
Mariscus brunneus (Sw.) Clarke. LG 2484, as Cyperus planifolius Rich. var. brunneus
w.) Kiik.; P 18690 (a), ae Os ).
Mariscus capillaris (Sw.) Vahl
Mariscus fulgineus (Chapman) rs ake LG 2512 (Ny), as Cyperus fulgineus Chapman; P
He (A), 18635 (A).
cus Squarrosus (L.) Clarke. P 18810 (a).
Sc feria lithosperma (L.) Sw. LG 2503; P 18803 (a).
GRAMINEAE
Aristida adscensionis L. LG 2488, 2489, 2509: P 18699 (A).
Bothriochloa ischaemum (L.) Keng. P 18784 (A).
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 117
Bothriochloa pertusa(L.) A. Camus. P 18784 (a), 18791 (A), both as Andropogon pertusus
(L.) Willd.
Bee americana (L.) Scribner. B 3533; P 18757 (A).
Bouteloua vaneedeni Pilger. B 3512 (type collection); LG 2474.
Brachiaria adspersa (Trin.) Parodi. P 18643, as Panicum adspersum Trin.
Brachiaria fasciculata (Sw.) S. T. Blake. Reported by Gould (1979).
Brachiaria reptans (L.) Gardner & Hubb. B 3543, as Panicum reptans L.
Cenchrus echinatus L., burr grass. P 18762 (A).
Cenchrus incertus M. Curtis. HK 20155 (a); P 18693 (a), as Cenchrus gracillimus.
Cenchrus tribuloides L. B s.n.
Chloris gayana Kunth, Rhodes grass. Reported by Harris as cultivated.
Chloris inflata Link. P 18621 (A)
Cymbopogon citratus (DC. ex Nees) Stapf, lemon grass. Cultivated and naturalized.
B 3454, as Andropogon schoenanthus.
Dactyloctenium aegyptium (L.) Beauv. P 18761 (A).
Digitaria bicornis (Lam.) Roemer & Schultes. P 18970 (A).
Digitaria decumbens Stent, pangola grass. Reported by Harris as cultivated.
Digitaria insularis (L.) Mez. LG sight (Dog Is.), as Trichane insularis (L.) Nees.
Digitaria sanguinalis (L.) Scop. B 3456; P 18790 (a).
Eleusine indica (L.) Gaertner. B s.n.; A).
Eragrostis ciliaris (L.) Link. [7K 20119 (A); LG 2518; P 18746 (A).
Eragrostis tenella (L.) Beauv. ex Roemer & Schultes. HK sight.
Heteropogon contortus (L.) Beauv. LG 2521.
Oplismenus hirtellus (L.) Beauv. subsp. setarius (Lam.) Mez. B s.n., as Oplismenus se-
tarius (Lam.) Roemer & Schultes.
Panicum diffusum Sw. B 3457, 3459, 3538; P 18684 (a).
Panicum geminatum Forsskal. B 3494.
Panicum maximum Jacq. P 18620 (A).
Panicum molle Sw. B 3453
Panicum paniculatum L. B 3539, 3550
Paspalidium geminatum (Forsskal) Stapf. B 3494, as Panicum geminatum Forsskal.
Paspalum fimbriatum Kunth. B 3455; P 18618 (A
Paspalum laxum Lam. B 3550; LG 2516; P 18652 (A), 18767 oA
Paspalum paniculatum L. B s.n., as Paspalum hemisphericum Poi
Rhynchelytrum repens (Willd.) C. E. Hubb., red-headed grass. LG cat as Tricholaena
rosea Nees; P 18789 (A)
Saccharum officinarum L. Cultivated. HK sight.
Sorghum halepense (L.) Pers. Cultivated. HK sight.
pee indicus (L.) R. Br. LG 2519, pro ae
Sporobolus jacquemontii Kunth. LG 2519, pro
Sporobolus pyramidatus (Lam.) A. Hitche. HK 20126 (A); LG 2519, pro parte.
Vetiveria zizanioides (L.) Nash. B te ek sight.
Zea mays L., corn. Cultivated. HK s
HyYDROCHARITACEAE
Thalassia testudinum Banks & Sol. ex Konig. P 18627 (A).
LILIACEAE
Aloé vera (L.) Burman, aloe, sempervive. Cultivated and naturalized. HK sight.
MUSACEAE
Musa sapientum L. Cultivated. HK sight.
118 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
ORCHIDACEAE
Epidendrum kraenzlinii Bello. HK 20149 (a), LG sight, as Epidendrum bifidum Aublet.
Tetramicra canaliculata (Aublet) Urban. LG sight.
PALMAE
Coccothrinax barbadensis (Lodd. ex ie Becc. Cultivated. HK sight.
Cocos nucifera L. Cultivated. HK sight
Phoenix dactylifera L. Cultivated. HK § sight.
Thrinax morrisii Wendl., broom palm, thatch palm. HK 20150 (a); H. A. A. Nicholls
s.n. (lectotype, kK); P 18667 (a).
PANDANACEAE
Pandanus utilis Bory. Cultivated. HK sight.
RUPPIACEAE
Ruppia maritima L. HK 20130 (a); LG sight (Scrub Is.); P 18773 (a).
DicoTYLEDONEAE
ACANTHACEAE
Asystasia gangetica (L.) T. Anderson. Cultivated. HK sight.
Blechum brownei L. P 18531 (A).
Oplonia spinosa (Jacq.) Raf. LG sight.
Pseuderanthemum carruthersii (Seemann) Guillaumin var. reticulatum (Bull) Fosb. Cul-
tivated. HK sight.
Ruellia tuberosa L., snagdragon. P 18598 (a).
Thunbergia fragrans Roxb. P 18758 (A).
AIZOACEAE
Sesuvium microphyllum Willd. P 18656 (Aa
Sesuvium portulacastrum L., pondweed. B 3526a.
AMARANTHACEAE
Achyranthes aspera L. B s.n., as Achyranthes obtusifolia Lam.; P toe (A).
Saris ae (L.) Kuntze. Cultivated and natur alized. HK s
rnanthera caracasana Kunth, yard-pussley. B 3555 (Ny), as pean repens
Kuntze: F ? TS754 (A
Amaranthus crassipes Schidl. P 18682 (a).
Celosia nitida Vahl. P 18732 (a).
Lithophila muscoides Sw. HK 20125 (a); LG sight (Dog Is.); P 18687 (A).
ANACARDIACEAE
Anacardium occidentale L., cashew. HK sight.
Comocladia dodonaea (L.) Urban, arn wood, wild mango. B 3556, as Comocladia
ilicifolia Sw.,; LG sight (Dog Is., Scrub Is.); P 18648 (a).
Mangifera indica L., mango. Cultivated. HK sight.
Spondias mombin L. , golden apple, plum. Cultivated. Parker sight.
Spondias purpurea L., fig, hog plum. Cultivated. Parker sight.
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 119
ANNONACEAE
Annona muricata L., soursop. Cultivated. HK sight.
Annona squamosa L., sugar apple. Cultivated. HK sight.
APOCYNACEAE
Catharanthus roseus (L.) G. Don, old maid. ee a
Nerium oleander L., oleander. Cultivated. HK s
Plumeria alba L. , pigeonwood, snakewood. LG nae (Dog Is., Scrub Is.); P 18625 (A).
Plumeria rubra ie , frangipani. Cultivated. HK sight.
Rauvolfia viridis Roemer & Schultes, Antigua balsam. P 18523 (A).
Tabernaemontana divaricata (L.) R. Br. Cultivated. HK sight.
Urechites lutea (L.) Britton, lice bush. B s.n., as Urechites suberecta Muell. Arg.;
HK 20071 (a); LG 2056 (Scrub Is.), sight (Dog Is.); P 186/J (A).
ARALIACEAE
Polyscias fruticosa (L.) Harms. Cultivated. HK s
Polyscias guilfoylei (Cogn. & Marchal) L. H. ae Cine HK sight.
ASCLEPIADACEAE
Asclepias curassavica L. E 47.
Calotropis procera (Aiton) R. Br., French cotton, milky-milky bush. B s.n.; P 18623 (A).
Cynanchum parviflorum Sw. P 18550 (a).
BATACEAE
Batis maritima L., pondweed. B 3545a; LG sight (Scrub Is.); P 18744 (A).
BIGNONIACEAE
Crescentia cujete L. Cultivated. HK s
Podranea ricasoliana (Tanf.) Sees Cc aaa: HK sight.
Spathodea nilotica Seemann. Cultivated. HK si
ates heterophylla (DC.) Britton, bark, cedar, white cedar. B 3482, 3512, 3541, all
ecoma leucoxylon Martius; GG 12 (Ny); HK 20104 (a); P 18555 (a).
ee pallida (Lindley) Miers. Cultivated. Parker sight.
Tecoma stans (L.) Kunth, fever bush, torchwood. E 42; P 18587 (a).
BOMBACACEAE
Ceiba pentandra (L.) Gaertner. Cultivated or naturalized. P 18792 (A).
BORAGINACEAE
Argusia Fane (L.) Heine, wild lavender. B s.n., as Tournefortia gnaphalodes
R. Br.; E 52; LG sight (Scrub Is.); P 78675 (A).
Poe hee Jacq., chink bush. B 35/8 (Ny); LG sight (Dog Is.); HK 20067 (a);
P 18579 (A).
Cordia collococca L., clamen cherry. = 18753 (A).
Cordia sebestena L. Cultivated. HK sight.
Heliotropium angiospermum ie eyebright. E 45; B s.n.; HK 20079 (a); P 18640
I mL.
Heliotropium curassavicum L. LG sight (Dog Is.); P 18657 (A).
Heliotropium indicum L., eyebright. Parker sight.
120 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
pene ees a Sw. B 3517 (Ny); HK 20137 (a); LG 2061, 2062 (both Scrub
Is.); P 18688 (A), 18808 (A).
Ro es acanthophora (DC.) Griseb. LG sight (Scrub Is.).
Tournefortia volubilis L. B 3521, 3540; P 18780 (a).
BURSERACEAE
Bursera simaruba Sarg., turpentine tree. B s.n.; P 18532 (a).
CACTACEAE
Cephalocereus nobilis (Haw.) Britton & Rose, doodle doo. P 18743 (a).
Epiphyllum oxypetalum (DC.) Haw. Cultivated. HK sight.
Hylocereus undatus (Haw.) Britton & Rose. Cultivated. HK sight.
ee nivosa Link. LG sight (Scrub Is.), as Neomammillaria nivosa (L.) Britton
& Ros
oa ee intortus (Miller) Urban, pope’s head. HK sight; LG sight (Scrub Is.), as Cereus
intortus Miller.
Opuntia cochenillifera (L.) Miller. Cultivated. HK sight.
Opuntia dillenii (Ker Gawler) Haw. LG sight (Scrub Is.); P 18783 (A).
Opuntia rubescens Salm-Dyck. LG sight (Scrub Is.).
Opuntia triacantha (Willd.) Sweet. LG sight (Scrub Is.).
CANELLACEAE
Canella winterana (L.) Gaertner, pepper cinnamint, pepper cinnamom. B 3479, as Ca-
nella alba Murray; LG sight (Scrub Is.); P 18558 (A).
CAPPARACEAE
Gi ete cynophallophora L., ey widdy, parrotbush, snake bush. B 3522; HK 20141
(A); LG sight (Dog Is.); P 18733 (A).
Capparis. SO ne L. LG sight (Dog Is.); P 18586 (a).
Capparis frondosa Jacq., whitescrub. B s.n. (Ny), as Capparis baducca L.
Capparis hastata Jacq. HK 20142 (a).
Cleome gynandra L. B s.n., E 59; HK 20110 (A).
CARICACEAE
Carica papaya L., pawpaw. Cultivated. HK sight.
CASUARINACEAE
Casuarina equisetifolia J. R. & G. Forster, lumber tree. Cultivated and naturalized. HK
CELASTRACEAE
C ee cel rhacoma Crantz, maidenberry. B 3489, 3500, both as Rhacoma crosso-
€ L.; E 56; HK 20045 (aA), 20065 (A); LG sight (Scrub Is.); P 18559 (a).
blacodendrn xylocarpum (Vent.) A. DC., cuttard. LG sight (Dog Is.); P 18534 (a),
Gmina aio (Sw.) Urban. B 3479; HK 20043 (a), 20156 (A), 20/68 (A); P 18664
A), 1
ie ae (Lam.) Krug & Urban. LG sight.
Schaefferia frutescens Jacq. P 18633 (A).
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 121
CHENOPODIACEAE
Atriplex ee (Jacq.) Standley. LG 2717 (Dog Is.).
Chenopodium murale L. HK 20131 (a).
Salicornia bigelovi Torrey. HK 20118 (A).
Salicornia herbacea L. B 3505a, 3571.
CHR YSOBALANACEAE
Chrysobalanus icaco L., coco plum. P 18778 (A).
COMBRETACEAE
Conocarpus erecta L., buttonwood, pond bush. B 3545; LG sight (Dog Is.); P 18568 (A).
Laguncularia racemosa (L.) Gaertner. B 3547; P 18745 (A).
Terminalia catappa L., almond. HK sight.
COMPOSITAE
Ambrosia hispida Pursh. Cultivated. 7 sight.
Bidens cyanapiifolia Kunth. P 18679
Borrichia arborescens (L.) DC. E 50, $5: ‘LG sight (Dog Is., Scrub Is.); P 18628 (A).
Cosmos sulphureus Cav. Cultivated. HK sight.
Dyssodia tenuifolia Cass. Naturalized. P 18578 (A).
Emilia peeren Nicolson. P 18742 (A
Eupatorium odoratum L. P 18588 (a).
Bea bidentata (L.) Kuntze. HK sight
Lactuca intybacea Jacq. HK 20114 (a); P 18787 (A).
Lagascea mollis Cav., catnip. B s.n.; P 18525 (a
Parthenium hysterophorus L., mule weed, whitehead, whitetop. B s.n.; E 30; P 18765
(A
Pectis humifusa Sw. B ee P 18660 (A).
Pectis linifolia L. P 18779
Pluchea symphytifolia (Miller) Gillis. B 3563, as Pluchea ee = Cass.; HK sight.
Pseudogynoxis confusus (Greenman) Cabrera. Cultivated. HK s
Solidago microglossa DC. Cultivated. P 18793 (A
Sonchus oleraceus L., sowthistle. HK 20140 (a); P 18629 (A), 18788 (A).
Synedrella nodiflora (L.) Gaertner. P 18674 (a).
Tridax procumbens L. HK sight.
Vernonia albicaulis Pers. HK 20164 (a); P 18543 (A).
Vernonia cinerea (L.) Less. HK 20161 (A); P 18641 (A).
Wedelia aay Rich., marigold. B s.n., as Wedelia buphthalmoides Griseb.; HK 20124
(A); P 7 (A).
Wedelia ee (L.) Hitche. Cultivated. HK sight.
Xanthium strumarium L. HK sight.
Zinnia multiflora L. Cultivated. HK sight.
CONVOLVULACEAE
Cuscuta americana L., dodder, love vine, yellow dod. B 3480; E 31; HK 20100 (a);
0
I (A).
Evolvulus antillanus Powell. B 3565, LG 2475, 2476, 2478, 2504, 2505, allas Evolvulus
argyreus Chois
Evolvulus convolvuloides (Willd.) Stearn. P 18686 (a).
Evolvulus glaber Sprengel. B 3564 (Ny); P 18686 (A).
Evolvulus sericeus Sw. P 18647 (a
ee JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Ipomoea batatas L., sweet potato. Cultivated. HK sight.
pomoea carnea Jaca. subsp. fistu/osa (Martius) Austin, morning glory. Cultivated. HK
sight.
Ipomoea eggersii (House) Austin, wild potato. B 3471, 351 5bis, both as Ipomoea arenaria
(Choisy) Steudel; & 35; HK 2006] (a); P 18542 (a).
Ipomoea nil (L.) Roth. HK sight.
Ipomoea pes-caprae (L.) R. Br. subsp. brasiliensis (L.) Ooststr., sea bean. B s.n.
Ipomoea triloba L. P 18520 (A).
Jacquemontia cayensis Britton. HK oe (A); P 18646 i) 18802 (a).
Jacquemontia pentantha (Jacq.) G. Don, black wiss
Jacquemontia solanifolia (L.) Hallier f "HK 20098 (a): : pie (A).
Merremia dissecta (Jacq.) Hallier f., nio, noyeaux, sprain bush. B s.n.; P 18590 (a).
CRASSULACEAE
Bryophyllum pinnatum (Lam.) Kurz, Christmas plant. B s.n.; HK sight.
Kalanchoé blossfeldiana Poelln. Cultivated. HK sight.
Kalanchoé tubiflora (Harvey) Raym.-Hamet. Cultivated and naturalized. HK 2005] (a).
CRUCIFERAE
Brassica carinata A. Braun. HK 20112 (a).
Brassica oleracea L. var. — L., cauliflower. oa i
Brassica oleracea L. var. capitata L., cabbage.
Cakile lanceolata (Willd.) O. Schulz. HK 20122 a a or (A).
Lepidium virginicum L. B s.n. (NY).
CUCURBITACEAE
Cucumis anguria L. HK 20075 (a).
Cucurbita moschata Duchesne ex Poiret, et Cultivated. HK sight.
Momordica charantia L., maiden apple. HK s
EUPHORBIACEAE
Acalypha amentacea Roxb. subsp. wilkesiana (Muell. ar Fosb. Cultivated. HK sight.
Acalypha chamaedrifolia (Lam.) Muell. Arg. HK 20127 (a).
Acalypha poiretii Sprengel. B 3451; P 18797 (a).
Argythamnia candicans Sw., tea. B 3466; HK 20167 (a); LG sight (Scrub Is.); P 18565
A).
Breynia disticha J. R. & G. Forster. Cultivated. HK s
Chamaesyce blodgettii (Engelm.) Small. HK 20058 i : 18651 (A), 18658 (a).
Chamaesyce hirta (L.) Millsp. HK 20080 (a); P 18576 (a).
Chamaesyce hypericifolia (L.) Millsp. P 18639 (a
Chamaesyce mesembrianthifolia (Jacq.) Dugand. ‘B 3561, 3567, both as Chamaesyce
buxifolia (Lam.) Small; E 48; HK 20121 (a); LG sight (Dog Is.); P 18809 (a).
Chamaesyce multinodis ie Millsp. P 1859/ (a).
lige pilulifera L. B s
Chamaesyce prostrata Aiton. LG sight (Scrub Is.); P 18597 (a).
nae variegatum (L.) Blume. Cultivated. HK sight.
Croton betulinus Vahl, nanny bunch. B 3465, 3499; HK 20159 (a); LG sight (Scrub Is.);
P 18553 (a).
Croton flavens L., balsam. B 3477, 3528, 3599; E 39; HK 20085 (a); LG sight (Dog Is.,
Scrub Is.); P 18522 (A), 18532 (A).
Croton lobatus L. B s.n.; HK 20147 (a); P 18636 (A).
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 123
Croton microcarpus Ham., sweet marjoram. B s.n., as Croton ovalifolius Vahl; HK 20109
(A); LG 2471, 2489, 2506, P 18665 (A), as Croton nummulariaefolius A. Rich.
Euphorbia cyathophora Murray. HK 20165 (a); P 18583 (a).
Euphorbia heterophylla L., Bethlehem star. P 18637 (A)
Euphorbia lactea Haw. Cultivated. HK sight.
Euphorbia leucocephala Lotsy. Cultivated. HK s
Euphorbia pulcherrima Willd., Christmas plant. Cavate. HK sight.
Euphorbia tirucalli L. Cultivated and naturalized. HK sight.
Gymnanthes lucida Sw., scrub bush. P 18805 (a).
Hippomane mancinella L., manchineel. B s.n.; LG sight (Dog Is., Scrub Is.); P 18807
A
Jatropha curcas L., barricata bush. HK sight.
ae sossypifolia L., physic nut. HK “sect - LG sight (Dog Is.); P 18680 (a).
opha integerrima Jacq. Cultivated. HK sig
oe multifida L. Cultivated. HK sight.
Manihot esculenta Crantz. Cultivated. HK sight.
ss laa tithymaloides (L.) Poit., Sie dae heart, candle flame bush. B 3572; P 15616
Phan amarus Schum. & Thonn., churchweed. HK 20115 (a), 20148 (a); P 18638
Phan nee L., bilbush. B 32566; E 33; LG sight (Scrub Is.); HK 20078
(A); P 6 (A).
Ricinus communis L., castor nut. Naturalized. HK sight.
GOODENIACEAE
Scaevola plumieri (L.) Vahl, candlewood. B 3563; HK 20123 (A); P 18781 (A).
GUTTIFERAE
Clusia rosea L., autograph tree, pitch apple. HK 201/54 (a); P 18631 (a).
LABIATAE
Bee tg (L.) Aiton, hollow stalk. P 18702 (a).
icranthum Willd., French basil. HK 20072 (a).
Be ies amboinicus (Lour.) Launert, stingy time. Cultivated. B s.n.
Plectranthus blumei (Bentham) Launert. Cultivated. HK sight.
Salvia occidentalis L., cat mint. P 18575 (a).
Salvia serotina L., Gat mint. B s.n.; HK 20144 (a); P 18662 (a).
LAURACEAE
Cassytha filiformis L. B 3523; P 18668 (a).
Persea americana Pers. Cultivated: B3491/a.
LEGUMINOSAE
Acacia farnesiana Willd., queen casha. LG sight (Dog Is.); P 18528 (a).
Acacia macracantha Humb. & Bonpl., kushar. HK 20087 (a); P 18774 (a).
Caesalpinia bonduc (L.) Roxb., nicker tree. P 18529 (A).
Caesalpinia coriaria (Jacq.) Willd. P 18671 (a).
Caesalpinia divergens Urban, red nicker. HK 20128 (a); LG sight (Dog Is., Scrub Is.),
as Guilandina divergens Urban; P 18562 (A).
124 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Caesalpinia pulcherrima Sw., pride of Barbados. Cultivated. P 18759 (a).
Cajanus cajan (L.) Huth, Angola pea, pigeon pea. Cultivated. HK sight.
Canavalia rosea (Sw.) DC. P 18782 (a
Centrosema virginianum Bentham. HK 20158 (A); P 18547 (A).
Chamaecrista glandulosa (L.) Greene var. swartzii (Wikstr6m) Irwin & Barneby, wild
tamarind. B 3516; P 18563 (A).
Crotalaria incana L. P 18698 (A), 18734 (A).
Crotalaria retusa L., shack-shack. P 18736 (A).
Crotalaria verrucosa L. HK 20152 (A); P 18655 (A).
Delonix regia (Bojer) Raf., flamboyant. Cultivated. HK sight.
Desmanthus virgatus (L.) Willd. HK 20107 (a); P 18604 (A
Desmodium frutescens Schindler var. angustifolium Schindler. LG 2515.
Erythrina variegata L. var. orientalis (L.) Merr. Cultivated. HK sight.
Galactia dubia DC. B 3503; HK 20069 (a); P ath la ).
Gliricidia sepium (Jacq.) Kunth, quick set. HK s
Indigofera suffruticosa Miller. P 18701 (a).
Indigofera tinctoria L. HK 20111 (a); P 18735 (a
Lablab purpureus (L.) Sweet, bonavist. Cultivated. HK sight.
Leucaena leucocephala (Lam.) De Wit, mimosa, wild tamarind. B s.n.; P 18764 (a).
Neptunia pubescens Bentham. P 1875
Parkinsonia aculeata L. Cultivated and ais: HK sight.
Pithecellobium unguis-cati (L.) Martius, bread and cheese, crabwood, grooven-eye.
Bs.n.; E 57; LG sight (Dog Is., Scrub Is.); P 18540 (a).
Riynehosia minima DC. B 3486; P 18574 (a).
Rhynchosia reticulata (Sw.) DC. HK 20153 (A); P 18549 (a).
Senna bicapsularis (L.) Roxb. E 58; P 18530 (A).
Senna italica Miller. B s.n., as Cassia obovata Colladon
Senna obcordata (Wikstrém) Britton. LG 2473, as Cassia obcordata Sw.
Senna obtusifolia (L.) Irwin & Barneby. P 18672 (a)
oe eae (L.) Link, bush coffee, stinkweed. B s.n., as Cassia occidentalis L.;
0133 (A); P 18795 (A), 18816 (A).
iar siamea (Lam.) Irwin & Barneby. Seen - sight.
Sesbania grandiflora (L.) Pers. Cultivated. HK s
Sophora tomentosa L. B 3490; LG 2481, 2494, 296. 2497, 2498, 2502; P 18580 (a).
— hamata (L.) Taubert, sweetweed, wild Isaac. B 3536; HK 20083 (a);
P 18763
eperi indica L., tamarind. HK sight.
Tephrosia cinerea Pers, P 18700 (a).
LOGANIACEAE
Spigelia anthelmintha L. B 3474, HK 20163 (a).
LORANTHACEAE
Dendropemon caribaeus Krug & Urban. HK 20132 (a).
Phoradendron trinervium (Lam.) Griseb., mistletoe. B s.n.; HK 20049 (a), 20050 (a);
P 18564 (A).
LYTHRACEAE
Lawsonia inermis L., mignonette. Cultivated. HK sight.
MALPIGHIACEAE
Byrsonima lucida Rich., gooseberry, goosie tree. B 3501, 3511, E 36; HK 20166 (a);
P 18536 (A).
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 125
Galphimia gracilis Bartling. one oe HK sight.
Heteropteris purpureus (L.) K EF 4].
Malpighia emarginata Sessé I ae ex DC., sherry. B s.n., as Malpighia punicifolia
L.; P 18777 (a).
Malpighia linearis Jacq. LG sight (Scrub Is.).
Be Mega diversifolium (Kunth) A. Juss. B 3513; HK 20120 (a); LG 2486, 2499;
Ben emarginatum (Cav.) Juss. HK 20093 (a), 20097 (a).
Stigmaphyllon ne (Poiret) Small. B 3452, as Stigmaphyllon periplocifolium
A. Juss.; P 18519 (a
MALVACEAE
Abelmoschus esculentus (L.) Moench, okra. Cultivated. HK sight.
Abutilon indicum (L.) Sweet. B s.n.
Abutilon umbellatum (L.) Sweet. P 18766 (A).
Bastardia viscosa (L.) Kunth. P 18678 we
Gossypium barbadense L. Cultivated. H
Herissantia crispa (L.) Briz. HK 20160 o " ey (A).
Hibiscus rosa-sinensis L. Cultivated. HK s a
Hibiscus sabdariffa L., sorrel. Cultivated. HK s
Malvastrum coromandelianum (L.) Garcke. B 3449, as Malvastrum tricuspidatum
A.
Sida abutilifolia ae HK 20116 (a); P 18681] (a).
Sida acuta Burman f., jingle weed. P 18756 (a).
Sida ciliaris L. B 3493, 3535; HK 20073 (a); P 18649 (a).
Sida spinosa L., wild Isaac. B 346la; HK Seer he P 18673 (a).
Thespesia populnea (L.) Sol. ex Correa. HK s
MELIACEAE
Azadirachta indica A. Juss., neem. Cultivated. HK 20113 (A).
Melia azedarach L. Cultivated. HK sight
Swietenia mahagoni Jacq. HK sight; LG sight.
MORACEAE
Artocarpus altilis (Parkinson) Fosb., breadfruit. Cultivated. HK sight.
Ficus citrifolia Miller. LG sight (Scrub Is.); P 18644 (a).
Ficus elastica Roxb., rubber tree. Cultivated. HK sight.
MorINGACEAE
Moringa oleifera Lam. Cultivated or escaped. HK sight.
MyrTACEAE*
Eugenia axillaris (Sw.) Willd., sneeze berry. HK 20056 (A); LG sight (Scrub Is.); P 18815
(A).
*An unknown member of this family was collected by Le Gallo as nos. 2480 and 2493 (May 9,
1955) near the Catholic church. Since this time a new church was poe fu ule old building 1 remains
and is surrounded by dense scrub vegetation. Our i
126 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Eugenia foetida Pers., white wattling. HK 20094 (a), 20143 (a); P 18518 (a), 18552 (a),
18689 (A), 18748 (A).
Eugenia monticola (Sw.) DC., cuttard. B 3461, 3477a, 3520.
Pimenta racemosa (Miller) J. Moore, bay leaf. P 18622 (a).
Psidium guajava L. B s.n., HK sight
Psidium longipes (Berg) McVaugh var. orbicularis (Berg) McVaugh. B 3509 (Ny), as
Myrtus anguillensis Urban, LG 2067, 2492, 2508, 2517; P 18801 (A).
NYCTAGINACEAE
Boerhavia coccinea Miller. HK 20117 (a); P 18579 (a).
Boerhavia scandens L., piecrust. B 3451; HK 20091 (a); LG sight (Dog Is.); P 18602 (a).
Bougainvillea glabra Choisy. Cultivated. HK sight
Guapira fragrans Dum.-Cours. P 18814 (A).
Mirabilis jalapa L. Cultivated. HK sight.
Pisonia subcordata Sw., loblolly. B s.n.; P 18785a (A), 187856 (A).
OLEACEAE
Forestiera eggersiana Krug & Urban. HK 20139 (a); P 18544 (a), 18798 (A).
Jasminum fluminense Vell. Cultivated. P 18768 (a).
PAPAVERACEAE
Argemone mexicana L., thistle. B s.n.; E 80; LG sight (Dog Is.); P 18741 (a).
PASSIFLORACEAE
Passiflora edulis Sims, passionfruit. Cultivated. HK sight.
Passiflora foetida L., pops. HK 20096 (a).
Passiflora suberosa om pops. B 3498; LG 2510; P 18538 (a).
PERIPLOCACEAE
Cryptostegia grandiflora R. Br. Cultivated and naturalized. HK sight.
PHYTOLACCACEAE
Rivina humilis L. P 18813 (a).
PLUMBAGINACEAE
Plumbago auriculata Lam. Cultivated. HK sig
Plumbago scandens L., doctor John. HK ire os P 18600 (A).
POLYGONACEAE
Antigonon leptopus Hooker & Arn. Naturalized. HK s
Coccoloba nie gii Lindau, wild grape. B 3472; HK 20052 7 20068 (A); LG 2058, 2069;
PIS52
Coccoloba baie Lindau = Coccoloba uvifera (L.) L., wild grape. HK 20135 (a), 20136
(A) (each of these collections represents a slightly different phase of the hybrid).
Coceoloba microstachya Willd. B 3458, 3483 (Ny), both as Coccoloba diversifolia Jacq.
Coccoloba uvifera (L.) L., sea grape. P 18612 (a).
PORTULACACEAE
Portulaca halimoides L., pussley. B 3552; E 69; HK 20084 (a); P 18572 (a).
Portulaca oleracea L. B 3553; P 18683 (a).
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 127
PUNICACEAE
Punica granatum L. Cultivated and naturalized. HK 20055 (a).
RHAMNACEAE
Colubrina arborescens (Miller) Sarg., mawby bark. E 34; HK 20066 (a); LG 2051, 2068
(both Dog Is.), both as Colubrina ferruginosa Brongn.; P 18645 (A
Krugiodendron ferreum (Vahl) Urban, ebony berry. HK 20162 (A); P 1875 I (a).
Reynosia uncinata Urban, sloe. B 3470; HK 20092 (a); P 18533 (A
Zizyphus mauritiana Lam., doms, pommeserrette. HK 20054 (a), P 18654 (A).
Zizyphus rignonii Delponte, thorn. B 3452a (Ny), 3488 (not located), 3506a (Ny), all as
Zizyphus havanensis, HK 20102 (a), LG 2053, 2066, sight (Scrub Is.), all as Sarcom-
phalus domingensis (Sprengel) Krug & Urban; P 18535 (a)
RHIZOPHORACEAE
Rhizophora mangle L., whistle. HK sight; LG sight.
ROSACEAE
Rosa indica L., rose. Cultivated. HK sight.
RUBIACEAE
Antirhea page (DC.) Urban, mutton polly. B 3475a; HK 20099 (a), 20101 (a),
LG 2071; P 18605 (a).
Feet ee L., candlebush. B 3486; E 40; HK 20044 (a); P 18650 (a).
Ernodea littoralis Sw., cough bush, stinging whip. B 3487, 3510; E 32; HK 20047 (a),
P 18545 (a).
Exostema caribaeum Roemer & Schultes, fustic. B 3476, 3502; E 37; HK 20060 (a);
P 18557 (a).
Guettarda scabra La m., chink, wild guava. B 3464, 3506; HK 20095 (a); P 18556 (a).
sight.
Randia aculeata L., five-finger tree. B 3450a; HK 20053 (a); P 18560 (a).
Rondeletia anguillensis R. Howard & E. Kellogg, sp. nov. Ficures 4, 5.
Frutex cruciatus, foliis minutissimis, minoribus quam 4 mm longis, floribus distylis
Stiff shrub up to 1 m tall; branches divaricate, spine tipped; bark smooth, grayish;
ulent; blade suborbicular when young, becoming ovate to elliptic, 1.5-3.8 x 1.2-2.7
mm, the apex rounded, the base rounded, the margin thickened-revolute, the adaxial
surface shiny, dark green, glabrous, the abaxial surface white-velutinous, the midvein
white abaxially. Flowers stiffly erect to horizontal, 4-merous, subtended by crateriform
m long; corolla salverform, pale pink, appressed-white-puberulent externally, the
tube 4-5.7 mm long in short-styled plants and 3.2-3.7 mm long in long-styled ones,
glabrous within, the lobes 0.8-1.6 mm long, puberulent above, with annulus of small,
raised tubercles at throat; stamens inserted on corolla tube, the filament < 0.2 mm
long, filiform, the anther dorsifixed, oblong to slightly cuneate, 1.4-1.6 mm long in
both forms, the pollen 3-colpate in short-styled plants, 3- and 4-colpate in area
ones; style linear, 2—-2.2 mm long and sparsely retrorse-pubescent in short-styled plants,
7 mm long and nearly glabrous in long-styled ones, the stigmas 2, 0.8 mm long
128 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
IGURE 4. Rondeletia anguillensis: a, habit (Proctor 18571, a), x 0.75; b, lower
surface of leaf (Proctor 18571), x 7; c, dehisced fruit (Proctor 18571), x 11; d, short-
styled flower (Howard & Kellogg 20105, A), x 7; e, long-styled flower (Howard & Kellogg
20103, A), x 10. (a drawn by M. Dykens, b-e by I. Al-Shehbaz.)
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 129
Figure 5. Rondeletia anguillensis (Howard & Kellogg 20103, a), seed. Scale =
100 um.
in short-styled plants and 0.3 mm in long-styled ones; the ovary 2-locular, obconical,
1.2 mm long, stiffly erect-pubescent. Fruit capsular, globose, 1.8-2.9 mm in diameter,
initially loculicidal, subsequently septicidal; calyx lobes persisting and becoming 0.8-
mm long; placenta peltate, reniform in longitudinal section, becoming massive,
hemispheric: seeds numerous, 0.72-0.75 mm long, imbricate upward, irregular in
outline, finely reticulate.
Type. Anguilla, east end of island, 6 Feb. 1985, R. Howard & E. Kellogg 20105
(holotype, A).
SPECIMENS SEEN. Angpuilla: E end of island, R. Howard & E. Kellogg 20103 (a), vic. of Little Bay,
near Flat Cap Point, Proctor 18571] (A).
Spermacoce confusa Rendle, chicken weed. B 3542; HK 20064 (a); P 18642 (A).
Strumpfia maritima Jacq. B 3551; E 49, LG sight (Dog Is., Scrub Is.), P 18617.
RUTACEAE
Ampris elemifera L., ironwood. B 3481; HK 20076 (a); LG 2056 (Scrub Is.); P 18517
Citrus plies (Christm.), Se lime. ne HK sight.
Citrus aurantium L., sour orange. Cultivated. HK s
Citrus panes Macfad.. ett Cultivated. HK «
Citrus sinensis (L.) Osbeck, sweet orange. Cultivated. ie sight.
Murraya paniculata (L.) Jack. Cultivated. HK sight.
Zanthoxylum flavum Vahl, alexander. B 3525; HK 20063 (a); LG sight (Scrub Is.);
P 18541 (a), 18670 (A).
Pe Bie punctatum Vahl, ironwood. B 3469, 3526, 3532, all as Fagara trifoliata
Sw.; HK 20077 (A).
Zanthoxylum spinifex (Jacq.) DC., ramgoat. B 3529, as Fagara spinifex Jacq.; HK 20090
(A); P 18584 (A), 18799 (A).
130 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
SAPINDACEAE
Cardiospermum corindum L. P 18554 (A).
Hypelate trifoliata Sw., ironwood. B 3508; LG 2410, hie 2479, 2514; P 18573 (a).
Meliococcus bijugatus L., genip. Cultivated. HK sig
SAPOTACEAE
Bumelia obovata (Lam.) DC., thorn tree. HK 20048 (a); LG sight (Dog Is.); P 18589 (A).
Bumelia salicifolia (L.) Sw., mass wood. B 3542; HK 20138 (a); P 18642 (A).
SCROPHULARIACEAE
Capraria biflora L., tasane. LG sight aa Is.); P 18697 (a).
Lindernia diffusa (L.) Wettst. HK 20074 (a).
SIMAROUBACEAE
Castela erecta Turpin, cockspur. HK 20089 (a); LG 2054, 2065 (both Scrub Is.), 2470
(Dog Is.), 2500; P 18592 (a), 18606 (a
Suriana maritima L. B 3560; E 51. LG sight (Scrub Is.); P 18614 (a).
SOLANACEAE
Capsicum frutescens L., pepper. Cultivated. HK sight.
Datura stramonium L. P 18738 (a).
Lycium americanum Jacq. B 3546; HK 20134; LG sight (Dog Is., Scrub Is.); P 18747
(A).
Lycopersicon ee (L.) Karsten, tomato. Cultivated. HK sight.
Physalis angulata L. P 18796 (a).
Solandra guttata D. Don. Cultivated. HK s
Solanum melongena L., bolonge, eggplant. cee HK sight.
Solanum racemosum Jacq. canker berry, cob berry, conka berry. B s.n.: E 53: HK 20088
(A); LG sight (Dog Is., Scrub Is.); P 18527 (a).
STERCULIACEAE
Melochia pyramidata L. P 18524 (a).
Melochia tomentosa L. B 3478; E 60, HK 20082 (a); LG sight (Scrub Is. ); P 18593 (a).
Waltheria glabra Poiret. P 18569 (a
Waltheria indica L., marshmallow. B 3496, as Waltheria americana L.; E 43: P 18603
(A).
TAMARICACEAE
Tamarix chinensis Lour. Cultivated. HK sight.
THEOPHRASTACEAE
Jacquinia arborea Vahl, scrub bush. HK 20046 (a), LG sight (Scrub Is.); P 18567 (a).
Jacquinia berterii Sprengel. B 3558; E 61; HK 20059 (a); LG 2060, 2063 (both Scrub
Is.); P 18537 (a), 18632 (A).
TILIACEAE
Corchorus hirsutus L., marshmallow. B 3473, 3519; LG sight (Scrub Is.); P 18570 (a),
S2(A
Corchorus siliquosus L., calaloo. B 3463; P 18561 (a).
1987] HOWARD & KELLOGG, FLORA OF ANGUILLA 131
TURNERACEAE
Turnera ulmifolia L. P 18731 (A).
ULMACEAE
Celtis iguanaea (Jacq.) Sarg. B s.n.
UMBELLIFERAE
Anethum graveolens L. P 18676 (A).
URTICACEAE
Pilea serpyllifolia (Poiret) Wedd. Cultivated. HK sight.
VERBENACEAE
Citharexylum fruticosum L. P 18624 (a).
Clerodendrum aculeatum (L.) Schldl. B s.n.; LG sight (Dog Is.); P 18740 (a).
Duranta erecta L. B s.n., as Duranta repens s Kin th.
Lantana camara L., sage cop. P 18786 (A).
Lantana involucrat L., sage, sage cop. B 3476, 3504 (ny); E 44; HK 20106 (a); P 18526
(A), 186
Lippia ae Kunth, B 3475; LG sight (Dog Is.).
Lippia strigulosa Martens & Gal. P 18677 (A).
Priva lappulacea (L.) Pers. P 18599 (a).
Stachytarpheta jamaicensis (L.) Vahl, worry wine. B 3497; HK 20157 (a), P 18775 (a).
VIOLACEAE
Hybanthus portoricensis Urban. LG sight (Scrub Is.).
VITACEAE
Cissus verticillatus (L.) Nicolson & Jarvis. B s.n., as Cissus sicyoides L.
ZYGOPHYLLACEAE
Guaiacum officinale L., lignum vitae. Cultivated. P 18800 (a).
Kallstroemeria maxima (L.) Torrey & A. Gray. P 18685 (A).
KELLOGG & HOWARD, POLLEN DIMORPHISM 133
UNUSUAL POLLEN DIMORPHISM IN
RONDELETIA ANGUILLENSIS (RUBIACEAE)
ELIZABETH A. KELLOGG AND RICHARD A. HowArp!
Long-styled plants of Rondeletia anguillensis bear a mixture of three- and
four-colpate pollen, whereas short-styled plants bear only three-colpate grains.
Short-styled plants have smaller pollen grains with lower pollen stainability
than long-styled plants.
In the preceding paper (Howard & Kellogg, 1987) we described Rondeletia
anguillensis, a new species collected on the Caribbean island of Anguilla in
1985. The plants were clearly distylous, a condition common in the Rubiaceae.
Measurements confirmed that short-styled plants had notably longer corolla
tubes than long-styled ones (4—5.7 vs. 3.2-3.6 mm, respectively); this difference
in corolla size has been reported for other distylous plants (Ganders, 1979). In
the process of preparing the description, we discovered that the pollen of the
two stylar forms was more strongly dimorphic than is commonly the case in
the family. We report our results here in the hope of stimulating further col-
lecting and investigation of the phenomenon.
METHODS
Our observations were based on material from three collections, two short-
styled plants (Howard & Kellogg 20105, a; Proctor 18571, A) and one long-
styled one (Howard & Kellogg 20103, a) (for full specimen citations, see Howard
& Kellogg, 1987). After preparing SEM photographs of pollen from each of
the two forms, we continued investigations with the light microscope. Pollen
was stained overnight with cotton blue in lactophenol. For each plant the
diameters of 200 pollen grains from a single flower were measured and averaged.
Pollen stainability was calculated from more than 200 grains for each flower
observed (four flowers for each of the Howard & Kellogg collections, one for
the Proctor collection). For each of these nine flowers, the number of colpi was
recorded for the first 200 grains from which it could be determined.
RESULTS AND DISCUSSION
The photographs in the Figure show that the long-styled plant has a mixture
of three- and four-colpate pollen (a, b), whereas the short-styled plant bears
‘Arnold Arboretum, Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 133-136. January, 1987.
134 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Rondeletia anguillensis, nonacetolyzed pollen (scale bars = 10 um): a, b, long-styled
plant, Howard & Kellogg 20103, c, d, short-styled plant, Howard & Kellogg 20105.
consistently three-colpate grains (c, d). Also, pollen from the short-styled plant
appears to be slightly smaller than that from the long-styled one.
These observations were confirmed by light microscopy, the results of which
are summarized in the TABLE. Pollen size and stainability were virtually iden-
tical for the two short-styled plants, despite the fact that they were collected
26 years apart and at different localities on the island. Unfortunately, we had
only one long-styled plant, and that had very few nearly mature buds from
which to take pollen, so we cannot be certain that the pollen dimorphism is
characteristic of the species. There is still the possibility that Howard & Kellogg
20103 is simply an anomalous plant.
These preliminary results show some surprising differences between the two
style forms. Statistical comparison of the mean sizes of pollen of the two
Howard & Kellogg collections produces tf = 8.96, df = 398, a difference sig-
nificant at p < 0.005. However, the short-styled plants have the smaller pollen,
contrary to the condition in most other distylous plants. Ganders (1979) re-
ported pollen of short-styled plants to be smaller than that of long-styled ones
in Fauria crista-galli Makino (Menyanthaceae); this was apparently the first
report of such a size relationship.
Variation in pollen stainability has been reported by Ornduff (1980) in pop-
ulations of Hedyotis caerulea (L.) Hooker. He reported variation both between
and within short- and long-styled plants, and variation between and within
years. There may be similar variability in Rondeletia anguillensis, but its sig-
nificance is unclear.
1987] KELLOGG & HOWARD, POLLEN DIMORPHISM 135
Variation in pollen size, colpus number, and stainability for three plants of
Rondeletia anguillensis.
Num-
BER
SIZE (um
a (um)
SPECIMEN coLtpl Range Mean SD STAINABILITY (%)
parte
oward & ease 20105 3* 11-15 13 0.75 72, 76, 93, 94
pane 18571 3 11-15 13 0.76 76
Long-style
Howard & Kellogg 20103 3,4 12-15 14 0.61 99, 99, 99.5, 99.5
*One 4-colpate grain observed out of more than 900. Most probably contamination.
The variation in colpus number is interesting for two reasons. First, mor-
phological differences between long- and short-style pollen are usually subtle,
particularly in the Rubiaceae. Major differences in shape have been reported
for Lithospermum L. (Boraginaceae; Johnston, 1952), and differences in exine
sculpturing occur in the Plumbaginaceae (Baker, 1966). The only really marked
difference in pollen morphology for distylous Rubiaceae was reported by Baker
(1956) for Rudgea jasminoides (Cham.) A. Rich., in which the pollen of long-
styled plants was smooth and that of short-styled ones was spiny.
Second, four-colpate pollen is one of the major characters used by Borhidi
and colleagues (1980, 1981) in distinguishing Rondeletia L. and the segregate
genera Roigella Borhidi & Zequeira, Neomazaea Urban, and Acuneanthus
Borhidi, Jarai-Komlodi, & Moncada; pollen of Rondeletia and Acuneanthus is
three-colpate, while that of the other two genera is four- or five-colpate. The
variation we have found in this character casts some doubt on its usefulness
at the generic level. Its variability within species should perhaps be investigated
more fully before it is relied upon for major distinctions among groups.
Four flowers from the long-styled plant were scored for percent of three-
colpate pollen. The percentages were 47, 51, 59, and 62. The pooled x? = 7.0,
indicating that the ratio of three- to four-colpate pollen was significantly dif-
ferent from 1:1 (p < 0.01). However, a test for homogeneity of x? values among
the flowers showed a significant lack of homogeneity, so pooling the values
may not be justified. If x* values are calculated separately, values for two of
the flowers are not significantly different from a 1:1 ratio, whereas values for
the other two are. If we assume that the two pollen morphs would indeed
appear in equal proportions if the sample were sufficiently large, then we could
explain the observed variation in pollen morphology by a one-locus gene with
two alleles, one of which conditions for three colpi and the other for four.
Under this explanation, the long-styled plant is heterozygous, the (haploid)
pollen grains therefore being half three- and half four-colpate, whereas the
short-styled plants are homozygous. If this proves to be the case, it would be
an interesting parallel with the gene for distyly itself, which in all reported cases
is also one locus with two alleles and complete dominance (Ganders, 1979).
136 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
One style morph is then homozygous recessive (ss), while the other 1s hetero-
zygous (Ss). The homozygous dominant does not occur because of self-incom-
patibility of the heterozygotes. Although the short-styled plants are the het-
erozygotes in most species, long-styled heterozygotes have been reported
(Ganders, 1979). It is conceivable that the gene for pollen shape could be linked
with the gene for distyly to produce the pattern we have observed in Rondeletia
anguillensis.
ACKNOWLEDGMENTS
This study was carried out with support from National Science Foundation
grant BSR 83-07701
LITERATURE CITED
Baker, H. G. 1956. Pollen dimorphism in the Rubiaceae. Evolution 10: 23-31
—. 1966. The evolution, functioning and breakdown of heteromorphic incom-
patibility systems. I. The Plu pareve Ibid. 20: 349-368.
Boruipi, A., M. JARAI-KOMLopI, & M. Moncapa. 1980. Acuneanthus, a new genus
of Rubiaceae. Acta Bot. Acad. Sci. ie 26: 277-287.
. ZEQUEIRA. 1981. Studies in Rondeletieae (Rubiaceae) I. A new genus:
Roigella. Acta Bot. Acad. Sci. Hung. 27: 309-312.
GANDERS, F. R. 1979. The biology of heterostyly. New Zealand J. Bot. 17: 607-635.
Howarp, R. A., & E. A. KELLOGG. 1987. Contributions to a flora of Anguilla and
adjacent islets. J. Arnold Arbor. 68: 105-131.
Jounston, I. M. 1952. Studies in the Boraginaceae. X XIII. A survey of the genus
Lithospermum. J. Arnold Arbor. 33: 299-363.
OrnburF, R. 1980. Heterostyly, population composition, and pollen flow in Hedyotis
caerulea. Amer. J. Bot. 67: 95-103.
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Journal of the Arnold Arboretum January, 1987
CONTENTS OF VOLUME 68, NUMBER |
Phylogenetic Implications of Leaf Anatomy in Subtribe Melittidinae
(Labiatae) and Related Taxa.
Mones S. ABu-ASAB AND PHILIP D. CANTINO ................ 1-34
The Genera of Pontederiaceae in the Southeastern United States.
SICA TTS ATY os a's con gs Ke Volo baad os LoS eed nig dines 35-71
Reproductive Structure of Lithocarpus Sensu Lato (Fagaceae): Cy-
mules and Fruits.
POSER GAUGE yoink igo kh Sy eed ed odd 140 nb ns baenae, 73-104
Contributions to a Flora of Anguilla and Adjacent Islets.
RICHARD A. HOWARD AND ELIZABETH A. KELLOGG ........... 105-131
Unusual Pollen Dimorphism in Rondeletia anguillensis (Rubi-
aceae).
ELIZABETH A. KELLOGG AND RICHARD A. HOWARD ........ geet 133-136
Volume 67, Number 4, including pages 371-512, was issued October 8, 1986.
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JOURNAL
OF THE
ARNOLD ARBORETUM
VOLUME 68 APRIL 1987 NUMBER 2
THE GENERA OF CINCHONOIDEAE (RUBIACEAE) IN THE
SOUTHEASTERN UNITED STATES!
GEORGE K. ROGERS?
The infrafamilial classification of the Rubiaceae is in an unsettled state, with
solid answers awaiting accumulation and interpretation of data on some 500
genera. Schumann’s system, the only clear, comprehensive one, is followed in
the present account. This is not to say that it satisfactorily reflects natural
relationships, for it does not—it rests upon heavy-handed application of a few
4h fel }
'Prepared for United States, a long-term project made possible
by grants from the National Science Foundation and at this eee supported by BSR-8415769
(Carroll E. Wood, Jr., principal investigator) and BSR-8415637 (Norton G. Miller, principal investi-
gator). This treatment, the 114th in ane. series, eae the format ea in the first paper (Jour.
Arnold Arb. 39: 296-346. 1958) inued to the present. The area covered by the Generic Flora
includes North and South Carolina, nes ae Tennessee, Alabama, Mississippi, Arkansas,
and Louisiana. The descriptions are based primarily on the plants of this area, with information
about extraregional members of a family or genus in brackets [ ]. References that I have not verified
are marked with an asterisk.
Treatments of the first four genera were prepared at the Arnold Arboretum while I held a post-
doctoral appointment th e remainder were prepared at the Missouri Botanical Garden. I owe
thanks to the Rubiaceae researchers of St. Louis, who meet occasionally for discussion. This g
has broadened my perspective on the family and has been th rce of a great deal of factual
helped generously with aspects of the de ae the heme base in C sane risnenecateay ies hile te
Carroll Wood supplied information, guidance, and ins
and Stephen Spongberg improved the manuscript with their good ‘deas, oe Missouri Botanical
Garden provided space and facilities.
The illustrations were drawn by Rachel A. Wheeler (Cephalanthus), Dorothy H. Marsh (Casasia),
and an Stoutsenberger (Hamelia) from materials prepared by Carroll Wood. The specimens of
Cep a rom the Arnold Arboretum (Wood) and Louisiana (Joseph Ewan, Gu); those of
Casi Bee Hamelia from Big Pine Key, Monroe County, Florida (Wood).
?Missouri Botanical Garden, P. O. Box 299, St. Louis, Miseaun 63166.
© President and Fellows of Harvard College, 1
Journal of the Arnold Arboretum 68: 137-183. pen 1987.
138 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
characters, and students of the Rubiaceae have since stressed that it breaks
apart obvious alliances.
Even Schumann’s fundamental division of the Rubiaceae into two subfam-
ilies, the Cinchonoideae and the Rubioideae (Coffeoideae), is based on a single
character, the number of ovules in each locule of the ovary (multiple in the
former, solitary in the latter). His classification provides, nonetheless, a con-
venient and useful framework.
The two foremost students of the Rubiaceae since Schumann, Verdcourt
(1958 and later works; see especially 1976) and Bremekamp (particularly 1966),
have proposed reforms of the infrafamilial classification. Although neither
assembled a comprehensive new scheme, both have added new insights, and
both have laid out their concepts of the tribes and subfamilies with character-
izations and discussion. Further, both have supplied thorough histories of the
subject. So that their contributions do not pass ignored, a summary of the
various dispositions of our genera in comparison with Schumann’s follows.
Bremekamp increased the number of subfamilies from Schumann’s two to
eight, of which three concern us. He redefined the Rubioideae as members of
the Rubiaceae having raphides and generally valvate corollas. With emphasis
shifted to these characters (especially the former), the Hedyotideae (including
our Hedyotis L. sensu lato and Pentodon Hochst.) were moved from the Cin-
chonoideae to the Rubioideae. Also, Hamelia Jacq., which has raphides, was
transferred along with Hoffmannia Sw. from tribe Gardenieae in the Cincho-
noideae to the resurrected Hamelieae DC. in the Rubioideae (see generic treat-
ment).
Bremekamp did not leave the remainder of Schumann’s tribe Gardenieae
in the Cinchonoideae; instead, he transferred it (containing our Randia, Ca-
sasia, and Catesbaea) to the Ixoroideae Raf., a subfamily he composed of tribes
showing the “ixoroid” pollination mechanism (pollen deposited on the shaft
of the style). My suspicion is that the ixoroid pollination mechanism is too
widespread, either by convergence or by persistence from distant common
ancestry, to be a reliable character in defining a subfamily of the Rubiaceae.
It shows up in Pentodon, clearly a member of the Hedyotideae, and in such
other families as the Loganiaceae, Campanulaceae, and Compositae. Breme-
kamp was uncertain of the placement of Cephalanthus.
Verdcourt’s strong Old World emphasis makes it difficult to apply his ideas
to our genera. He recognized three subfamilies, including the Cinchonoideae
and the Rubioideae, defined primarily by the presence or absence of raphides.
Verdcourt (1958, 1976), like Bremekamp, placed Hamelia and our genera of
Hedyotideae in the Rubioideae. He departed from Bremekamp and Schumann
by merging tribe Condamineeae (containing Pinckneya) with the Rondeletieae
(DC.) J. D. Hooker & Bentham (FI. Nigritana, 378. 1849; note earlier au-
thorship than that given by Darwin). Verdcourt agreed with Schumann but
disagreed with Bremekamp, placing Cephalanthus in the Naucleeae and re-
taining the Gardenieae (minus Hamelia) in the Cinchonoideae. Among the
authors of interest, he is unique in segregating tribe Catesbaeeae J. D. Hooker
from the Gardenieae (see treatment of Catesbaea).
To summarize the present state of affairs, in my view the size of the family
1987] ROGERS, CINCHONOIDEAE 139
Rubiaceae forces botanists concerned with its infrafamilial subunits to sub-
divide it ‘from the top down,” stressing differences found in a few characters.
Much discussion connected with the problem centers around the comparative
(not convincingly substantiated) “importance” of various characters for this
purpose. Only massive collection of new data and a new, more evolutionary
emphasis will eventually allow infrafamilial groups to be built “from the bottom
up,” buttressed by shared derived similarities.
For those workers interested in determining the correct names of taxa of the
Rubiaceae above the rank of genus, S. P. Darwin’s thoroughly researched
nomenclator for subfamilies, tribes, and subtribes in the family is indispensable.
RUBIACEAE eg CINCHONOIDEAE Rafinesque, Ann. Gén. Sci. Phys.
1 (p. 66 in reprint). 1820, ““Cinchonaria.”
Trees or shrubs (except Hedyotis sensu lato and Pentodon) with usually
opposite, sometimes whorled or fascicled, leaves. Stipules interpetiolar, gen-
erally with 1 (sometimes bifid) lobe between adjacent petiole bases (to fimbriate
in Hedyotis and Pentodon, becoming shredded in Randia), usually bearing
colleters on the adaxial side. Flowers pentamerous or tetramerous, with tubular
corollas. Ovary inferior, usually bilocular (but with up to 5 locules in Hamelia;
Casasia unilocular but appearing bi- or trilocular), the locules generally mul-
tiovular (uniovular in Cephalanthus, Randia sometimes with a single seed in
the fruit). Type GENUS: Cinchona L.
REFERENCES:
Apams, C. D. Flowering plants of Jamaica. 848 pp. Mona, Jamaica. 1972. [Rubiaceae,
699-733.]
Atatn, Hno. [Liocier, E. E.]. Rubiaceae. Fl. Cuba 5: 13-146. 1962.
ANGELY, J. Flora ae e geben a do Estado de Sao Paulo. Vol. 4. Pp. [1-16 +]
17-36 + 685-892 + x. S40 Paulo. 1970. [Rubiaceae, 767-800.
BaILLon, H. Rubiacées. Hist PL 7: 257-503. 1880. English translation, Rubiaceae. /n:
The natural history of plants 7: 257-503. 1881.
Barker, H. D., & W. S. DarpeaAu. Flore d’Haiti. viii + 456 pp. Port-au-Prince. 1930.
BENTHAM, G., & J. D. Hooker. Rubiaceae. Gen. Pl. 2: 7-151, 1227-1229. 1873. [Ru-
biaceae in “‘series,”’ ““subseries,”” and tribes.]
BREMEKAMP, C.E. B. The African species of O/den/andia L. sensu Hiern et K. Schumann.
Verh. Nederl. Akad. Wet. Afd. Natuurk. 2. 48(2): 1-297. 1952. [Position of Hedy-
otideae, 1 1-25; includes revision of Pentodon, comments on typification of Hedyotis,
and characterization of Oldenlandia.]
. Remarks on the position, the delimitation and the subdivision of the Rubiaceae.
Ac ta Bot. Neerl. 15: 1-33. 1966.
Brizicky, G. K. aa and sectional names: their starting points and early sources.
Taxon 18: 643-660.
BuswELL, W. M. Native erin of south Florida. Bull. Univ. Miami 20(3): 1-48. 1946.
[Rubiaceae, 43—47.]
CANDOLLE, A. P. pe. Rubiaceae. DC. Prodromus 4: 341-622, 672, 673. 1830.
CorreELL, D. S., & H. B. Correy. Flora of the pares Archipelago. a +] 1692 pp.
Vaduz, Licehtestin. 1982. [Rubiaceae, 1366-14
—— &M. NSTON. Manual of the vascular ae of Texas. xv + 1881 pp.
Dallas. 1979. ieee 1479-1496.]
140 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Darwin, S. P. The ea tribal and subtribal nomenclature of the Rubiaceae.
Taxon 25: 595-610. 1976.
Dwyer, J. D. ee Fl. Panama. Ann. Missouri Bot. Gard. 67: 1-522. 1980.
Goprrey, R. K., & J. W. Wooten. Aquatic and wetland plants of southern United
States. Dicotyledons. x + 933 pp. Athens, Georgia. 1981. [Rubiaceae, 712, 714-
721,
HALLé, F. Etude biologique et morphologique de la tribu des Gardéniées (Rubiacées).
Mém. ORSTOM 22: 1-146. pis. 1-5. 1967.
Haypen, M. V. Systematic morphological study of New World rubiaceous — hae
bioideae sensu saeeeaTy Unpubl. Ph.D. thesis, St. Louis Univ. v pp.
[+ biography of baa 19
Hepper, F. N., & R. W. J. Keay. en In: F. N. Hepper, ed., Fl. W. Trop. Africa.
ed. 2.2 : 104-223. 1963.
Hom, T. Rubiaccse: anatomical studies of North American representatives of Cepha-
lanthus, pate Houstonia, Mitchella, Diodia, and Galium. Bot. Gaz. 43: 153-
186. pls. 7-9.
Jones, F. B. Flor | ae Texas Coastal Bend. xxxvi + 262 pp. Sinton, Texas. 1975.
(Randle Cephalanths Hedyotis, Galium, Borreria, Richardia, Diodia, Sperma-
coce, 188-192.]
Jones, S. B. Mis os flora. VI. Miscellaneous families. Castanea 41: 189-212. 1976.
“Rubiaceae, ae 1.]
KISAKUREK, M. V., A. ; M. LEEUWENBERG, & M. Hesse. A chemotaxonomic investi-
gation of . plant families of Apocynaceae, Loganiaceae, and Rubiaceae by their
indole alkaloid content. 7m: S. W. PELLETIER, ed., Alkaloids: chemical and biological
perspectives 1: 211-376. 1983.
Koek-Noorman, J. A contribution to the wood anatomy of the Cinchoneae, Copto-
sapelteae and Naucleeae (Rubiaceae). Acta Bot. Neerl. 19: 154-164. 1970.
. Hocewec. The wood anatomy of Vanguerieae, Cinchoneae, Condamineae,
and Rondeletieae uaneetaes Acta Bot. Neerl. 23: 627-653. 1974 [1975]. ead
photos of Pinckneya wood; four species of Exostema studied. ]
Krause, K. Uber esa enet Driisen an den Nebenblattern von Rubiaceen. Ber.
Deutsch. Bot. Ges. 27: 446-452. 1909.
Lewis, W. H. Cytopalynological study of African Hedyotideae (Rubiaceae). Ann. Mis-
souri Bot. Gard. 52: 182-211. 1965a.
. Type collections of African rubiaceous taxa at the Missouri Botanical Garden
Herbarium. /bid. 212, 213. 1965b. [Includes several of Hedyotis, sensu lato, and
Pentodon.]
Chromosome numbers cores ag 1. Ibid. 53: 100-103. 1966.
Lit TLe, E. L. Atlas of United States trees. Vol. 4. Minor eastern hardwoods. U.S. De
Agr. Forest Serv. Misc. Publ. 1342. v + 17 pp. + 3 base maps + 166 species maps
[+ 2 pp. indices]. 1977. [Cephalanthus, maps 32-NE, 32-SE, 32-N; Pinckneya, map
93.] Vol. 5. Florida. Jbid. 1361. vi + 22 pp. + 6 base maps + 256 species maps.
1978. els map 42; Exostema, map 202: Pinckneya, map
WORTH. Common trees of Puerto Rico and the Virgin Islands.
U.S. a Agr. Handb. 249. x + 548 pp. Washington, D. C. 1964. [Rubiaceae,
504-525.]
Lona, R. W., & O. Lakera. A flora of tropical Florida. new ed. xvii + 962 pp. Miami.
1976. iRubiacene. 792-809.]
Lunk, W. A. svar of West Virginia. Castanea 12: 27-38. 1947
Martin, A.C, e comparative adn ay Am. Midl. Nat. 36: 513-
660. 1946. nena 582, 592, 596-599, pls. 44, 45.]
Morton, J. F. Atlas of medicinal plants of ee a rica. Bahamas to Yucatan.
xxviii + 1420 pp. Springfield, Illinois. 1981. [Rubiaceae, 852-879.
PFEIFFER, L. Nomenclator botanicus. Vol. 2(1). 760 pp. Kassel, Germany. 1874.
1987] ROGERS, CINCHONOIDEAE 14]
Porcuer, F. P. Resources of the southern fields and forests, medical, economical and
agricultural. new ed. xv + 733 pp. Charleston, South Carolina. 1869. [Exostema,
Pinckneya, Cephalanthus, and other Rubiaceae, 442-445.]
Proctor, G. R. Flora of the Cayman Islands. ix [+ 4 maps] + 834 pp. London. 1984.
[Rubiaceae, 720-743.
RapForp, A. E., H. E. AHLES, & C. R. BELL. Guide to the vascular flora of the Carolinas,
with Here biion in the Southeastern States. Preface + 383 pp. Chapel Hill, North
Carolina. 1964. Leen oe 309.
RosBerTSON, C. Flowers and 1 . Lists of visitors of four hundred and fifty-three
flowers. 221 pp. Carlinville, nee 1928. [Cephalanthus, 175, long- and short-
tongued bees, other Hymenoptera, various Lepidoptera, Coleoptera, and Hemiptera;
Houstonia purpurea, 177, bees, flies, butterflies, beetles.
SARGENT, C. S. The silva of North America. Vol. 5. viii + 189 pp. pls. 198-251. Boston
and New York. 1893. [Exostema, Pinckneya, Guettarda, 103-114, pls. 226-229.]
SCHUMANN, K. Rubiaceae. Nat. Pflanzenfam. IV. 4: 1-156. 1891
ScoGGAN, H.J. The flora of Canada. Part 4— Eee mes (Loasaceae to Compositae).
Pp.
1117-1711. Ottawa. 1979. [Rubiaceae, 1407-1414]
SMALL, J. K. Manual of the southeastern flora. xxii + 1554 pp. New York. 1933.
(Reprinted Univ. N. ae pe Chapel Hill.) Rubee 19512 1269.]
Soukup, J. Las Rubiaceas del P sus géneros y lista de especies. Biota 9: 315-346,
377-398. 1973. [Includes Bieaandi 337; Exostema, 330.]
STANDLEY, P. C. Rubiaceae. N. Am. FI. 32: 3-300. 1918.
. Trees and shrubs of Mexico (Bignoniaceae—Asteraceae). Contr. U.S. Natl. Herb.
23: 1313-1721. 1926. [Rubiaceae, 1349-1394.]
. Rubiaceae. Fl. Yucatan. Publ. Field Mus. Bot. Ser. 3: 157-492. 1930.
—. The Rubiaceae of Venezuela. /hid. 7: 343-485. 1931.
—. Rubiaceae. FI. Peru. /bid. 13(6): 1-261 + index. 1936.
—. Rubiaceae. Fl. Costa Rica. /bid. 18: 1264-1380. 1938.
L. O. WituiAMs. Rubiaceae. Fl. Guatemala. Fieldiana Bot. 24(2, nos. 1-3):
1-274. 1975.
STEYERMARK, J. A. Flora of Missouri. Ixxxiii + 1728 pp. Ames, Iowa. 1963. [Rubiaceae,
—140
. The botany of a sa Highland. Part 9. Rubiaceae. Mem. New York Bot.
Gard 23: 227-832.
Rubiaceae. FI. ees, 1-2070 + errata. 1974.
TOMLINSON, P. B. The biology of trees native to tropical Florida. v + map + 480 pp.
Published by the author, Allston, Massachusetts. 1980. [Rubiaceae, 331-354; in-
cludes original data and detailed illustrations.]
VERDCOURT, B. Remarks on the classification of the Rubiaceae. Bull. Jard. Bot. Bruxelles
28: 209-290. 1958.
. Rubiaceae (part 1). Fl. Trop. E. Africa. 414 pp. + map. 1976. [Includes com-
ments on infrafamilial classification, with synopsis of subfamilies and tribes.
Vines, R. A. Trees, shrubs and woody vines of the Southwest. xii + 1104 pp. Austin,
Texas; and London. 1960. [Rubiaceae, 936-940.
WELLS, J. R., & A. J. SHARP. The Coffeoideae (Rubiaceae) of Tennessee. Jour. Tenn.
Acad. Sci. 41: 147-153. 1966.
WUNDERLIN, R. P. Guide to the vascular plants of central Florida. iv + 472 pp. Tampa,
St. Petersburg, Fort Myers, and Sarasota. 1982. [Rubiaceae, 344-349.]
KEY TO THE GENERA OF CINCHONOIDEAE IN THE
SOUTHEASTERN UNITED STATES
A. Plants herbs or pene subshrubs; raphides present; placentae peltate; fruits dry
and less than 0.5 cm
142 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
B. Flowers pentamerous; placentae bilobed apically; plants hygrophilous ee fleshy.
geese us shattie std uae gecasa yest ean aid hye dcatee 91 dea oh eae ey eee 3. Pentodon.
Bese se de ore tees et cae esse eas tay este etaarey aaa utente Hedyotis.
A. Plants shrubs or trees; raphides absent (except in Hamelia); placentae ia axile,
sometimes parietal (nearly peltate in E.xostema), or the ovules pendulous; fruits fleshy
and/or over 0.5 cm long.
C. Flowers and fruits in globose heads; locules of ovary uniovular. .............
Fade eh da etie Le en oes este ok du eee ae . Cephalanthus.
C. Flowers and fruits not in globose heads; locules of ovary usually multiovular
(Randia sometimes with only | seed in a fruit).
D. Plants armed with paired spines; leaves largely in fascicles clustered along
stems.
E. Flowers mostly tetramerous; aestivation of corolla valvate; stamens in-
fruit
D. Plants unarmed; leaves decussate, whorled, or in terminal clusters.
F. Fruits dehiscent; seeds winged: anthers exserted.
G. Calyx lobes more or less uniform; seeds vertical or nearly so; flowers
Ex
BONIAY cone Gave iy rota ces SG ee on enue deed pap ae ostema.
G. Some calyx lobes expanded into leaflike pink to white * “flags”: seeds
rizontal or oblique; flowers in compound cymes. ... 1. Pinckneya.
F. Fruits cae none seeds unwinged; feria eeleded or partly exserted.
H. Flowers perfect; corolla red or orange, lobes a small fraction of length
of tube; ovary usually 5-locular; plants pubescent; raphides present.
y
Seaside tab a Soe Seng ote ee eee a ea . Hamelia.
H. Flowers imperfect, plants dioecious; corolla white, lobes see
y unilocular (or apr £ nts
migetly glabrous; raphides absent. beep anes ree ataae 7. Casasia
Tribe CONDAMINEEAE Bentham & Hooker, Gen. Pl. 2: 8, 12. 1873.
1. Pinckneya A. Michaux, Fl. Bor. Am. 1: 103. p/. 73. 1803.
Shrubs to small trees, sometimes in colonies from root suckers. Leaves de-
ciduous, opposite, the blades lanceolate or ovate to usually nearly elliptic,
obtuse or rounded to caudate at the base, acuminate or less often acute at the
apex, lateral nerves usually rather arcuate-ascending, the petiole and midrib
often reddish (color fading in pressed specimens); stipules narrowly deltoid to
lanceolate with acuminate apices, acting as bud scales, caducous, bearing col-
leters adaxially toward the base; abaxial side of blades of young leaves and the
petioles, young stems, inflorescence axes, ovaries, calyces, and corollas usually
abundantly provided with variably kinked to straight and spreading or parallel-
appressed, tawny to almost white, incompletely septate and nonseptate, uni-
seriate trichomes; adaxial side of leaf blades often strigose to glabrate. Inflo-
rescence a pyramidal or hemispheric compound cyme with a straight central
axis, the lateral units sometimes repeating the form of the main axis, the
branching opposite or distal pedicels alternate; distal bracts linear or greatly
expanded to resemble the flaglike sepals, the basal bracts often intergrading
with foliage leaves. Flowers fundamentally pentamerous, nearly actinomorphic
1987] ROGERS, CINCHONOIDEAE 143
(except for the flaglike calyx lobes), fragrant. Calyx lobes briefly connate above
the ovary, the nonflaglike lobes ca. /4—¥4 the length of the corolla and subulate
or linear, or somewhat broadened toward the base, pink or partly green, in
certain flowers 1-3 (or all 5) calyx lobes clawed and with greatly expanded
blade(s) much exceeding the corolla in length and breadth, these resembling
foliage leaves in shape, but smaller and pink to white, then sometimes with
reddish borders. Corolla creamy or greenish yellow to pink, mottled with (pink
or) purple or brown, with a long, narrow, cylindrical or slightly flared tube and
(4 or) 5 (or 6) ligulate or narrowly elliptic, reflexed lobes about '4—'» the length
of the tube, the lobes imbricate or some valvate, with particularly coarse tri-
chomes within. Stamens exserted, the filiform filaments inserted near the base
of the tube in a pilose ring, anthers dorsifixed, sagittate, elliptic-oblong or
broadened below the middle; pollen grains tricolpate and reticulate (fide Verd-
court). Ovary surmounted by an epigynous disc, containing numerous ovules
arranged more or less in 2 ranks along an axile placenta in each locule; style
filiform, the stigma exserted and barely divided into 2 broad lobes. Capsules
persistent, slightly longer than broad to slightly broader than long, lightly com-
pressed perpendicular to the septum (this often appearing as a sunken vertical
line), predominantly loculicidal, speckled with lenticels, the endocarp made up
of light-colored fibrous cells, the apical perianth scar a broad ring around a
sunken center. Seeds waferlike, with a broad wing around the embryo (except
often at the hilum), wedge or fan shaped, the hilum opposite the broadest edge,
the surface area considerably less than cross-sectional area of the locule, stacked
horizontally or obliquely along a broadened placenta raised on a ridge running
nearly the entire length of the middle of the septum (ridge and placenta T-shaped
in transverse aspect, the seeds attached at various points across the head of
the T), surface of seeds reticulate from outlines of testa cells, these with re-
ticulate, straplike reinforcements on the outer walls. Embryo in a tough sa
(presumably endosperm), spatulate or with cotyledons very slightly sri,
the radicle about as long as cotyledons or shorter. Type species: Pinckne
bracteata (Bartram) Raf. (P. pubens Michx.). (Name commemorating cane
Charles Cotesworth Pinckney, 1746-1825, South Carolinian, veteran of the
American Revolutionary War, statesman, presidential candidate, and bene-
factor of André Michaux and his son Francois-André.)— GEORGIA BARK, FEVER
TREE, POSSUM POD.
A monotypic genus confined to the two southernmost counties of South
Carolina, the southern half of Georgia (including the Okefenokee Swamp), and
scattered localities in northeastern to northwestern Florida (several counties
from Nassau to Volusia, west to Gulf and Jackson), but not in the western
portion of the Florida Panhandle (see Little, 1977, for map). The distribution
lies mostly, but by no means overwhelmingly, in the Altamaha Grit region of
Georgia and is probably largely determined by edaphic factors.
Pinckneya is encountered in low, sandy, wet situations, especially at margins
of swamps, stream banks, and low spots in pine barrens. According to Taylor
and Uphof (independently?), it thrives best on river hummocks, where its trunk
is periodically submerged.
144 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
The flowers open sequentially (possibly rarely as early as late April) through
May and June (to July).
For explanation of the displacement of the well-known name Pinckneya
pubens Michx. by P. bracteata, consult Merrill and Wilbur.
The most salient characteristic of these shrubs or small trees is that on many
flowers one or more calyx lobes are expanded into large pink or sometimes
white “flags.” This occurs frequently, but sporadically, in the Rubiaceae, al-
though not in any of the other genera indigenous to our area. Kurz & Godfrey
remarked that it 1s “one of the most spectacularly beautiful [trees or shrubs]
occurring in northern Florida.” The less conspicuous, typically greenish yellow
corollas are marked with purple or brown and have TCHEACS, senate pu-
bescent lobes on the long tubes. The slightly flattened loculi ist
for long periods on the branches; upon opening they reveal qiimenbe wa-
ferlike seeds stacked horizontally in the two locules. Interpetiolar stipules with
abundant colleters on the adaxial side help to distinguish Pinckneya from
nonrubiaceous genera. The young stems, inflorescence axes, corollas, and some-
times foliage are typically conspicuously pubescent. Midribs of living leaves
tend to be reddish.
Most botanists place Pinckneya either in the tribe Condamineeae or in in-
frafamilial groups named differently but consistent with the same general circle
of affinity. Shared tribal or subfamilial characteristics include absence of raph-
ides, presence of endosperm in the seeds, incompletely septate uniseriate hairs,
mostly entire stipules, often “pitted” testa cells, woody habit, and—chiefly—
capsular fruits containing numerous horizontal seeds. While most members of
the tribe have valvate corolla lobes, an attribute sometimes ascribed to Pinck-
neya, | found the lobes to be imbricate or partly valvate in buds from the one
collection available for dissection.
Among the genera of the Condamineeae, Pogonopus Klotzsch emerges from
the literature as likely the closest relative for Pinckneya. Bentham & Hooker
erected the subtribe “Pinkneyeae” for the pair, and Baillon merged the two
genera. Their most conspicuous similarity, expanded flaglike sepals, is too
widespread in the Rubiaceae to stand as strong evidence for relationship, yet
Pinckneya and Pogonopus agree further in shape and size of corollas (the lobes
are reflexed in Pinckneya only), position of anthers and stigmas, shape of
capsules (although much smaller in Pogonopus), and indument. Their habit
and leaves are similar but do not set them apart from other arborescent Ru-
biaceae. Beyond the differences indicated parenthetically above, Pogonopus has
smaller seeds less drawn out marginally into wings and has stamens inserted
higher in the corolla tube, although the latter difference is hardly appreciable
when Pogonopus speciosus (Jacq.) K. Schum. is compared with Pinckneya. I
found the basal portion of the corolla tube of flowers of Pogonopus speciosus
and P. tubulosus (DC.) K. Schum. to be thickened into a woody cylinder, a
feature not found in Pinckneya. (See Oersted for an illustrated floral dissection
of P. speciosus, as P. exsertus.) In contrast with authors who list internally
glabrous corolla lobes in Pogonopus as a distinction from Pinckneya, I en-
countered internally pubescent lobes in both genera
Koek-Noorman & Hogeweg, in an investigation of wood anatomy of the
1987] ROGERS, CINCHONOIDEAE 145
Condamineeae, evidently perceived no particular connection between Pinck-
neya and Pogonopus. They called Pinckneya “exceptional” among its relatives
in having semi-ring-porous wood with tangential pore chains and concentric
parenchyma bands. (At least the first of these exceptional features is probably
due to the temperate distribution of the genus, which is in itself very unusual
among woody Rubiaceae.)
A second possible close relative is the newly described monotypic Brazilian
genus Kerianthera Kirkbride. Kirkbride held the new genus to be most similar
phenetically within the Condamineeae to Pinckneya. He listed their shared
features as foliar calyx lobes, dense pubescence on the inner faces of the corolla
lobes, and winged seeds but peed Kerianthera om both Pogonopus and
Pinckneya by its ““4-merous calyx, 7-8 rous corolla ting from
the apex of the corolla tube, anthers with approximately 300 locelli, septicidal
capsules, and seeds irregularly biwinged” (p. 109).
It is doubtful that frequent mention of Pinckneya in old botanical-medical
literature as a remedy for malaria has any meaningful basis. Cornatzer and
colleagues related secondhand that pharmaceutical tests on extracts from Pinck-
neya revealed no antimalarial effects on infected canaries. Application of Pinck-
neya against malaria probably grew out of the perception of its relationship to
Cinchona L., the source of the familiar antimalarial alkaloid quinine. Whether
alkaloids form in Pinckneya remains a debatable question: Sumerford and
Naudain tried and failed to detect any, but Wall and colleagues indicated the
presence of at least one unnamed alkaloid. Further work is desirable.
REFERENCES:
Under subfamily references see BAILLON; BENTHAM & HOOKER; KOEK-NOORMAN &
Hocewec; LitTLe (1977); SCHUMANN; and VERDCOURT (1958).
ANONYMOUS. ee pubens Michx. Natl. Hort. Mag. 29: 184, 185. 1950. [Flowered
in age , D. C.; includes horticultural and descriptive notes and photo of
er.]
aes J. J. The birds of America. xii pp. + 500 pls. + ee xill-xxvi. New York.
1937 (originally published 1827-1830). ee Divs
BARTRAM, W. Travels through North & South Carolina, eae East & West Florida.
xXxxiv + 522 pp. 1791. [Bignonia bracteata, 16, 468.
CLARK, R. C. Woody plants of Alabama. Ann. pe a Gard. 58: 99-242. 1971.
[Pinckneya absent, despite a close ee in Geo
CORNATZER, W. E., M. M. McEwen, & J. C. ANDREWS. Schizonticidal tests on Rauwolfia
ee and some other proposed antimalarial plants. Jour. Elisha Mitchell Sci.
c. 60: 167-170. 1944. [Pinckneya, 170.]
eee W.H. Preliminary reports on the flora of Georgia. 2. Distribution of 87 trees.
Am. Midl. Nat. 43: 742-761. 1950. [Pinckneya, 749, 750, 761 (map).
Harper, F. Two more available plant names of William Bartram. Bartoni , 8.
1942. [Pinckneya bracteata incorrectly published here as a new Combination (cf
MERRILL, WILBUR).
Harper, P. A rare small tree—Pinckneya pubens. Jour. Roy. Hort. Soc. 102: 222. 1977.
[Includes color photograph of inflorescence, descriptive notes, and habitat notes;
mentions “‘pure white form” and hardiness in zone 8.]
Harper, R. M. A phytogeographical sketch of the Altamaha Grit region of the coastal
plain of Georgia. Frontisp. + 414 pp. + 28 pls. 1906. Repaged reprint from Ann.
146 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
New York Acad. Sci. 17: 1-414. 1906. [Pinckneya, 63, 65, 91, 156, 192, 209, 329,
332.]
KiRKBRIDE, J. H., Jk. Manipulus Rubiacearum IV. Kerianthera (Rubiaceae), a new genus
from Amazonian Brazil. Brittonia 37: 109-116. 1985. [Includes distribution of foliar
sepals in Rubiaceae. ]
Kurz, H., & R. K. Goprrey. Trees of northern Florida. xxxiv + 311 pp. Gainesville,
Florida. 1962. [Pinckneya, 286-288.]
Lawrence, E. Pinckneya pubens. Am. Hort. Mag. 40: 232, 233. 1961. [Includes hor-
ticultural and descriptive notes and common names.
Litt.e, E. L., Jk. Rare and local trees in the national forests. U. S. Dep. Agr. Forest
Serv. Conserv. Res. 7 21. ii + 14 pp. 1977. [Pinckneya, 4.]
MELLINGER, M. B. Some plant associations of Pinckneya pubens. Castanea 31: 310-
313. 1966 [1967]. Visited ten localities in Georgia and South Carolina; for associated
plants see TAYLOR, UPHOF.
MERRILL, E. D. In defense of the validity of William Bartram’s binomials. Bartonia 23:
10-35. 1945. [P. bracteata (Bartram) Raf. (P. pubens Michx.), 23, 24; includes
nomenclatural history.
MicHAux, F. A. Georgia bark. The North American sylva. Vol. 1. Pp. 260-262. pil. 49.
Paris. 1819. [Includes color plate, origin SE ean name, and manner of use.]
ida; includes descriptive notes, common names, “ane black-and-white photographs
showing flowers and habit (see frontisp.).
Naupain, E. H. Pinckneva pubens, Michaux (Georgia bark). Am. Jour. Pharm. 57: 161-
163. 1885. [Chemical tests yielded a suspected glucoside, “pinckneyin,” but no
alkaloids. ]
oe A.S. L’Amérique centrale. iii + 18 pp. map + 3 pls. + 18 pls. Copenhagen.
3. [Pogonopus exsertus” (P. speciosus), 17, pl. 13.
Re! C.S. Pinckneya bracteata. Casket 1827: 193 (fig.), 194. 1827.
SUMERFORD, W. T. A note on Pinckneya pubens (Michaux). Jour. Am. Pharm. Assoc.
Sci. Ed. 32: 101, 102. 1943. [Alkaloids not found; suggests one artifact that may
have caused erroneous reports of alkaloids in Pinckneya |
_— E. B. The Georgia bark or quinine tree (Pinckneya pubens). Pl. World 9: 39-
1906. [Includes notes on habitat, appearance, origin of name, medicinal use,
a associates (see UPHOF, MELLINGER).
Upnor, J. C. T. Pinckneya pubens Rich. Mitt. Deutsch. Dendrol. Ges. 49: 1-4. 1937.
[Includes history, origin of name, habit, habitat, medicinal use, flowering time,
description, illustration, and pea plants (see omen eae OR).
Wa Lt, M. E., C. S. FENSKE, J. W. Garvin, J. J. WILLAMAN, Q. Jones, B. G. SCHUBERT,
& H.S. Gentry. Steroidal oe LV. Survey of plants an nena sapogenins
and other constituents. Jour. Am. Pharm. Assoc. Sci. Ed. 48: 695-722. 1959. [Pinck-
neya, 718; alkaloid(s) in leaf, stem, and fruit.
Wixbsur, R. L. A reconsideration of Bartram’s binomials. Jour. Elisha Mitchell Sci. Soc.
87: 562 73. 1971. [P. bracteata (Bartram) Raf., 70, 71; includes nomenclatural his-
tory. ]
WriGutT, A. H., & A. A. Wricut. The sets and composition of the vegetation of
Okefinokee Swamp, Georgia. Ecol. nogr. 2: 110-232. 1932. [Pinckneya, 137,
138, 150, 169, 194; along St. Mary’s es and in cypress bays, cypress ponds, and
moist pine barrens.]
Tribe HEDYoTIDEAE DC. Prodromus 4: 342, 401. 1830.
2. Hedyotis Linnaeus, Sp. Pl. 1: 101. 1753; Gen. Pl. ed. 5. 44. 1754.
Annual or perennial, delicate to coarse, prostrate to stiffly erect herbs or
weak subshrubs [or shrubs], highly variable in habit, sometimes rosette forming,
1987] ROGERS, CINCHONOIDEAE 147
with | or few delicate ascending axes, these (infrequently) unbranched to (fre-
quently) highly branched throughout, or extensively branched at base and
scoparioid, axillary growth strongly developed and often overtopping terminal
growth, the branching frequently widely divergent and symmetrical. Stems
winged or angled, often square, occasionally with adventitious roots when
procumbent. Roots thick and woody or fasciculate. Plants usually with con-
spicuous raphide bundles, and with stems, leaves, and calyces pilose to glabrous.
Leaves petiolate or sessile, opposite [or fasciculate or whorled], (frequently)
nearly linear to (infrequently) broader than long, commonly more or less nar-
rowly elliptic, entire or scabrous around the margins, infrequently cordate
basally; stipules interpetiolar, membranaceous, emarginate or bilobed to del-
toid or rounded, or frequently fimbriate, with multicellular glandular heads
either adaxial or marginal. Flowers on long, threadlike peduncles or pedicels
to sessile, terminal or axillary, solitary or, more often, in fundamentally cymose
but highly variable inflorescences, these (usually) compound dichasial, some-
times simple dichasial or partly monochasial, lax and uncrowded to fasciculate,
then sometimes tightly clustered into hemispheric heads or pseudoumbellate,
flowering axes often between pseudodichotomous branches or forming pseu-
dodichotomies with other axes. Flowers tetramerous, homostylous, hetero-
stylous, or cleistogamous. Calyx lobes separate to top of ovary or briefly connate,
usually deltoid or elliptic to subulate, exceptionally with claw and limb. Corolla
white or greenish, or blue with a yellow or reddish eye, or pink, or variably
purplish, extremely variable in length, usually pubescent within, the tube ob-
solete or very nearly so to several times longer than calyx, abruptly expanded
at the level of the anthers or not expanded; in species with well-developed
corolla tubes the corolla most often salverform to funnelform or sometimes
obconical, the lobes ca. 4 as long as tube to much longer, spreading or erect,
variable in shape. Anthers included or exserted, sessile or on epipetalous fil-
aments, fusiform to orbicular, dorsifixed; pollen grains 3- or 4-colporate, re-
ticulate. Ovary inferior, each of the 2 locules with a peltate placenta bearing
numerous reportedly hemianatropous or anatropous ovules; stigmatic lobes 2,
included or exserted, long and threadlike to short and stubby, nearly sessile or
on a long, filiform style. Fruit a capsule usually compressed perpendicular to
the generally sunken septum, much broader than long to cuneiform, often
apically emarginate, inferior to almost superior, usually conspicuously belted
by calyx sinuses and/or corolla scar, adorned with persistent calyx lobes, pri-
marily loculicidally dehiscent but not rarely also septicidal:; dehiscence usually
restricted to the apex (but sometimes indehiscent); seeds numerous, minute,
rugose to fairly smooth, dark, subglobose to angular or flattened, containing
initially nuclear [or exceptionally cellular] endosperm. Megagametophyte (em-
bryo sac) of the Polygonum type. (Including Oldenlandia L., Houstonia L.)
Lectotype species: H. Auricularia L. (discussion in text). (Name from Greek,
hedys, sweet, and otos, ear, in reference to habit of plants; see Linnaeus, Phi-
losophia Bot. 179. 1751
A vaguely circumscribed, polymorphic genus, possibly with 400 species when
defined broadly, almost worldwide in warm regions and with extensions into
temperate areas, although nearly absent from Europe and the Soviet Union:
148 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
present in Australia, Asia (including Japan and the Malay Archipelago), the
Middle East (very poorly represented), almost the entire length of Africa, and
the Americas from central Argentina to southern Canada. Roughly 60 species
occur in the New World, about 50 of them on mainland North America and
approximately 30 in the continental United States, with about two-thirds of
these reaching the range of the Generic Flora. Most North American species
belong to the group often recognized as the genus Houstonia, and with a few
exceptions, the West Indian and Central and South American species belong
to the group often recognized as the genus Olden/andia.
The interrelationships and taxonomic status of Hedyotis, Houstonia, Ol-
denlandia, and a number of additional extralimital genera have been contro-
versial for centuries and remain inadequately investigated, especially from a
worldwide perspective. The disparate circumscriptions and diagnostic char-
acteristics given by different authors cloud the usage of all three names and
make it impossible to characterize the segregate genera crisply. The following
sketch comes from the literature (see especially Gray, 1860; Lewis, 1961). It
must be stressed that the validity of the distinctions changes with the varying
concepts of the groups, that much of the variation is continuous, that most of
the distinctions rest upon inadequate sampling, and that exceptions and overlap
abound.
Oldenlandia sensu stricto is variously estimated to have from 80 to around
300 species, depending on its delimitation when recognized as a genus. Its
distribution is almost worldwide in warm regions; it is best represented in the
Old World tropics, with a center of diversity in Africa (see Bremekamp, 1952,
for a revision of African species; also see Lewis, 1965, under subfamily ref-
erences). About 15 species are distributed in America from the southern limit
given above for Hedyotis to New York (//. uniflora (L.) Lam.). Hedyotis co-
rymbosa (L.) Lam., H. lancifolia Schum., and H. herbacea L. are Old World
species reported as weeds scattered in the American tropics. No fewer than
three endemic species have been named from Cuba (see Alain). Five or six
species (listed below) are found in the continental United States, all of them
reaching the area of the Generic Flora.
Tendencies toward a slender, herbaceous habit, narrow leaf blades, ho-
mostylous flowers (for a list of 39 exceptions, see Bahadur, 1963), short corolla
tubes, hemispheric placentae partitioned and sessile or inconspicuously stalked
from the center of the septum (vs. placentae of irregular shape and stalked from
base of septum in other species of Hedyotis, according to Hayden), completely
inferior ovaries, thin, loculicidal capsules, and numerous tiny, angled or nearly
spherical seeds lacking hilar ridges and containing fleshy endosperm have been
set forth as distinctive features of O/denlandia. (Hayden (p. 21) rejected the
endosperm character as ““completely useless.”
Houstonia comprises about 40 species nearly limited to North America; a
few of them are rare and possibly introduced in the West Indies, and //.
serpyllacea Schlecht. thrives in Guatemala. Roughly half the species reach the
continental United States, and slightly over half of these occur in the area of
the Generic Flora. The others are confined to the Southwestern States. Three
species extend from the Southeast as far north as southern Canada, with the
1987] ROGERS, CINCHONOIDEAE 149
natural northern limit being about 54 degrees north latitude (see Scoggan).
North of our range, Carr described from southwestern Virginia Houstonia
setiscaphia, which Terrell (1959; also see Uttal) reduced to synonymy with
oustonia canadensis Willd. ex Roemer & Schultes (Hedyotis canadensis (Willd.
ex Roemer & Schultes) Fosb.).
Species of Houstonia tend to have an herbaceous habit, comparatively wide
leaf blades, heterostylous flowers, long corolla tubes, partly superior, fairly thin,
loculicidal capsules, and relatively few, large seeds flattened parallel to the
placenta, these concave toward their peltate attachments, often with hilar ridges,
and containing corneous endosperm. Fosberg (1941, 1954), Fosberg & Terrell,
Greenman, Lewis (most papers cited here), Lewis & Terrell, Shinners (1949),
Standley (1918), Terrell (most cited papers), Terrell and colleagues, and Yelton,
among others, have studied the taxonomy and related aspects of Houstonia.
Potentially of interest in connection with the relationship between Houstonia
and Oldenlandia, the two studied species of Houstonia have “‘naked” or ‘“‘un-
differentiated” ovules not showing an obvious integument separated from a
nucellus. Homologies of the exposed layer are not certain (cf. Lloyd; Fagerlind;
Roth & Lindorf). Numerous sources (Fagerlind; Siddiqui & Siddiqui; Farooq,
1953, 1958; Farooq & Inamuddin; Raghavan & Rangaswamy; Rao & Babu;
Shivaramaiah & Rajan; Shivaramaiah & Rao), on the other hand, agree that
species of O/denlandia have ovules with one integument and a reduced nucellus
of one or a few cells. More study in Houstonia is needed before the difference
can be given much taxonomic weight.
Hedyotis sensu stricto, comprising over a hundred species restricted to warm
Asia, 1s ordinarily more woody and shrubby than the two preceding “‘potential”
genera. Additional characteristics are fimbriate stipular lobes, axillary inflo-
rescences, short corollas, sometimes hard, thick, indehiscent or septicidal fruits,
and variably shaped (but not concave) seeds. Sinuses between the persistent
calyx lobes on the capsules have been said to be narrower than in O/denlandia.
The principal proponent of maintaining all three genera as distinct is Terrell,
whose conclusions (1975b) are given credence by his study of a broad spectrum
of herbarium specimens, mostly from the New World. He pointed out that
Oldenlandia and Houstonia differ in base chromosome numbers, except in
morphologically divergent species. His comparison of type species of the three
groups does demonstrate a level of variation consistent with the recognition
of three genera but leaves the question of intermediates untouched. (Note, as
explained below, that Terrell and I accept different lectotype species for Hedy-
otis.) Subdividing the assemblage into three or more genera requires a will-
ingness to draw rather arbitrary lines to break up a large, awkward, hetero-
geneous assemblage. Verdcourt (1976) indicated that the cumbersome nature
of the complex and its heterogeneity justified partitioning it into multiple
genera.
With some trepidation I interpret the case for a broad view of Hedyotis as
slightly more convincing. In 1961 Lewis (p. 221) concluded with detailed
documentation that “‘no character currently in use”’ distinguishes Houstonia
from Oldenlandia and added that admittedly incomplete cytological evidence
favors the union. His efforts focused chiefly on American species, and he
150 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
appears to have had mixed feelings about the status of species from the Old
World (see pp. 217 and 221 (footnote)). By incorporating the lectotype species
of Oldenlandia, Hedyotis corymbosa, under Hedyotis, he made it necessary to
regard Oldenlandia as a name in synonymy, although later (1964) he separated
O. corymbosa from Hedyotis and recognized Oldenlandia as a genus. Along
with Terrell and others, he coauthored a paper in 1986 explicitly holding
Oldenlandia to be distinct (but see p. 113 for doubts).
Lewis cited palynological evidence in 1965 to support joining Houstonia
with Hedyotis. a Fosberg (1937, 1941, 1943b, 1954; Fosberg & Terrell),
stressing that the differences are weak and/or break down, followed by Shinners
(1949), has maintained that Oldenlandia, Houstonia, and Hedyotis are insul-
ficiently distinct to stand separately, a position that I find especially convincing
in view of the geographic breadth of the sampling that stands behind it. McVaugh
(p. 160) dismissed the differences between Hedyotis and Houstonia as evidently
“largely traditional rather than morphological.”
Since all three generic names have equal priority, the name to be adopted
for the genus encompassing the trio depends on the choices made by the earliest
authors to unite them. Lamarck selected Hedyotis over Oldenlandia in 1792,
and Kunth likewise chose Hedyotis in 1820 upon placing Houstonia in syn-
ymy.
Encircled by a crowd of potentially separate genera, mostly from the Old
World, Hedyotis is not a sharply defined unit, even containing both of our
potential segregates, and cannot be readily characterized in a universally ac-
ceptable manner. Fosberg (1943b) listed the attributes of the genus taken broad-
ly. The following enumeration of characters 1s based mostly upon Fosberg’s.
Hedyotis sensu lato has tetramerous flowers with valvate corollas and equal
calyx lobes: stigmatic lobes or branches receptive ventrally; expanded, fleshy,
peltate placentae; and capsular or dry indehiscent fruits moderately flattened
and with sclerified endocarps. The numerous seeds are often inserted peltately
or are taller than broad and are neither imbricate nor horizontal. They lack
lateral wings, except for thin edges at the angles. For a discussion of the position
of Oldenlandia among its African relatives, see Bremekamp (1952).
Hedyotis and Pentodon are our representatives of the sizable tribe Hedyoti-
deae (for comparison see Pentodon). Bremekamp (1966) and Verdcourt (1976)
differed in their characterizations of the tribe, although they agreed that mem-
bers usually have bilocular ovaries containing numerous ovules. Bremekamp
further characterized the tribe as having valvate corolla lobes, peltate placentae
inserted at the middle of the septum (Verdcourt said at the base), relatively
thin testa cells, and nonconnivant anthers opening by slits. Verdcourt included
capsular fruits. (See introduction for remarks on the position of the Hedyoti-
deae.
A handful of species in our area and several others from outside of it have
been included in Anotis DC. (or Anotis auct.), which Lewis (1966b) determined
to be an unnatural assemblage containing American species better placed in
Hedyotis.
In 1962 and 1965 Lewis developed a phylogenetic hypothesis for five in-
formal subgroups of subg. Houstonia in North America, taking into consid-
1987] ROGERS, CINCHONOIDEAE es
eration chromosome numbers, apertural fine structure in pollen grains, distri-
butions, and relative levels of advancement as judged from morphological
characters. Soon thereafter, Hayden added characters from seed coats. The
trunk of Lewis’s phylogenetic tree (1965, p. 263) culminates in ‘“‘“Group 2,”
having the base chromosome number of x = 11, a widespread number among
Rubiaceae, and thus thought likely to have remained unchanged from the
original stock of the subgenus. “‘“Group 2” is confined to southwestern North
America, the most likely port of entry and hub of radiation from the American
ropics.
Lewis (1962) attributed the level of morphological specialization lower than
that of “Group 2” to “Group 1,” hypothetically isolated by ancient climatic
changes to Baja California, an area possibly ‘‘not requiring major adaptations”
(1962, p. 864). He went on in 1965 to interpret the pollen of “Group 1” as
likewise least specialized and probably relictually similar to pollen in other
subgenera of Hedyotis and other genera of Hedyotideae. If Lewis is correct,
the base chromosome number of x = 13 in “Group 1” reflects an aneuploid
climb from the ancestral x= 11.
An apparent d d series along with presumed morphological
and palynological specialization i in the species toward the end of the series led
Lewis to derive “Group 3” (x = 11-9), found in the United States and Mexico,
from the stem of “Group 2,” and “Group 5” (x = 7, 8) from the stem of
“Group 3.” At first glance, the eastern North American “Group 4” might be
assumed to be closely related to ““Group 5” since the base chromosome number
of x = 6 (as counted by Lewis) in “Group 4” suggests the next step of the
descending aneuploid sequence, but the seemingly unspecialized gross mor-
phology, seeds, and comparatively large chromosomes observed in ““Group 4”
contradict such a position. In 1965, Lewis used pollen structure to link ‘““Group
4” to “Group 3,” and IJ infer support for this from Hayden
In 1986 Terrell, Lewis, Robinson, & Nowicke reevaluated species relation-
ships within Houstonia, using mostly characters from seed morphology, chro-
mosome numbers, and pollen (with special attention to ora). They set up a
dozen “‘species-groups,”’ seven of which consist of only one or two species. The
others correspond roughly to Lewis’s groups |-5, although there were several
differences in membership, and the authors of the 1986 paper did not formally
connect the new groups with the old. They did conclude that the new groups,
except for the intermediate “7. nigricans group,” fall into two “‘basic series.”
To paraphrase their summary, one series (not a formal nomenclatural series)
has a haploid chromosome number of 7 = 13 or more (vs. n = 11 or less),
ellipsoid or sublenticular noncrateriform (vs. crateriform) seeds, and colporate
pollen with the nexine merely thin in the equatorial portion of the aperture
(vs. grains colporate or the ora with thickened margins). They deferred making
taxonomic changes until more data were gathered.
Examining 116 collections from the Hedyotis purpurea and H. caerulea
“groups,” Lewis & Terrell came across frequent intraspecific euploid variation
in ploidy level but very little intraspecific aneuploidy. In two species the poly-
ploids were separated geographically from the diploids and appeared to be
colonizers—no marked geographic separation between the ploidy levels was
152 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
detected in the remaining species. The authors could not distinguish individuals
with different ploidy levels morphologically, which led them to attribute mul-
tiplication of chromosome sets to autoploidy rather than alloploidy, even though
meiosis was mostly normal. Variability in chromosome number seemed to be
connected with heterostyly and a perennial habit.
Divergent generic concepts have contributed to the profusion of names of
species and infraspecific taxa recorded as occurring in the range of the Generic
Flora. Beyond the problem of species and their varieties appearing under mul-
tiple generic names, botanists have achieved so little agreement concerning
ranks, definitions, and names of taxa in our area that the most recent revision
covering our species (by Standley, 1918) is obsolete, and subsequent sources
disconcertingly contradictory. Therefore, a complete list of the species in the
Southeast is currently impossible. The summary that follows rests heavily on
the work of Fosberg, Lewis, and Terrell. (It is based entirely on literature —I
have conducted no comparative study at the species level.) Full synonymy and
consideration of questionable species lie beyond the scope of the present effort.
Subgenus OLDENLANDIA (L.) Fosb. (not accepted here as validly published
by Torrey & Gray) includes in our area Hedyotis Boscii DC., n = 18; H.
callitrichoides (Griseb.) Lewis, n = 11, also in Africa, probably as an intro-
duction from the New World tropics; H. corymbosa, n= 9, 18,27; H. Salzmanii
(DC.) Steudel (Oldenlandia thesiifolia (St.-Hil.) K. Schum., introduced from
South America; see Fosberg & Terrell), 7 = 15; and H. uniflora (including H.
fasciculata Bertol. or not), n = 18, 36.
Subgenus HoustoniA (L.) A. Gray (Man. ed. 1. 180. 1848, see Brizicky)
(subg. Edrisia (Raf.) Lewis*) corresponds to Houstonia, if recognized at the
generic level, and as discussed above, has been broken down into informal
subgroups.
“Group 3” in subg. Houstonia is represented by H. nigricans (Lam.) Fosb.
(Houstonia angustifolia Michx.;, see Fosberg, 1954, and Long & Lakela), n =
9 (10
Subgenus Houston, Group 4, is the Hedyotis or Houstonia purpurea “group”
revised by Terrell (1959), who remarked on a high percentage of intergradation
and geographic variation involving every species. Terrell suspected hybridiza-
tion and introgression to have played significant roles in producing the pattern
of variation; pairs of species seemed to interbreed at some places but not at
others. In connection with the probable hybridization, it is of interest to note
that Lewis (1962) encountered almost uniformly normal meiosis in his cyto-
logical survey of the genus in North America, and Fosberg (1943b, p. 15)
described hybridization as “‘little evident” among Hawaiian saa despite
‘tremendous evolutionary activity.” Most species of the H. purpurea group
have polyploid races in addition to diploids (Lewis & Terrell). Terrell took a
sUpon publishing subgenera in Houstonia, Rafinesque (Ann. Gén. Sci. Phys. 5: 225 (13 in reprint).
1820) automatically created subg. Houstonia, which he called Houstonia subg. Edrisia. By ICBN
Article 57.3, the combination in Hedyotis formed by merging the original subg. Houstonia and
Rafinesque’s other subgenera into one subgenus must be called by the generic name, not subg. Edrisia
(Raf.) Lewis (Am. Jour. Bot. 49: 858. 1962).
1987] ROGERS, CINCHONOIDEAE 153
relatively narrow view in recognizing four species as opposed to Fosberg’s
(1954) placement of the entire complex in H. purpurea (L.) Torrey & Gray.
Whether or not most components of the complex should be treated as varieties
of H. purpurea or as distinct species, our representatives can be listed as follows:
Hedyotis purpurea (including or not Houstonia montana Small; cf. Yelton;
Terrell, 1978; Kral), n = 6, 12; H. longifolia (Gaertner) Hooker (including or
not Hedyotis Nuttalliana Fosb. = Houstonia tenuifolia Nutt.; see especially
Smith; the latter accepted as a species by Terrell in 1959), n = 6, 12; H.
canadensis, n = 6, 12; and H. ouachitana E. B. Smith (here presumed to belong
to “Group 4’’)
“Group 5” is represented by Hedyotis australis Lewis & Moore (Houstonia
micrantha (Shinners) Terrell; see Terrell, 1975a; Lewis & Moore), n = 16; H.
caerulea (L.) Hooker (including or not Hedyotis crassifolia Raf. = Houstonia
pusilla Schoepf and Houstonia patens Ell., according to Lewis & Moore, n =
8, 9, 16, 24 (but see Love & Love for reservations); H. Michauxii Fosb. (Hous-
tonia serpyllifolia Michx.), n = 16, 24; H. procumbens (J. F. Gmelin) Fosb.,
= 14 (see Gaddy & Rayner); and H. rosea Raf., n = 7 (see J. E. Moore; Taylor
& Taylor; Waterfall).
Seeds of Hedyotis corymbosa have been the subject of a series of studies (see
Corbineau & Céme for an entry to the literature). While the physiological
results are outside the scope of the present paper, a few salient ecological
discoveries deserve mention. The seeds are dimorphic in that for germination
some are “dormant” and require stratification while others do not. Artificial
selection led to ne lines of plants, one of which produces seeds showing no
need for stratification. The other produces a mix of the two types of seeds,
with the percentage of “dormant” seeds increasing as the season progresses.
All demand warm temperatures and must be activated by exposure to light,
although (at least in those not requiring stratification) the effects of light are
digi with a aaa of para tS ‘Dormant” seeds are strongly inhibited
fro of oxygen as high as that in the atmosphere,
eee after a cee period of stratification.
The citation of a lectotype for Hedyotis still requires choosing between al-
ternatives. Of three species comprising the genus in Linnaeus’s Species Plan-
tarum, H. herbacea can be eliminated from consideration first. Although it
dates back, along with H. Auricularia and H. fruticosa, to the year Linnaeus
first published Hedyotis, it is missing from one of the two generic treatments
appearing that year (in 1747a but not 1747b). For this reason and also since
authors (see Bremekamp, 1939, 1952) have removed it to Oldenlandia (see
ICBN T.4.e), since it was least known to Linnaeus, and since two different
lectotype species have already been proposed, it is unsuitable as a choice. Ruling
out H. herbacea has never provoked disagreement—the problem lies in settling
on one member of the remaining pair.
As background for discussing the conflict, it is worthwhile to note that Lin-
naeus’s description of Hedyotis is repeated essentially verbatim in all Linnaean
publications cited in the present context, including the laturally decisive
fifth edition of the Genera Plantarum.
The best choice for lectotype does not shine forth from recognition of Lin-
154 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
naeus’s frequent practice of basing generic descriptions on single species. Both
potential lectotype species were well known to Linnaeus from literature and
specimens when he wrote the generic description, and examination of the works
he cited reveals neither species as focal. (The only source I have not examined
is ‘‘Marlow. obs.,” cited more extensively by Dale and probably the ““Marloe”
discussed by Jackson.)
Nor is a single species revealed as central by Bremekamp’s (1939, 1952)
selection of Hedyotis fruticosa as lectotype, chiefly on the grounds that it, but
not H. Auricularia, agrees with the generic description in having dehiscent
fruits. (He pulled H. Auricularia out of Hedyotis as type species of his new
genus E-xallage in 1952.) Dehiscence, however, could not have entered the
generic description via H. fruticosa, about which Linnaeus (1747a, p. 26, no.
63) admitted, ‘“De fructu nulla nobis certitudo.”
Fruits of Hedyotis Auricularia were described (although with no mention of
dehiscence) in works Linnaeus cited (e.g., Burman). Bremekamp (1939) himself
suggested quite plausibly that Linnaeus’s failure to register fruits of H. Auric-
ularia as indehiscent could have resulted from misinterpretation of them as
immature, assuming their presence on the original specimens.
That Hedyotis Auricularia deviates from the generic description in this pos-
sibly minor character does not show the description to rest on H. fruticosa:
the information in the generic description that is at odds with H. Auricularia
did not originate with H. fruticosa, and Bremekamp did not show H. fruticosa
to match the generic description better. Bremekamp’s case, then, 1s based
mostly on an error and is incomplete. As explained below, I reject his supple-
mentary contention that Blume rendered H. Auricularia “illegitimate” as lec-
totype in 1826 by placing what Bremekamp regarded as a synonym under the
generic name Metabolos Blume. Bullock and Terrell (1975b) accepted Bre-
mekamp’s lectotypification.
The 1983 International Code of Botanical Nomenclature (Art. 8.1) rules that
the first lectotype chosen can be unseated only if demonstrated to be “‘in serious
conflict with the protologue.”’ If it is agreed that Hedyotis Auricularia has not
been thus exposed, it cannot be displaced (even if it was placed in Metabolos
under a different name), having been cited twice as typifying the genus before
er-Arnott), although it can be objected that the early use of “‘typus” is not
equivalent to the modern designation of a lectotype. That, however, may be a
moot objection, since Hitchcock & Green selected H. Auricularia as “standard
species” a century later but still ahead of Bremekamp.
In the interest of future investigations, 1t may be useful to stress that the
large number of species of Hedyotis in the broadly stated type locality for both
potential lectotypes, Sri Lanka, intensifies the hazard of working with incor-
rectly identified specimens. Types are presumably in the Hermann herbarium
at BM (see Trimen). Several specimens of Hedyotis, including one labeled H.
Auricularia by Linnaeus and another labeled H. fruticosa, are in the Linnean
Herbarium. The latter disagrees with the foliis /anceolatis Linnaeus attributed
to H. fruticosa in the Species Plantarum, for it has broad, mostly ovate leaf
1987] ROGERS, CINCHONOIDEAE 55
blades. (According to Stearn (p. 94), Linnaeus applied “‘lanceolatus” to blades
“oblong, but gradually tapering towards each extremity and terminating in a
point, the greatest width being at the middle, not below” (also see p. 91,
fig. 6).)
Preparations from species of Hedyotis sensu lato serve as folk remedies
around the world. Oldenlandia affinis (Roemer & Schultes) DC. (Hedyotis
a Roemer & Sehuites), which i is given to hasten childbirth 1 in Africa, con-
nst t and two oxytocic proteins. Practical
ae usage 1S hampered by the toxicity of serotonin and at least one of the
proteins, and both compounds are ineffective when administered orally to
laboratory animals (Gran, 1973a, b, d). Topical uses for oldenlandias are com-
mon and could, at least in some cases, as exemplified by Hedyotis diffusa Willd.,
be related to the presence of antiinflammatory iridoids.
The red dye “Indian madder” or “‘chay-root” from the commercially cul-
tivated Oldenlandia umbellata L. colors turbans and other products in India.
Extracts from this species are also used in treating tuberculosis. Roots of Hedy-
otis corymbosa yield the green (after chemical treatment) dye gerancine, and
bark from roots of H. herbacea, as well as leaves from H. scandens Roxb.,
likewise color fabrics. Capsules from H. scandens have been used to blacken
teeth.
Leaves of Hedyotis Auricularia, H. scandens, and H. nitida Wight & Arnott
are eaten in Asia. Hedyotis fruticosa 1s a minor source of wooden rods. For
further information on Hedyotis as a medicine and on its other uses, see Datta
& Sen, Lin et a/., Morton, Sastri et a/., Simmonds, and Usher.
LI UAY
REFERENCES:
Under subfamily references see ALAIN; BAILLON; BENTHAM & HOOKER; BREMEKAMP
(1952, 1966); Brizicky; DE CANDOLLE; HAYDEN; HoLM; Lewis (1965a, b, 1966); LONG
LAKELA; MorTON; SCOGGAN; SOUKUP; STANDLEY (1918); and VERDCOURT (1958, 1976).
Attims, Y. Influence de l’age physiologique de la plante mére sur la dormance des
graines d’Oldenlandia corymbosa L. (Rubiacées). Compt. Rend. Acad. Sci. Paris D.
275: 1613-1616. 197
BAHADUR, B. Heterostylisti in Oldenlandia umbellata L. Jour. Genet. 58: 429-439.
1963. [List of over 150 heterostylous species of Rubiaceae includes 39 species of
Oldentandia.|
. Heterostyly in Hedyotis nigricans (Lam.) Fosb. [bid. 60: 175-177. 1970a. [Ma-
terials from Texas; pins and thrums compared over a number of characters; dem-
onstrated incompatibility in illegitimate crosses, although pins selfed yielded some
seeds.
. Homostyly and heterostyly in O/denlandia umbellata L. Ibid. 192-198. 1970b.
[Homostyles with short styles and short stamens, some self-compatible, partly fertile
with heterostyles.
BENJAMIN, D. S. Estudo das Rubiaceae Brasileiras—II. Arg. Jard. Bot. Rio de Janeiro
18: 223-227. 1964 [1965]. [Hedyotis doer (H. Salzmanii), 224.
Braun, E. L. Two members of the Rubiaceae new to Ohio. Rhodora 78: 549-551. 1976.
[Houstonia setiscaphia possibly nae with H. canadensis; see also CARR.]
BREMEKAMP, C. E. B. Pleiocraterium genus novum Rubiacearum Hedyotidearum. Rec.
Trav. Bot. Néerl. 36: 438-445. 1939. ie eee Ue “‘and its nearest
allies” (p. 438) and gives this species as lectotyp ssion in present text);
156 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
excludes H. Auricularia from Hedyotis, for continued argument against H. Auric-
ularia as lectotype for Hedyotis, see BREMEKAMP (1952), pp. 29, 30
: new species of Oldenlandia (Rubiaceae) from India with remarks on its
inflorescence morphology. Kew Bull. 29: 359-361. 1974. [See BENNET, Jour. Econ.
Taxon. Bot. 4: 592. 1983, for combination in Hedyotis; includes characterization of
Oldenlandia sensu Brem. and descriptive discussion of sympodial growth and floral
arrangements in Oldenlandia.|
Buttock, A. A. Nomenclatural notes: VI. Type species of some generic names. Kew
Bull. 13: 97-100. 1958. [Hedvotis, 99.]
BURMAN, J. Thesaurus Zeylanicus. [14 +] 235 pp. + appendices + //0 [11/1] pls.
Amsterdam. 1736. [Hedvotis fruticosa (Valerianella foliis nervosis ...), H. Auricu-
laria (V. i ..), 227, 228, pls. 107, 108 (fig. 1).]
Carr, L.G. An w species of Houstonia from the cedar barrens of Lee County, Virginia.
Rhodora ‘e 306-310. 1944. [Houstonia setiscaphia Carr = Hedyotis canadensis,
see also BRAUN.
CHAMBERS, K.L. Hedyotis australis in Georgia. Rhodora 65: 271-273. 1963. [Compared
—. A. DE, & D. DE SCHLECHTENDAL. De plantis in expeditione speculatoria
manzofhana ee Rubiaceae. Linnaea 4: 129-202. 1829. [H. Auricularia as
, 153.)
CLute, W. N. The bluet. Am. Bot. 38: 57-59 + frontisp. to issue of journal. 1932.
[Popular account of H. caerulea. |
CorRBINEAU, F., & D. Come. Some particularities of the germination of Oldenlandia
corymbosa L. seeds (tropical Rubiaceae). Israel Jour. Bot. 29: 157-167. 1980. [In-
cludes scanning electron micrographs of seeds; effects of temperature and light,
scarification, hormones, and oxygen concentration; observations on parts of seeds
responsible for germination requirements.
& . Effect of the intensity and duration of light at various temperatures
n the germination of Oldenlandia corymbosa L. seeds. P|. Physiol. 70: 1518-1520.
1982. [Includes references not listed in the present paper. Authors studied seeds that
do not require stratification for germination but do require light; at cool temperatures
strong light of sufficient duration inhibits germination (under certain conditions even
longer exposures reverse inhibition).]
DALE, S. Pharmacologia, seu manuductio ad materiam medicam. ed. 3. [i +] frontisp.
[+ uu] + vil [+ vi] + 460 pp. London. 1737. [Auricularia, 146, 147.
Datta, P. C., & A. SEN. Pharmacognosy of O/denlandia corymbosa Linn. Quart. J
Crude Drug Res. 9: 1365-1371. 1969. [Includes medicinal uses, histology, —
tion of pollen, chemical tests, and ae
Dennis, W. M., D. H. Wess, B. E. Worrorp, & R. K State records and other recent
noteworthy salen of Tennessee plants. III. ae 45: 237-242. 1980 [1981].
ora
FAGERLIND, F. Embryologische, zytologische und bestaubungsexperimentelle Studien
in der Familie Rubiaceae nebst Bemerkungen tiber einige ha ae a
Acta Horti Berg. 11: 195-470. 1937. [Houstonia caerulea and H. longifolia lack
ovule integuments, 206; see also LLoyp and Rotn & oe |
FARMER, R. E., JR. Seed propagation of the Roan Mountain bluet. Jour. Tenn. Acad.
Sci. 54: 126-128. 1979. [Houstonia purpurea var. montana. ]
FarRoog, M. The endosperm and seed structure of O/denlandia corymbosa Linn. Curr.
Sci. Bangalore 22: 280-282. 1953. [Endosperm nuclear; before formation of walls,
vesicles a in central area of endosperm, a feature previously unknown in the
Rubiac
The ea biialee of Oldenlandia corymbosa Linn. Jour. Indian Bot. Soc. 37:
358- 364. 1958. [Ovules hemianatropous, embryo sac Polygonum type, pollen grains
trinucleate when shed, endosperm nuclear; germinated pollen encountered in closed
1987] ROGERS, CINCHONOIDEAE Let
— & M. INamuppIN. The embryology of Oldenlandia nudicaulis. Ibid. 48: 166-
173. 1969.
FosBerG, F. R. Some Rubiaceae of southeastern Polynesia. Occas. Pap. Bishop Mus.
13: 245- 293, 1937. [Includes discussion of merging Oldenlandia and Hedyotis.]
—. Observations on Virginia plants. Part 1. Virginia Jour. Sci. 2: 106-111. 1941.
[Hedyotis, 110, 111; includes new combinations, a new name in Hedyotis, and brief
discussion of generic limits; also see ibid., 284.
Notes on North American plants. IV. Am. Midl. Nat. 29: 785, 786. 1943a.
[Hedyotis Michawxii Fosb., nom. nov. (Houstonia serpyllifolia Michx.).
e Polynesian species of Hedyotis (Rubiaceae). Bishop Mus. Bull. 174: 1-102.
ols 1-4. 1943b. [Includes taxonomic history of Hedyotis, discussion of generic
definition, distinguishing features of Hedyotis s.l., and infrageneric classification. ]
Notes on plants of the eastern United States. Castanea 19: 25-37. 1954. [Hedy-
otis, 29-37; includes reiteration of position on merging Houstonia and Oldenlandia
with Hedyotis, synonymy, new combinations, and distributional information. ]
Observations on Hedyotis caerulea var. minor. Ibid. 20: 104-106. 1955. [Ob-
served in Alabama and Georgia; includes comments on habitats, habit, flowering
period, floral variation, taxonomic position, and absence of heterostyly.]
& E. ERRELL. A recently established exotic in west Florida and Alabama
(Hedyotis Salzmanii or Oldenlandia Salzmanii, Rubiaceae). Cee 50: 49-51
1985.
Fukuoka, N. Studies in the floral anatomy and morphology of Rubiaceae. II. Hedy-
otideae (Hedyotis). Acta Phytotax. Geobot. 29: 179-185. 1978. [Floral anatomy of
nine species described and illustrated.]
Gappy, L. L., & D. A. RAYNer. Rare or overlooked recent plant collections from the
Coastal Plain of South Carolina. Castanea 45: 181-184. 1980. [Houstonia procum-
ens. |
GRAN, L. Isolation of oxytocic peptides from Ol/denlandia affinis by solvent extraction
of tetraphenylborate complexes and chromatography on sephadex LH-20. Lloydia
36: 207, 208. 1973a
—. On the effect of a polypeptide isolated from ‘‘kalata-kalata” (Oldenlandia affinis
DC.) on the oestrogen dominated uterus. Acta Pharmacol. Toxicol. 33: 400-408.
1973b. [Used to hasten childbirth in Africa.]
n the isolation of tetramethylputrescine from Ol/denlandia affinis. Lloydia 36:
209, 210. 1973c. [Not oxytocic, not a true alkaloid; also found in Solanaceae. ]
Oxytocic principles of Oldenlandia affinis. Ibid. 174-178. 1973d.
GRAY, i Notes upon some Rubiaceae, collected in the United Dane South-Sea ex-
ee expedition under Captain Wilkes, with characters of new species, &c. Proc.
Am . Arts Sci. 4: 33-50, 306-318. 1858, 1860. [Includes Femrreens of rela-
pee among Hedyotis, Houstonia, and Oldenlandia.]
GREENMAN, J. M. Revision of the Mexican and Central American species of Houstonia.
roc. Am. Acad. Arts Sci. 32: 283-293. 1897. [Taxonomic treatments of several
species accompanied by very little discussion; includes sections (see p. 292).
Hatusima, S. On some species of Hedyotis soe Japan and Formosa. (In Japanese and
Latin.) Jour. Jap. Bot. 36: 296-298. 1961.
Hitcucock, A. . L. GREEN. Standard-species of Linnaean genera of Phaner
gamae (7532 54). Pp. 111-195 in International Bot. Congr. rca (England),
1930. Nomenclature proposals by British botanists. London.
Jackson, B. D. Guide to the literature of botany. Facsimile of the ae of 1881. xl +
626 pp. New York and London. 1964. [‘Marloe,” 199.
Kuastoir, H. N., S. K. SeNGupTA, & P. SENGupTA. Notes on the constituents of the
Indian medicinal plant Oldenlandia corymbosa Linn. Jour. Am. P Pharm. Ass soc. Sc cl.
Ed. 49: 562, 563. 1960. [Gamma-sitosterol, ursolic acid, and
esters and/or acetates); alkaloids not encountered.]
Krat, R. Areport on some rare, threatened, or endangered forest-related vascular plants
158 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
of the South. Vol. 2. U. = me Agr. Tech. Publ. R8-TP 2: [iv +] 719-1305. 1983.
[Houstonia montana (co red with H. purpurea), Hedyotis nigricans var. pulvi-
nata, 1074-1081; fae "distribution maps, descriptive and ecological informa-
tion, su si eee ee
Kuntu, C. Jn: A. HumMBoLpt, A. BONPL & C. Kuntu, Nova genera et species
on Vol. 3. [iv +] 456 pp. pls. “193 300. 1820. [Hedyotis, 388.
Lacey, J. B. Hedyotis corymbosa (L.) Lamarck (Rubiaceae) in Nacogdoches, Texas.
Field Lab. 25: 33, 34. 1957. [Dooryard.]
—— J. Tableau encyclopédique et aa des trois régnes de la nature. Vol.
.xvi[+ 1] + 496 pp. Paris. 1791-1793.
Lamon E. Hedyotis minima f. a ee Rhodora 59: 95. 1957. [Kansas.]
oy, J. Le mode de développement dans le genre O/denlandia (ee Hedyoti-
ae). Adansonia II. 15: 89-94. 1975.
eel D. A. Spatial segregation of pins and thrums in populations of Hedyotis nigricans.
Evolution 28: 648-655. 1974 [1975]. [Self- and intramorph-incompatible; apomixis
absent; the two morphs generally somewhat segregated spatially, but this varying
between populations; segregation possibly explainable as “ecological dimorphism”;
also see ORNDUFF (1980).]
Lewis, W. H. Chromosomes of East Texas Hedyotis (Rubiaceae). aaa Nat. 3:
204-207. 1958 [1959]. [Chromosomes illustrated for four specie
Rhodora 63: 216-223. 1961. [Includes extensive discussion and new transfers. |
Phylogenetic study of Hedyotis (Rubiaceae) in North America. Am. Jour. Bot.
49: 855-865. 1962. [Includes original vouchered chromosome counts for 39 taxa
covering about 85 percent of the species in North America, literature review for
additional counts, maps to show geographic distributions of chromosome numbers,
discussion of evolutionary trends in morphological characters for subg. EpristA,
ranking of species for advancement level related to chromosome numbers and ge-
ography, and validation of four varietal names; see Lewis (1965), below, for follow-
up with eye sas eee a ae and ieee et al. for continuation;
see Lewis & T LL for expan of chromosome c s.]
Oldenlendia ae smbosa Rees Grana Paleo: 5: 330-341. 1964. [In-
cludes chromosome numbers (diploid, tetraploid, hexaploid), pollen morphology,
systematic treatment, and distribution map.]
Pollen morphology and evolution in Hedyotis subgenus Edrisia (Rubiaceae).
Am. Jour. Bot. 52: 257-264. 1965. [Thirty-one species divided into five groups
based mostly on the structure of the tia a in pollen grains; phylogenetic scheme
from 1962 adjusted (includes dendrogram); see TERRELL et al. for continuation. ]
. Chromosome numbers of O/denlandia corymbosa (Rubiaceae) from southeast-
ern Asia. en Missouri Bot. Gard. 53: 257, 258. 1966a. [2n = 18, 36, 54
an genus Neanotis nomen novum (Anotis) and allied taxa in the rape
icas (Rubiaceae ies 53: 32-46. 1966b. [Correct names — in Hedyotis for
species once placed in Anotis; Neanotis proposed as new e for Asian oe
pollen of Hedyotis compared with Neanotis providing ae ‘for keeping the two
genera apart.]
. Typification of Hedy (Rubi a new variety from south-
eastern United States. bid 277, 378. 1966c. ae new var. hirsuta, but see
WILBUR.
. Notes on Hedyotis ead in North America. /bid. 55: 31-33. 1968a. [See
correction in Lewis, 1968b.]
—. Hedyotis acerosa var. ee comb. nov. (Rubiaceae). /bid. 397. 1968b [1969].
Hedyotis. Pp. 1487-1490 in D. S. CorreLt & M. C. JoHNsTon, Manual of the
eae plants of Texas. Renner, Texas. 1970. [15 species.]
Additions to the flora of the Bahama Islands. Rhodora 73: 46-50. 1971. [Hedy-
Bae nigricans, 48.
1987] ROGERS, CINCHONOIDEAE [59
-——. Hedyotis Correllii (Rubiaceae): a new Texas species. Brittonia 24: 395-397.
1972.
. Pollen size of Hedyotis caerulea (Rubiaceae) in relatio I number
and heterostyly. Rhodora 78: 60-64. 1976. [Contrary a Lewis’ s earlier opinion,
Oldenlandia accepted as genus; pollen from thrums on average larger than pollen
from pins; size of grains not related to ploidy level.]
. Moore. Hedyotis australis (Rubiaceae), a new species from the south
central United States. Southwest Nat. 3: 208-211. 1958 [1959]. [Compared with H.
crassifo re
. TERRELL. Chromosomal races in eastern North American species of
Hedyotis (orstona Rhodora 64: 313-323. 1962. [116 collections in H. caerulea
and H. purpurea groups examined; includes * ‘putative hybrids or intergrading col-
lections”’ and erat of cytology for subg. Edrisia.]
Lin, Y. C., W. C. Liao, Y. M. Lin, & F.C. CHEN. Cine of Hedyotis corymbosa.
Planta Med. 39: 278. 1980. [Includes uses in China and India; plants contain N-ben-
zoyl-L-phenylalanyl-L-phenylalaninol acetate, oleanolic acid, ursolic acid, gamma-
sitosterol, and stigmasterol.]
Linnaeus, C. Flora Zeylanica. 240 pp. + index + 15 pp. + 4 pls. Stockholm. 1747a.
[Hedyotis, 26, 27 (and 5, 6 in appendix).]
a plantarum genera. (Dissertation defended by C. M. Dassow.) [5 +] 14
pp. Stockholm. 1747b. [Hedyotis, 7, 8, reissued virtually unchanged in Amoen.
Acad. 1: 381-417. 1749.]
Lioyp, F. E. The comparative embryology of the Rubiaceae. Mem. Torrey Bot. Club
8: 1-112. 1899. [Absence of integument in Houstonia corroborated by FAGERLIND;
see also ROTH & LINDORF.]
Love, A., & D. Love. Tee oie remarks on some American alpine plants. Univ.
Colorado Stud. Biol. 17: 1-43. 1965. [Houstonia caerulea, 28, 29.
Ma taisse, F., J. GréGorre, L. Nyemso, & E. Rosprecut. A propos d’une recherche
d’alcaloides dans les Rubiaceae du Shaba méridional (Zaire). Bull. Jard. Bot. Bru-
xelles 49: 165-177. 1979. [Includes Oldenlandia and table showing subfamilies,
tribes, and genera with alkaloids.]
McVauGu, R. The vegetation of the gramuc ot ocks ore southeastern United States.
Ecol. Monogr. 13: 119-166. 1943. [ a, H. crassifolia, H. Nuttalliana,
160, 161.]
MEEHAN, T. Dimorphic flowers in Houstonia. Proc. Acad. Nat. Sci. Phila. 32: 349, 350.
1880. Tee caerulea, H. serpyllifolia, H. purpurea.]
MERRILL, E. D., & F. P. Metcatre. Hedyotis Linnaeus versus O/denlandia Linnaeus
and the status of Hedyotis lancea Thunberg in relation to H. consanguinea Hance.
Jour. Arnold Arb. 23: 226-230. p/. J. 1942.
MoHLENBROCK, R. H., & L. E. Hatsic. The annual species of Houstonia in Illinois.
Rhodora 64: 28-31. 1962. [Annual species compared with perennial; includes treat-
ment of H. caerulea, H. pusilla, and H. minima.]
Moore, D. M. New records for the Arkansas flora. IV. Proc. Arkansas Acad. Sci
9-16. 1958. [Hedvotis crassifolia var. micrantha Shinners probably deserves ae
status; 7. rosea.
Moore, J. E. Hedyotis rosea in Arkansas. Rhodora 58: 331. 1956.
MuE Ler, C. H., & M. T. MueLLer. A new Houstonia in southcentral Texas. Bull.
Torrey Bot. ae 63: 33, 34. 1936. oo pygmaea, sp. nov. (= Hedyotis rosea
Raf. fide SmitH; also see WATERFAL
Ornourr, R. An unusual homostyle in " Hedyotis caerulea (Rubiaceae). Pl. Syst. Evol.
127: 293-297. 1977. [Compares pins, thrums, and homostyles; homostyles rare—
only one plant known (cf. FosBerG, 1955);
themselves; homostyle self-incompatible but compatible as seed parent with het-
erostyles and as pollen donors with thrums (fertility much reduced with pins); in-
cludes comparison with homostyles in other typically heterostylous genera.]
160 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
. Heterostyly, population composition, and pollen flow in Hedyotis caerulea. Am.
Jour. Bot. 67: 95-103. 1980. [Pin/thrum ratio equal or pins predominant; ratio may
change from year to year or even within a year; distributions of the two morphs
random or deviating variably from random; pollen production about equal for the
two morphs or biased in favor of pins; pollen sterility moderate and highly variable
between morphs in some populations over time, and between populations (but,
overall, about equal for the two morphs); pollen flow greater from pins to thrums,
but seed set nearly identical (plants virtually self- and intramorph-incompatible);
intramorph pollen flow substantial (and largely intrafloral?); plants seemingly most
often pollinated by bombyliid flies.]
Patrick, T. S., & H.R. DeSeLm. Floristics of an East Tennessee cedar barren. (Abstract.)
ASB Bull. 32: 77. 1985. L[Houstonia nigricans.]
Pease, A. S., & A. H. Moore. An alpine variety of Houstonia caerulea. ‘ou 9:
209, 210. 1907. [Var. Faxonorum from Mt. Washington, New Hampshir
RAPINESOUE, C.S. Sur le genre Houstonia et description de plusieurs espéces aolee
etc. Ann. Gén. Sci. Phys. 5: 224-227. (Repaged as pp. 12-15 in reprint.) 1820. [14
species in four subgenera.]
RAGHAVAN, T. S., & K. RANGASWAMyY. Studies in the Rubiaceae. Part I. Development
of female pametophyte and oe Ene Dentella aes Forst. and Ol-
enlandia alata Koch. and some cyto Jour. Indian Bot
Soc. 20: 341-356. 1941. [Includes eel oo concerned with distinguishing
nucellus and integuments in reduced o
Rao, P. S., & K. S. Basu. oe of ae biflora Linn. Proc. Indian Sci.
Congr. Assoc. 62(3): 77.
Reep, C. F. Dentella repens me Hedyotis corymbosa, new to Aine United States. Phy-
tologia 19: 311, 312. 1970. [In Florida; also see Lewis (19
. Houstonia pusilla in Maryland and Virginia. etn 45: 35. 1980. [Spreads
Rocers, H. J. A new Houstonia from Chatham-Randolph County, N. C. (Abstract.)
Jour. Elisha Mitchell Sci. Soc. 69: 89. 1953. [No name supplied.]
Roth, I., & H. Lrinporr. La ea morfologica de la semilla de las Rubiaceae
y especial del café. Acta Bot. . 9: 141-147. 1974. [Houstonia with highly
reduced ovule, vestige of apna 145; see also FAGERLIND and LLoyp.]
SastTRI, S. B. N., chief ed., & COLLABORATORS. The wealth of India. Raw materials. Vol.
5. xxv + 332 + xii pp. 16 pls. New ce 1959. [Includes chemistry, uses, descrip-
tions, and references for several specie
SCHOENBECK, E. Houstonia minima in Peoria County. Trans. Illinois Acad. Sci. 40: 60.
1947.
—— L. H. Transfer of Texas species of Houstonia to Hedyotis (Rubiaceae). Field
Lab. 17: 166-169. 1949.
—. Hedyotis crassifolia Raf. var. lh Cas ee var. nov. Ibid. 18: 100.
1950. [= Hedyot ee see Lewis & Moor
SHIVARAMAIAH, G., & S. S. RAJAN. A on ito ee the embryology of Oldenlandia
mbellata Linn. Proc. Indian ee Sci. B. 77: 19-24. 1973. [Includes short literature
review for embryology of Rubiaceae.]
& K.S. Rao. Studies in Rubiaceae Il. Structure and oo of seed of
Oldenlandia gracilis DC. Curr. Sci. Bangalore 46: 662-664. 1977.
sr S. A., & S. B. Srppigur. Studies in the Rubiaceae I. ‘ contribution to the
mbryology of Oldenlandia dichotoma Hook. f. Beitr. Biol. Pflanzen 44: 343-351.
1968.
Stmmonps, P. L. Tropical agriculture. A treatise. ed. 3. xvi [+ i] + 539 + 33 pp. New
York and London. 1889. _tedyotis umbellata, ae 373.]
Situ, E. B. Hedyotis (Rubia : a new species from the Ouachita Moun-
tains of Arkansas and Oklahoma. Britionia 28: 453. 459. 1976 [1977]. [Compared
1987] ROGERS, CINCHONOIDEAE 161
and artificially crossed (failed) with Hedyotis longifolia; includes distribution map
for the new species (27 = 12) and H. longifolia var. longifolia in Arkansas and eastern
Oklahoma. ]
STEARN, W. T. An introduction to the Species Plantarum and cognate botanical works
of Carl Linnaeus. xiv + 176 pp. Jn: Ray Society facsimile of C. LINNAEUS, Species
Plantarum. Vol. 1. London. 1957. (Species planta y published in 1753.)
STEYERMARK, J. A. Bluets as summer flowers. Missouri Bot. Gard. Bull. 36: 93. 1948.
[Houstonia minima, H. pusilla, H. caerulea.
TAKAGI, S., Y. YAMAKI, K. MAsupa, Y. NISHIHAMA, & K. SAKINA. Studies on the herb
medical Coen used for some tumors. IJ. On the constituents of Hedyotis cor-
ymbosa Lam. (In Japanese; English summary.) Jour. Pharm. Soc. Japan 101: 657-
659. 1981. [Six iridoids, asperuloside, scandoside methyl ester, asperulosidic acid,
geniposidic acid, scandoside, deacetylasperulosidic acid.
TAYLor, R.J.,& C. TAYLor. The vascular flora of Oklahoma—additions and comments.
Rhodora 71: 215-219. 1969. [Hedyotis rosea, 218.
TERRELL, E. E. A revision of the Houstonia purpurea group (Rubiaceae). Rhodora 61:
157-180, 188-207. 1959. [Includes taxonomic history, chromosome counts, a
cussion of intergradation (with intergrading species pairs listed), key, taxonom
treatments of species, and distribution maps; for cytology cf. Lewis (1962) and i
RRELL
—. New combinations in Houstonia (Rubiaceae). Phytologia 31: 425, 426. 1975a.
Fo oustonia Correllii, H. mic hk (Hedyotis australis) not conspecific with Hous-
tonia pusilla (Hedyotis ti)
. Relationships of Hedyotis He L. to Houstonia L. and Oldenlandia L. Ibid.
418-424. 1975b.
xonomic notes on Houstonia purpurea var. montana (Rubiaceae). Castanea
43: 25-29. 1978. [Refutes YELTON’s treatment of Houstonia montana as a species,
Zee authorship, and gives synonym
w species and combinations in Howstoni (Rubiaceae). Brittonia 31: 164—-
169. 1978, [All in Lianne Texas, or New Mex
; oO new species a combination 1 nee ustonia ee from Mexico.
RaueRs 32: 490-494. re [1981], (Houstonia Sharpii, ing
New combinations in Houstonia and Oldenlandia Gabe: Phytologia 59:
79, 80. 1985. [Four new combinations -
, H. Lewis, H. Rosinson, & J. W. Nowicke. Phylogeneti of diverse
seed types, chromosome numbers, and ie morphology in i aeatona (Rubiaceae).
Am. Jour. Bot. 73: 103-115. 1
Trimen, H. Hermann’s Ceylon nerbaniui and Linnaeus’s “Flora Zeylanica.”’ Jour, Linn.
Soc. 24: 129-155. 1887. [Hedyotis, 137.
Usuer, G. A dictionary of plants used by man. 619 pp. New York. 1974. [Oldenlandia,
421.
Urrta_, L. J. Five amendments to the flora of southwest Virginia. Castanea 36: 79-81.
1971. Te setiscaphia, 79, 80; agrees with TERRELL’s reduction of this to
synonymy under Houstonia canaden Sis. ]
& R.S. Mircuett. Amendments to the flora of Virginia—II. Castanea 37: 96-
118. 1972. [Hedyotis Boscii, H. uniflora, 118.]
WATERFALL, U. T. The identity of Hedyotis rosea Raf. Rhodora 55: 201-203. 1953.
[Also see TAYLor & TAyYLor; synonyms: Houstonia pygmaea Mueller & Mueller
(Hedyotis Taylorae Fosb. with same type), Houstonia patens Ell. var. pusilla Gray.]
WIGHT, R., & G. . WALKER-ARNOTT. Prodromus florae peninsulae Indiae orientalis.
Vol. 1. xxxvii + 480 pp. facsimile ed. Dehra Dun and Delhi, India. 1976. (Originally
published in a 1834.) [Hedyotis, 405-418, in sections; H. Auricularia “‘the
acknowledged type of the genus,” 411.]
Wivpur, R. L. The status of Hedyotis procumbens var. hirsuta (Rubiaceae). Rhodora
162 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
70: 306-311. 1968. Sane ae recognition of the variety and dubious of Lewis’s
(1966b) selection of neo
& W
WUNDERLIN, R. P., _E. eee A new form of Houstonia pusilla from Mlinois.
Trans. Illinois Acad. Sci. 59: aes pee [H. pusilla f. albiflora. ]
Wyatt, R., & R. L. HELLwic. Factors determining fruit set in heterostylous bluets,
Houstonia caerulea (Rubiaceae). can Bot. 4: 103-114. 1979 [1980]. [Includes pol-
linators, comparison of pins and thrums, crosses, and selfing; authors consider re-
lationchin between fruit set and sizes of populations, ratios of morphs within pop-
ulations, and distances to nearest compatible populations. ]
YELTON, J. D. Houstonia montana, a species, not an ecological variety. Castanea 39:
149-155. 1974. [Includes crossing experiments; refuted by TERRELL (1978).]
3. Pentodon Hochstetter in Krauss, Flora 27: 552. 1844.
Hygrophilous, prostrate or feebly erect, fleshy, glabrate herbs, usually exten-
sively branched, frequently pseudodichotomously so, often tufted with nu-
merous basal branches; branches more or less quadrangular. Raphide bundles
conspicuous on surfaces of most organs when dry. Leaves opposite, nearly
sessile or on short, winged petioles, the blades (obovate to) lanceolate or ovate,
penninerved, usually minutely scabrous adaxially and marginally, rounded to
more often acute or acuminate at the apex, the base usually acute to cuneate
or sometimes rounded; stipular sheaths continuous with the flanges on the
petioles, membranaceous, interpetiolar, usually fimbriate, occasionally entire,
sometimes cuspidate in the center. Inflorescences mostly terminal, sometimes
axillary, usually between a pair of pseudodichotomous branches, fundamentally
dichasial or monochasial, sometimes with only | or 2 flowers, lax with long
axes, sometimes compound and sometimes paniculate with straight main or
branch axes; bracts and bracteoles mostly distinctly reduced [or foliose]. Flow-
ers pedicellate, pentamerous, small and inconspicuous, perfect, homostylous
[or heterostylous in P. /aurentioides and P. pentandrus var. minor, or ““pseu-
doheterostylous” in some African members of P. pentandrus var. pentandrus
having the anthers in fairly uniform position in the throat of the corolla but
the styles varying in length]. Calyx lobes connate basally into a short tube
topped with lanceolate or deltoid teeth 4-4 the length of the corolla. Corolla
nearly cylindrical but slightly [to broadly] flared, white [or reddish or blue],
pubescent or (reportedly) glabrous in the throat, the lobes usually about 4-4
the length of the corolla. Stamens inserted near the throat of the corolla tube
[or low in the tube in heterostylous flowers], uniform in length and included
[or exserted in short-styled flowers]; anthers dorsifixed, elliptic-oblong; fila-
ments shorter than anthers; pollen grains prolate or subspheroidal, tricolporate,
reticulate. Ovaries bilocular, containing numerous ovules on apically bilobed,
peltate placentae inserted on the septum; styles long enough to bear slightly
exserted [or included] stigmas, at least sometimes markedly thickened at the
level of the anthers beneath the stigmatic lobes, the thickening covered with
pollen and, in conjunction with a pilose ring at the same level, occluding the
throat of the tube; stigmatic lobes 2, linear. Capsules bilocular, crowned with
persistent calyx tube and teeth, thin walled and papery, obconical or obtur-
binate, somewhat compressed contrary to the septum, bearing 5 longitudinal
1987] ROGERS, CINCHONOIDEAE 163
keels corresponding to the midlines of the adherent sepals, dehiscing loculi-
cidally across the summit. Seeds numerous, minute, angular, brown, fairly
isodiametric, on the surface reticulate from outlines of testa cells, these with
irregular thickenings in the lateral walls. Type species: P. decumbens Hochst. =
P. pentandrus (Schum. & Thonn.) Vatke fide Bremekamp (1952); this the sole
original species. (Name from Greek, pente, five, and -odon, toothed, presumably
in reference to the five toothlike calyx lobes.)
Probably consisting of only two species, Pentodon laurentioides Chiov., en-
demic to Somalia, and P. pentandrus, 2n = 18, distributed in the Old World
across much of tropical Africa and on the southern Arabian Peninsula, Mad-
agascar, the Seychelles, and the Cape Verde Islands. The latter, or possibly a
third species, P. Halei (Torrey & Gray) Gray (Hedyotis Halei Torrey & Gray,
Oldenlandia Halei (Torrey & Gray) Chapman) is scattered across much of
Florida and occurs in southern Georgia, Louisiana, Texas, the West Indies (at
least Cuba, the Bahamas, and Guadeloupe), and according to Verdcourt (1976),
Nicaragua and Brazil. (I have seen no trustworthy documentation of Pentodon
from either Mississippi or Alabama.)
Opinion is divided as to whether Pentodon Halei is conspecific with P.
pentandrus. Standley (1918) held the latter to differ from P. Halei in having
pubescence within the corolla, longer peduncles relative to the leaves, racemose
(vs. cymose) inflorescences, and more slender (vs. “‘clavate’’) pedicels longer
relative to the capsules. This list probably exaggerates the differences —corollas
from P. Halei that examined are distinctly pubescent within, and Bremekamp
(1952, p. 180) found the distinctions to break down so far as to be “of little
importance,” if the range of variation in African specimens is considered. He
attributed differences in the inflorescence characters largely to differences in
the vigor of the plants, which he assumed to be reduced in the marginal North
American climate. Noting that the American material has small, elliptic leaves
and shorter inflorescences than most African specimens, Verdcourt (1976, p.
263) agreed that P. Halei “cannot be specifically distinct” from P. pentandrus
and agreed further with Bremekamp in suspecting introduction from Africa as
lying behind the New World populations of Pentodon. Its widely scattered
stations speak in favor of an appreciable ability to disperse. As Verdcourt has
already noted, better data on the distribution of modifications to the style, as
described below, could shed some light on the relationships among the widely
separated populations.
Pentodon appears to be most closely related to Hedyotis (especially subg.
OLDENLANDIA), in which it has been included, and from which it differs by the
pentamery (vs. tetramery) of its flowers and the distinctive thickenings on the
lateral walls of testa cells. Additional features that help to characterize Pentodon
are its apically bilobed placentae; thin, papery pericarps; and seeds not pro-
ducing mucilage upon moistening. (This paragraph is based largely on Bre-
mekamp, 1952, and Lewis, 1965a, and verified for Pentodon through herbarium
specimens.)
Pentodon laurentioides and P. pentandrus var. minor are heterostylous (for
an illustration of the two floral morphs in var. minor, see Verdcourt, 1976).
164 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
The other members of the genus show two curious variations of the breeding
system that call for further research. In the simpler case, the two flowers of P.
pentandrus from our area that I have been able to examine internally (Duncan
21650, Georgia, A, and Thomas et al. 72765 & 474, Louisiana, GH) have had
the style swollen apically and coated with pollen at the level of the anthers just
below the stigmatic lobes. The swelling was so positioned that, in conjunction
with the pilose ring borne on the tube, it would partly block entrance to the
corolla tube. Except fora thickened stylar apex (with stigmas missing) illustrated
in Godfrey & Wooten, I have seen no other indication of the thickening or of
adherent pollen for either African or American specimens. The functional role
of this condition, if any, will be best elucidated by field observations.
The second curiosity comes from Bremekamp (1952; also see Verdcourt,
1976), who described two floral morphs in African plants of P. pentandrus var.
pentandrus. The styles on different individuals are either of two lengths, in-
cluded or exserted, but the plants are not heterostylous in the conventional
sense of the term, since all flowers have included stamens. Bremekamp indi-
cated that the two morphs were geographically separated, although only on a
local scale; both are widespread in Africa.
This raises the question of the condition(s) in American populations. By
using bright transmitted light, I have consistently seen the anthers to occupy
about the same level in the corolla throats in all examinable flowers from our
area in the Harvard herbaria; all of the stigmas that I saw projected slightly
beyond the anthers. Moreover, the relative positions of stamens and stigmas
in the flower from the Bahamas illustrated by Correll & Correll are the same
as I observed on the mainland specimens; this seems also to be true of the
flowers shown by Small and by Godfrey & Wooten, although the long style is
depicted in each as detached, making its exact position relative to the stamens
indiscernible. Still, because the sampling so far is scanty, and because short,
included styles could be overlooked in an examination by transmitted light, it
would be premature to rule out the presence of such styles in the United States.
Pentodon pentandrus flowers in our area from May into October along shores
and in periodically flooded spots, swampy woods, and other low, wet sites.
An incidental note potentially useful in the field, pointed out by Dr. Robert
Kral (pers. comm.), is that in habit and overall appearance, Pentodon looks
deceptively like Lindernia crustacea (L.) F. Mueller, an introduced scrophu-
lariaceous weed in Florida.
Economic uses for this genus are negligible.
REFERENCES:
Under subfamily references see BREMEKAMP (1952); CORRELL & CORRELL; GODFREY
& Wooren; Lewis (1965a); SMALL; STANDLEY (1918); and VerDcourtT (1976)
Acnew, A. D. Q. Pentodon. Upland Kenya wild flowers. ix + 827 pp. London. 1974.
[Pentodon, 401.]
Dyer, R. A. The genera of southern African te plants. Vol. 1. Dicotyledons.
{3 +] 756 pp. Pretoria. 1975. ca 08.]
Fournet, J. Flore illustrée des phanérogames de Guadeloupe et de Martinique. 1654
pp. Paris. 1978. [Pentodon, 1160, 1161; ee copied from HALLE and based
1987] ROGERS, CINCHONOIDEAE 165
on an African specimen, this probably also true of mention of exserted anthers in
addition to included ones.
HA.ié, N. Rubiacées. Pt. 1. Fl. Gabon 12: 1-278. 1966. [Pentodon, 105, 106; detailed
illustrations, 77, 107
ScHWARTZ, O. Flora des tropischen Arabien. Mitt. Inst. Allg. Bot. Hamburg 10: 1-393.
1939. [Pentodon, 261.]
Woop, J. M., & M. S. Evans. Natal plants. Vol. 1. 83 pp. /00 pls. Durban. 1898.
[Oldenlandia macrophylla (P. pentandrus), 31, pl. 36, stamens slightly exserted.]
Tribe CINCHONEAE DC. Ann. Mus. Hist. Nat. Paris 9: 217. 1807,
““Cinchonacées, Cinchonaceae.”’
4. Exostema (Persoon) L. C. Richard ex Humboldt et Bonpland, Plantae Ae-
quinoctiales 1: 131. 1808 [1807]
Vegetatively glabrous to less often hispidulous or hirsute shrubs or small
trees, the branches symmetrical, sometimes supported by surrounding vege-
tation. Leaves opposite, petiolate [or nearly sessile]; stipules interpetiolar [or
reportedly intrapetiolar], broadly deltoid to drawn out into attenuate apices,
marginally ciliate, keeled when young [sometimes bilobed]. Flowers borne
singly on short pedicels in axils of upper leaves [or terminal; in cymes, thyrses,
or panicles in some species], pentamerous [or tetramerous], actinomorphic or
nearly so, fragrant. Calyx teeth broadly deltoid [to subulate], much shorter than
corolla tube. Corolla with slender cylindrical tube [less than 1 cm to] several
cm long (ca. 2-5 cm in our species) [20 cm or more in E. /ongiflor'um Roemer
& Schultes], white, yellowish, or pinkish [red or purplish], said to change from
white to darker hues in some species including ours, the 5 [4] linear-ligulate
lobes about as long as the tube or a little [or much] shorter, twisted-imbricate
in bud. Stamens exserted [rarely included], epipetalous near base of tube [or
reportedly inserted on receptacle], the linear, basifixed anthers long (10 mm
or more in our species). Style filiform, much exserted [or infrequently included],
thickened apically beneath a pair of stubby stigmatic lobes [or stigma reportedly
unlobed]. Capsule ellipsoid, truncate apically, crowned with persistent calyx
teeth [or teeth deciduous], dark colored, rugulate, septidical (and sometimes
splitting loculicidally to varying degrees); placentae large, flat, detached from
septum of dry and dehisced capsule. Seeds numerous, wafer thin, surrounded
by a narrow marginal wing, vertically imbricate; endosperm abundant; embryo
with radicle longer than the elliptic cotyledons. LecrotyPe species: E. carl-
baeum (Jacq.) Roemer & Schultes.* (Name from Greek, exo, out, and stema,
stamen, in reference to the exserted stamens.)— PRINCEWOOD.
A genus of some 35 or more species in tropical and subtropical America,
mostly in the West Indies, but also with a poorly studied group of roughly
‘Britton & Millspaugh’s choice of Exostema parviflorum A. Rich. as lectotype (in Bahama FI. 409.
1920) cannot be followed. This species is ruled o ut by ICBN (1983) Article 7.10, since this not
na
(Syn. Pl. 1: 196. 1805; cited by Richard on p. 135). (See Brizicky for comments on Persoon’s
infrageneric taxa.) Exostema caribaeum 1s here ee as lectotype—Persoon included it, and it
is the most-widespread and best-known specie
166 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
seven species on the mainland in southern Florida (see below), Mexico, Central
America, and (chiefly western) South America as far south as Peru (four species?)
and southern Brazil (one species; see Angely).
Exostema caribaeum ranges along the full length of the Florida Keys, is
unusual on the southern tip of mainland Florida (Tomlinson), occurs through-
out much of the West Indies, Mexico, and Central America, and has been
reported from scattered localities along the northern coast of South America
(probably present in Colombia, but doubtfully so in Venezuela and ““Guiana”’).
Features that help with recognition of Exostema caribaeum are elliptic leaves
pointed at both ends; solitary, axillary flowers; short, stubby calyx teeth (less
than 1 mm long); fragrant, white (or pinkish or yellowish) corollas to ca. 8 cm
long, including the long, nearly linear lobes, and with slender, cylindrical tubes;
long (1 cm or more), basifixed anthers conspicuously exserted; and ellipsoid,
apically truncate capsules elliptic, waferlike seeds to about
5 mm long completely surrounded by a narrow wing.
Exostema 1s our only member of the tribe Cinchoneae (woody plants with
bilocular capsules containing numerous vertically or nearly vertically arranged,
imbricate seeds having pitted testa cell walls). The genus was once included in
Cinchona L., from which it differs in its exserted stamens and its imbricate
(vs. valvate) corolla lobes. Koek-Noorman & Hogeweg found Exostema to
differ further from Cinchona in having fiber tracheids in the wood, rather than
fibers transitional between fiber tracheids and libriform fibers, although broad-
ened sampling is needed to bolster the strength of this character. Additional
features that help to separate Exostema from other members of the Cinchoneae
are uniform calyx lobes, five or sometimes four corolla lobes, and slender,
round, symmetrical corolla tubes. Koek-Noorman cited personal communi-
cation with C. Bremekamp in noting that the relationships of Exostema are
unclear.
Taxonomic study of Exostema 1s both outdated and fragmentary. The most
recent revision of the entire genus dates back to De Candolle, who divided it
into three sections that have been ignored by more recent authors. Most of the
species are covered in Standley’s treatment in the North American Flora (1918),
a picture that can be rounded out by an examination of some of his later
floristic studies in the New World (1926, 1930, 1936, 1938: 1975, with L. O.
Williams).
During the eighteenth century, medicinal interest in Cinchona, the original
source of quinine as a medicine for malaria, extended to numerous species of
Exostema. | know of no modern study aimed at relating the alleged curative
properties of Exostema to bona fide pharmacologic effects or to its chemistry.
Exostema caribaeum and undoubtedly other species yield a hard, strong, heavy
wood that polishes well and is used for turning, cabinet work, and applications
requiring durability. Because it burns readily, it has been used for torches.
Species of Exostema with showy flowers are sometimes cultivated in the West
Indies.
REFERENCES:
Under subfamily references see ALAIN; ANGELY; BrizicKy; DE CANDOLLE
KoOEK-NoorMAN; KOEK-NOORMAN & HoGEweEG; Lirtce (1978); Little & WADSWORTH:
1987] ROGERS, CINCHONOIDEAE 167
LonG & LAKELA; SARGENT; SOUKUP; Slane 1926, 1930, 1931, 1936, 1938);
STANDLEY & WILLIAMS; and TOMLINSO
Boruipt, A., & O. Muniz. New plants in Cuba II. Acta Bot. Acad. Sci. Hungar. 18: 29-
48. 1973. [Exostema caribaeum var. pubescens, var. nov.,
EQUEIRA. Studies in Rondeletieae @upiacese) IL. A new gen
beranthus. Acta Bot. Acad. Sci. Hungar. 27: 313-316. 1981. [Exostema neriifolium
is type species.]
BRITTEN, a An overlooked Cinchona. Jour. Bot. London §3: 137, 138. 1915. [Also see
ORGAN, WARNER, and KenrISH. Includes no biological montane oe uses; con-
eye er history in literature and synonyuly; makes ema Sanc
tae-luciae with four “‘cinchonas” as synon
HEcKEL, E. Sur la présence et la nature des cystolithes dans le genre Exostema (Rubi-
aceae). Bull. Soc. Bot. France 35: 400-403. 1888. [Cystoliths present in E. floribun-
dum but not encountered in FE. caribaeum.]
Hooker, W. J. Exostema longiflorum. Bot Mag. 71: pl. 4186. 1845.
KentTisH, R. Experiments and observations on a new species of bark. xii + 123 pp.
London. 1784. [Cinchona sanctae-luciae (see BRITTEN for combination in Exoste-
ma: also see WARNER and Moraan); a series of chemical experiments described;
the species as the subject of earlier writings identified; application against malaria
and other complaints; case sea and preparation, effects, and uses of Cinchona
(including the species in questio
LEMESLE, R., & R. LAFAYE. Conn budel a l'étude anatomique et microchimique de
lV Exostema floribundum Roem. et Schult. ee piton). Bull. Soc. Sci. Bretagne
19: 30-42. 1946. [Includes comparison with Cinc
Moraan, J. Medical history of the cortex ruber, or a bark. Trans. Am. Philos. Soc.
2: 289- 293. 1786. [Also see BRITTEN, WARNER, and KENTISH. Includes letters by T.
a Ducue and G. Davipson concerned with what are evidently species of Exostema,
Saat Cinchona caribaea as used here is probably not the modern Exostema
caribaeu
PRAIN, D. ane ee subcordatum. Bot. Mag. 135: p/. 8274. 1909.
SANCHEz-VIESCA, F. The structure of exostemin, a new 4-phenyl coumarin isolated from
Exostema caribaeum. Phytochemistry 8: 1821-1823. 1969.
Warner, M. F. Exostema sanctae-luciae. Jour. Bot. London 56: 55. 1918. [See also
BRITTEN, MorGan, and Kentisu; clarification of bibliographic histo
WEBERLING, F. Beitrage zur Morphologie der Rubiaceen— Infloreszenzen. Ber. Deutsch.
Bot. Ges. 90: 191-209. 1977. [Includes F. floribundum and E. caribaeum
WRIGHT, W. Description of the Jesuits bark tree of Jamaica and the Ca rib beEs, Philos.
Trans. Roy. Soc. London 67: 504-506. pl. 10. 1778. [Cinchona jamaicensis, Cin-
chona caribaea (Exostema caribaea).]
Tribe NAucLEEAE J. D. Hooker, FI. Nigrit. 377. 1849.
5. Cephalanthus Linnaeus, Sp. Pl. 1: 95. 1753; Gen. Pl. ed. 5. 42. 1754.
Deciduous (or somewhat evergreen in tropical Florida), sympodially branched
shrubs (or infrequently small trees) of wet soil. Leaves opposite or in whorls
of 3 (or 4), elliptic to ovate or lanceolate, usually acuminate and often cuspidate
apically, the bases variable; stipules usually with 1 deltoid or ovate lobe between
bases of adjacent petioles, sometimes bifid, or occasionally with 2 separate
lobes between pairs of petioles, the lobe(s) with adaxial and frequently marginal
colleters; foliage and twigs (especially abaxial surfaces) glabrous to densely
pubescent, the indument sometimes storied and sometimes strigose; buds often
multiple in leaf axils. Flowers fragrant, usually tetramerous, protandrous, tight-
168 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
ly clustered into distinctly globose heads on long peduncles, the heads terminal
or axillary, sometimes solitary, more often in racemose (or infrequently pa-
niculate) clusters at ends of branches. Calyx much shorter than corolla, the
tube topped with short, blunt teeth persistent in fruit. Corolla white or nearly
so, with a narrow, cylindrical or slightly flared tube several times longer than
the oblong to deltoid or ovate, imbricate, usually internally bearded lobes,
these alternating with exposed glands (colleters?) in the bud and sometimes
after expansion. Anthers sagittate, borne at throat of corolla tube on short,
epipetalous filaments. Style filiform, about twice the length of the corolla,
expanded apically into a scarcely (or not perceptibly) bifid or 4-lobed knob
(fide Tomlinson); ovary bilocular, containing a pendulous ovule in each locule.
Fruits dry, indehiscent, crowded on spherical head, each with | or 2 seeds,
cuneiform, the halves often separating along the septum, intermixed with long,
narrow bractlets, these as long as the fruits and expanded apically into pubescent
knobs. Seed matching shape of locule, with a conspicuous corky caruncle (aril).
LECTOTYPE SPECIES: C. occidentalis L.; see Haviland, Jour. Linn. Soc. Bot. 33:
2, 3, 37. 1897; Britton & Brown, Illus. Fl. No. U.S. & Canada. ed. 2. 3: 255.
1913; Merrill, Jour. Wash. Acad. Sci. 5: 532. 1915. (Name from Greek, kephale,
head, and anthos, flower, in reference to the spherical floral heads.) — BUTTON
BUSH.
A genus of six species as circumscribed in Ridsdale’s revision: Cephalanthus
natalensis Oliver (South Africa), C. tetrandra (Roxb.) Rids. & Bakh. (India to
Taiwan), C. angustifolius Lour. (southeastern Asia), C. glabratus (Sprengel)
K. Schum. (South America), C. salicifolius Humb. & Bonpl. (Texas, Mexico,
Central America), and our C. occidentalis L. (In the revision preceding Rids-
dale’s, Haviland recognized seven species; Ridsdale transferred two of these
to Ixora L., changed the name of one, and added one.)
Cephalanthus occidentalis, 2n = 44, ranges across North America virtually
throughout the area defined by New Brunswick (or possibly Prince Edward
Island, according to Scoggan), Cuba, Texas, southeastern Nebraska, southern
Minnesota, southern Ontario, and southern Quebec. The species is absent or
nearly so from the Florida Keys. A spottier distribution farther west excludes
the Rocky Mountains but includes New Mexico, Arizona, Utah, California,
and northern Mexico. Standley & Williams noted it in Guatemala and Hon-
Cephalanthus occidentalis is almost exclusively an inhabitant of freshwater
shores and low, wet places. It usually grows in full sun but tolerates some
shading. The stands can be dense and extensive.
Distinguishing Cephalanthus from other shrubs in the Generic Flora area is
not difficult; the restriction to wet sites is a useful character in itself. The pointed
leaves are opposite or whorled and are associated with interpetiolar stipules
that bear adaxial and often marginal colleters. The small, tubular, fragrant,
white or nearly white flowers with long, exserted styles are packed into globose
heads, a shape that remains unaltered as the fruits mature. Individual fruits
are indehiscent (the halves often separate but do not open) and cuneiform; they
generally contain a conspicuously carunculate seed in each locule.
1987] ROGERS, CINCHONOIDEAE 169
Figure 1. Cephalanthus. a-i, C. occidentalis: a, pendent flowering branchlet, x ‘2;
b, flower at anthesis, style not yet expanded—note squamule between 2 corolla lobes, x
6; c, flower with mature style, the pollen shed—note bractlets at base of ovary, squamule
between 2 corolla lobes, x 6; d, corolla laid open to show adnate staminal filaments, =
6; e, ovary in longitudinal section, | ovule (at left) in section, x 10; f, mature fruit, x
6; 2, fruit splitting into 2 indehiscent 1-seeded parts, x 6; h, seed, abaxial side ta
adaxial side at right—note corky caruncle, x 6; i, embryo, oriented as in seed,
Western populations that have narrow leaves on short petioles have been
set apart as Cephalanthus lentalis var. californicus Bentham (C. occidentalis
subsp. californicus (Bentham) E. Murray), another segregate that Ridsdale placed
in synonymy. Fernald recognized plants with lanceolate leaves attenuate at
both ends and only 1-3 cm broad as forma /anceolatus. Different individuals
of C. occidentalis range from being more or less glabrous to thickly pubescent
on twigs and abaxial leaf surfaces, a condition that has led some authors (e.g.,
Steyermark, 1963) to recognize C. occidentalis var. pubescens Raf., which 1s
170 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
found primarily in the southern United States but has been reported from as
far north as Quebec. Neither Haviland nor Ridsdale recognized the pubescent
taxon at any rank, and Wells & Sharp rejected it with the observation that the
two putative varieties grow together in Tennessee. My examination of her-
barium specimens at the Missouri Botanical Garden inclines me to agree with
these authors.
Curious threadlike structures (called bracteoles by Haviland and Tomlinson,
illustrated in FiGure | and in Tomlinson) are borne at the base of each flower.
These are roughly as long as the ovary and are expanded apically into pubescent
knobs that appear to plug the spaces between the fruits protectively.
The flowers within each head mature simultaneously and are protandrous,
the pollen being released in the bud. Some grains catch in the hairs inside the
corolla, and others are carried out of the tube on the apical region of the strongly
exserted style. Whether all pollen delivery takes place from the style is not
clear. Some wind pollination is suspected. (For more on pollination, see Rob-
ertson and Tomlinson.)
Ants may possibly play some role(s) in the life cycle of Cephalanthus occi-
dentalis. ‘“Squamules” readily interpretable as nectaries (colleters?) are con-
spicuous in the sinuses between unexpanded corolla lobes in the bud (see
Ficure |b, c), and the seeds are capped with large, corky caruncles (arils). It
is not inconceivable that the adaxial colleters on the stipules, too, provide
nourishment for ants.
In their revisions, written in the last century, both Schumann and Haviland
placed Cephalanthus in the tribe Naucleeae, where Haviland regarded it as
closely related to the African and Asian genus Adina Salisb. Cephalanthus
differs from Adina in having only one ovule in each locule of the ovary, in-
dehiscent fruits, and wingless seeds. In 1976 Ridsdale revised Cephalanthus
and isolated it as the monotypic Cephalantheae Ridsdale.
Ridsdale defended his isolation of Cephalanthus by arguing that the tribe
Naucleeae is in part artificially held together by too much emphasis on the
conspicuous clustering of flowers into heads. He thought Cephalanthus possibly
to be most closely related to Mitragyna Korth. and Uncaria Schreber, two
genera he transferred from the Naucleeae to the Cinchoneae. Cephalanthus
differs from these two in its indehiscent fruits and its single seed per locule.
In their survey of alkaloids in the Naucleeae sensu lato, Phillipson, Hem-
ingway, & Ridsdale found Cephalanthus, along with Uncaria and Mitragyna,
to deviate from the Naucleeae sensu stricto in producing “significant quantities”
ofnonquaternary nonglycosidic alkaloids of the heteroyohimbine and oxindole
types. Aware of the same set of alkaloids in Cephalanthus, Kisakurek and
colleagues agreed that the data support maintaining all three genera apart from
the Naucleeae. Further, Koek-Noorman interpreted the wood structure of C.
occidentalis and C. salicifolius as anomalous in the tribe, and Bremekamp
*He attributed the authorship to Kunth in HBK., where Cephalantheae appeared, as Ridsdale
acknowledged, as a “‘sectio.”” Although ‘‘Cephalantheae” has been used repeatedly as a name for
subgroups of the Rubiaceae (see Darwin, Pfeiffer), Ridsdale appears to have been the first to call it
a tribe.
1987] ROGERS, CINCHONOIDEAE 171
(1966) disfavored a place for Cephalanthus in his narrowly conceived Nau-
eeae.
Cephalanthus is of minimal consequence in human affairs. The plants are
amply supplied with alkaloids and, not surprisingly, are bioactive. They are
blamed for killing livestock, but Sperry and colleagues noted that losses are
negligible in Texas, probably on account of unpalatable constituents. Cepha-
lanthus occidentalis has long been used in folk medicine by American Indians,
among others, against such complaints as sore eyes, arthritis, toothache, fevers,
and diabetes, and it has found use as a laxative. Sometimes C. occidentalis is
grown ornamentally. According to Fernald, C. angustifolius Hort. (non Lour.)
may be C. occidentalis f. lanceolatus Fern. The fruits serve as food for water
birds, and the sweet-smelling flowers are valued by beekeepers as sources of
nectar.
REFERENCES:
Under subfamily references see BREMEKAMP (1966); CORRELL & CORRELL; DARWIN;
GoprreY & Wooten; Hoim; KisAKUREK et al.; KOEK-NOORMAN; LUNK; PFEIFFER, RAD-
FORD et al.; SCHUMANN; SCOGGAN; oo & WILLIAMS; STEYERMARK (1963):
ToMLINson; Vines; and WELLS & SHAR
Bonner, F. T. Cephalanthus occidentalis L. Common buttonbush. U. S. Dep. Agr. Agr.
Handb. 450: 301, 302. 1974. [Includes photos of fruiting heads and of single fruits,
drawings of longitudinal section through fruit and of germination; also includes
comments on distribution, flowering and fruiting periods, and germination tests,
suggestions for collecting fruits.]
Britton, N.L. The button-bush a tree. Jour. New York Bot. Gard. 1: 54. 1900. [Includes
photo of arborescent individual in New York; also see TOMLINSON for mention o
arborescence in Florida.
. Cephalanthus occidentalis. Addisonia 5: 17, 18. pl. 169. 1920. [Includes color
plate, distribution, habitat, brief pre-Linnaean history, common names, use as feb-
rifuge, description, and native regions for other species. ]
Capuron, R. Sur Pidentité du Cephalanthus chinensis Lam. soreave II. 13: 471-
473. 1973 [1974]. [Breonia chinensis (Lam.) R. Capuron, comb. n
DEANE, W. Remarkable persistence of the button-bush. Rhodora 4: 343, 244. 1902.
[Plants thriving 37 years after being ae when a wet area was filled. ]
Duncan, W. H. Preliminary reports on the flora of Georgia. 2. Distribution of 87 trees.
Am. Midl. Nat. 43: 742-761. 1950. [C. nes 750, 761.]
FERNALD, M. L. Additions to and subtractions from the flora of Virginia (concluded).
Rhodora 49: 175-194. pls. 1078-1085. 1947. [C. occidentalis, 181, 182; new forma
lanceolatus.|
ae A. F. Notes on — buds and leaf scars. 2. Cephalanthus occidentalis.
t. Gaz. 20: 79. pl. 8, fig. 1. 1895.
Fox, w ie & J. H. Soper. The ae ies of some trees and shrubs of the Carolinian
of southern Ontario. Part IJ. Trans. Roy. Canad. Inst. 30: 3-32. 1953. [C.
veetdemai 3, 28, 29 (distribution map for southern Ontario), 30, 31; limited to
uthern part of Ontario.]
on G. D. eae of the tribe Naucleeae. Jour. Linn. Soc. Bot. 33: 1-94. pis.
—4, 1897.
Hom, T. Medicinal ae os North America. 54. Cephalanthus occidentalis L. Merck’s
Rep. 20: 216-218.
KAMMEYER, H. F. Die hae sfolum e. Mitt. Deutsch. Dendrol. Ges. 60: 97. 1957 [1958].
[Includes horticultural fee
V2 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Lima, O. A., & J. Potonsky. Les constituants flavonoides de Cephalanthus spathelliferus.
Phytochemistry 12: 913-916. 1973. [C. spathelliferus Baker = Breonardia micro-
cephala (Del.) Ridsd., fide RIDsDALE (p. 187).
VAUGH, R. Suggested phylogeny of Prunus serotina and other wide-ranging phylads
in North America. Brittonia 7: 317-346. 1952. [C. occidentalis, 343, 344; includes
distribution map for southwestern North America; C. salicifolius should be merged
with C. occidentalis. ]
PHILLIPSON, J., & S. HEMINGWAy. Indole and oxindole alkaloids from Cephalanthus
occidentalis. inal 13: 2621, 2622. 1974. [Also see PHILLIPSON ef a
,& C. E. Riwspate. The chemotaxonomic significance of alkaloids in
the Naucleeae s. 1 (Rubiaceae) Lloydia 45: 145-162. 1982. [Includes chemical data
on five species of Cephalanthus, among them C. occidentalis, which contains
akuammigine, tetrahydroalstonine, isopteropodine, pteropodine, speciophylline,
ajmalicine, isorhynchophylline (and its N-oxide), rhynchophylline (and its N- -oxide),
Se bea and hirsutine. Includes taxonomic implications. Also see
KISAKUREK ef @
Ripspace, C. E. A revision of the tribe Cephalantheae (Rubiaceae). Blumea 23: 177-
ROBERTSON, C. Flowers and insects. VI. Bot. Gaz. 16: 65-71. 1891. [Cephalanthus
occidentalis, 65-67; includes description of pollen presentation (cf. TOMLINSON),
records diverse insect visitors to flowers.
SPERRY, O. E. Poisonous range ees XVIII. St. Johnswort and buttonball bush. Sheep
& Goat Raiser 39: 20, 21.
, J. W. DoLLanite, G. A HorrMan, & B. J. CAmp. Texas plants poisonous to
livestock: 57 pp. + index. College Station, Texas. 1974(?). [Cephalanthus, 18.]
STRICKER, M. H. Cephalanthus—synonym for dependability. Am. Bee Jour. 89: 522,
523. 1949.* [Honey plant.]
TAytor, L. A. Plants used as curatives by certain south-eastern tribes. xi + 88 pp
Cambridge, Mae ge 1940. [C. occidentalis, 58; includes list of medicinal
uses. |
Tribe GARDENIEAE A. Richard ex DC. Prodromus 4: 342, 367. 1830,
““Gardeniaceae.”’
6. Randia Linnaeus, Sp. Pl. 2: 1192. 1753: Gen. Pl. ed. 5. 74. 1754.
Spiny [or unarmed] shrubs or small trees bearing opposite branches and short
shoots. Spines axillary, paired, sharp, stiff, usually inserted at ca. 45-degree
angle, generally shorter than leaves. Bark on twigs breaking up into conspicuous
untidy scales or taking the form of longitudinal flanges separated by long fis-
sures. Plants glabrous to strigillose [or more heavily pubescent] on twigs and
stipules. Leaves sessile or on short petioles, opposite or fascicled on short
shoots, small (not often longer than 3 cm), (infrequently) ovate to (frequently)
oblanceolate or obovate [sometimes trilobed], mostly rounded and mucronate
apically, the margins usually revolute when dry. Stipules with a single variably
shaped (usually deltoid and apiculate) lobe centered between adjacent petiole
bases, often split or shredded by growth of twig and/or by weathering. Plants
typically dioecious, the flowers subsessile in leaf axils, solitary or occasionally
clustered on short shoots among leaves, mostly pentamerous, imperfect, with
the nonfunctional organs reduced (or possibly flowers sometimes perfect, fide
Tomlinson) [or flowers perfect]. Calyx lobes variable in size and shape, deltoid
1987] ROGERS, CINCHONOIDEAE 173
to obovate [or foliose to suppressed], coalescent basally into a short tube.
Corolla white [or yellowish], cylindrical [flared or campanulate], the imbricate-
contorted lobes spreading and roughly as long as the tube, thickly pubescent
in and near the throat [or internally glabrous]. Stamens on very short filaments
in the corolla throat [or included or exserted]. Ovary inferior, usually bilocular;
style expanded and cleft apically into a pair of thick, exserted lobes [or undi-
vided]. Berries globose to ellipsoid, crowned with the persistent calyx, variably
reported as white or greenish to purple when ripe, the pulp dark toward the
inside. Seeds | or few, discoid. LEcTOTYPE SPECIES: R. mitis L. (see Britton, FI.
Bermuda, 361. 1918), this regarded by most modern authors as a synonym of
R. aculeata L., the only other species of Randia in the Species Plantarum.
(Named for Isaac Rand, ?-1743, British apothecary and botanist, director of
the Chelsea Physic Garden; for biographical notes see Trimen & Thiselton-
Dyer.)— INDIGO BERRY.
A rather vaguely defined genus usually estimated to have 200-300 species
and with a pantropical distribution (see below). Randia aculeata, the only
species indigenous to the area of the Generic Flora, occurs in South Florida at
the northern edge of its range, which extends to Mexico, Central America,
northern South America, and the West Indies. Texan populations are inter-
pretable as belonging to R. aculeata (for commentary see Vines, who tentatively
favored this stance), although Correll & Johnston and F. B. Jones referred them
to R. rhagocarpa Standley.
In our area Randia aculeata inhabits hammocks, shores (sometimes asso-
ciated with mangroves), oceanside dunes, pinelands, and thickets. The soil is
sometimes marly and is sometimes dry. As described by Tomlinson, the flow-
ers, chiefly borne April-June, are for the most part functionally imperfect by
abortion, although possibly some perfect ones may form. In 1966 Bremekamp
reported staminate flowers in some Gardenieae to have abortive styles that act
to hold pollen. The extent of involvement, if any, of the abortive styles in R.
aculeata in the pollination system is a question worthy of new observations.
Randia aculeata is recognized and differs from other genera of Rubiaceae
treated in this paper, except Catesbaea (see treatment of this genus for com-
parison), in being a shrub or small tree armed with paired axillary spines, each
of which diverges from the stem at roughly 45 degrees. Further, our Randia
has small, frequently apiculate leaves most often broadest above the middle
and usually fascicled on short shoots. The small flowers are solitary or clustered
on the short shoots. They have white, tubular corollas, and the thick stigmatic
lobes protrude from the pistillate flowers. The few-seeded, globose to ellipsoid
berries are conspicuously topped by calyx remnants.
Defining Randia from a global perspective is hard to accomplish. At present
the generic boundaries remain unsettled, especially in the Old World. Authors
disagree severely in their generic circumscriptions and synonymy. In a treat-
ment fundamental to taxonomic accounts that followed, Bentham & Hooker
conceived of Randia as polymorphic, pantropical, and made up of about 90
species in six sections. They named a new genus allied to Randia, Basanacantha
J. D. Hooker, which they thought to differ in being dioecious (an invalid
174 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
distinction), and in having glumaceous stipules, terminal flowers, membra-
naceous leaves, and other distinctive characters. Schumann held nearly the
same concept of Randia but added a seventh section
Critical of Schumann’s treatment, Fagerlind regretted that Randia had be-
come a “refuse dump” for Gardenieae of uncertain position. Emphasizing
branching relationships and using diverse additional characters, he pruned
Randia back to Schumann’s sect. Eurandia (sect. RANDIA), emended this, added
Basanacantha, and limited Randia to American species. Even if Fagerlind’s
work has not been particularly influential, the merger of Basanacantha with
Randia has been supported by a number of later authors (see especially Stand-
ley, 1919), and it is more or less in harmony with a tendency among recent
authors to transfer Old World species from Randia to other genera
Concentrating on West African species, Keay dismissed Fagerlind’s taxo-
nomic conclusions about Randia as “not altogether satisfactory,” stressed the
need (that persists) for a full revision, and recognized as distinct 21 genera,
“all of which have at one time or another been included, wholly or partly, in
Randia or Gardenia.’ Keay listed new or resurrected generic placements for
126 species previously included in Randia. More recently, Hepper & Keay
attributed no species to Randia in the Flora of West Tropical Africa. Tirven-
gadum, after considering “practically all taxa described under Randia,” likewise
confined the genus to America and characterized it as having [paraphrased]
unilocular ovaries with parietal placentae, a nonwaxy bluish pericarp, imperfect
flowers, pollen grains remaining in tetrads, and distinct testa cells, and as lacking
serial bud formation. (In contrast with Tirvengadum, American floristic authors
tend to describe the ovary as generally bilocular.) Yamazaki sorted the Asian
species out among five other genera, leaving none in Randia. However, it must
be emphasized that acceptance of such exclusive boundaries is not unanimous.
Authors working on floras in the New World (Standley; Standley & Williams;
Steyermark; Dwyer) have regarded Randia as pantropical but have avoided
assertions about its limits. They have not adopted infrageneric categories, ex-
cept that Williams and Standley & Williams recognized subgenus BASANA-
CANTHA (J. D. Hooker) L. O. Williams, which they distinguished from subg.
RANDIA by the former’s longer corollas, larger fruits, more often imperfect
flowers, and terminal quartets of spines (vs. spines paired and scattered). Like
Keay, they stressed the need for revisionary work, not only in terms of delim-
iting the genus, but also of redefining our R. aculeata, which they perceived as
too inclusive.
Randia aculeata has been used as a folk remedy for dysentery, and the fruit
has been the source of a blue dye. Fruits from at least one extraregional species
have served as food for humans. Randia formosa (Jacq.) K. Schum. is cultivated
as an ornamental in tropical regions, and it yields an essential oil used in
making perfume (see Prance & Da Silva for an illustrated account of this
species)
REFERENCES:
Under subfamily references see BENTHAM & Hooker; BREMEKAMP (1966); a cee
& CORRELL; CORRELL & JOHNSTON; Dwyer; HALLE; HEPPER & Keay; F. B. Jones;
1987] ROGERS, CINCHONOIDEAE 175
SCHUMANN; STANDLEY (1918); STANDLEY & WILLIAMS; STEYERMARK (1972, 1974);
TOMLINSON; VERDCOURT (1958, 1976); and VINES.
Boruipr, A. Rubiaceas cubanas, I. Randia L. y Shaferocharis Urb. Acta Bot. Acad. Sci.
Hungar. 27: 21-36. 1981. [Revision of six Cuban species, including R. aculeata, and
description of three new ones; discussion of infraspecific variation in R. aculeata.]
DEWo Fr, G. P. Randia for southern gardens. Baileya 2: 46. 1954. [R. macrantha, R.
macrophylla.]
FAGERLIND, F. Die Sprossfélge in der Gattung Randia und ihre Bedeutung fiir die
Revision der Gattung. Ark. Bot. 30(3, paper 7): 1-57. 1943. [R. mitis (R. aculeata),
22-24, 30, fig. 8, a, j; systematic conclusions summarized, 41-43.
Hume, E. P. Puerto Rico’s Christmas tree. Bull. New York Bot. Gard. 49: 284-287.
1948. [R. aculeata.]
Keay, R. W. J. Randia and Gardenia in West Africa. Bull. Jard. Bot. Bruxelles 28: 15-
72. 1958. [Includes Seecics of generic circumscription. ]
Opter, P. A., & K. S. Bawa. Sex ratios in tropical forest trees. ENO ae ae 812-821.
1978. [R. subcordata, staminate/pistillate = 2.04; R. spinosa, s/p =
PELLEGRIN, F. Une Rubiacée du Gabon qui sert a narcotiser le poisson. ra Bot.
ppl. Agr. Trop. = 498-501. 1938.*
ee G. T., & M. F. pa Sitva. Arvores de Manaus. 312 pp. Manaus. 1975. [R.
tomentosa, 226- on
Quersu, M. A., & R. S. THAakur. Chemical constituents of Randia tetrasperma. Pl.
Med. 32: 229-232. 1977.*
SAHaRIA, G. S., & V. SESHADRI. Chemical investigation on Randia saponins. Isolation
and characterisation of randioside A beta-D-galactopyranosyl (1 to 3)-oleanolic acid.
Indian Jour. Forestry 3: 6-8. 198
SaINTY, D., P. DELAVEAU, F. BAILLEUL, & C. Moretti. 10-cafeyl desacetyldaphylloside,
noavel iridoide de R andia formosa. Lloydia 45: 676-678. 1982 [1983]. [In addition
to iridoid in title, ioe gardenoside, and deacetylasperulosidic acid are foun
in bark.]
STANDLEY, P. C. A note concerning the genus Randia, with descriptions of new species.
Contr. U. S. Natl. Herb. 20: 200-203. 1919. [Basanacantha merged with Randia.]
TIRVENGADUM, D. D. A synopsis of the Rubiaceae-Gardenieae of Ceylon (Sri Lanka).
ull. Mus. Hist. Nat. Paris III. Bot. 35: a= 33. ill
& C. SAstReE. La signification ification de Randia
et genres affinés. Mauritius Inst. Bull. 8: oe 98. 1979.
TrimMEN, H., & W. T. THIsELTON-DyerR. Flora of Middlesex. Map + xli + 428 pp.
London. 1869. [Isaac Rand, 388, 389.]
UesatTo, S., E. Aut, H. NisHmmura, I. KAwAMURA, & H. INouye. Four iridoids from
Randia canthioides. Phytochemistry 21: 353-357. 1982. [New iridoids: 10-dehy-
drogardenoside (artifact?), dimeric dehydrogardenoside, randioside, deacetylas-
perulosidic acid methyl ester aglycone. Previously known iridoid glucosides: gar-
denoside, deacetylasperulosidic acid methyl ester, scandoside methyl ester. Authors
interpret results as supporting placement of Randia among Gardenieae. ]
WILDEMAN, E. pe. La myrmécophilie du Randia eetveldeana De Wild. et Dur. (Rubi-
- ées). Bull. Acad. Roy. Sci. Belg. Cl. Sci. 18: 52-58. 1932. [Keay placed species of
andia mentioned in this paper in synonymy as follows: R. eetveldeana = Roth-
mannia Whitfieldii (Lindley) Dandy, Randia Lujae = Rothmannia Lujae (De Wild.)
Keay, Randia myrmecophylla = Rothmannia macrocarpa (Hiern) Keay, Randia
A Ae ee = Gardenia imperialis K. Schum
WILLIAMS, L.O. Randias from Central America. Phytologia 24: 159-163. 1972. [Includes
comments on nitereatonsip and peace of Central American species;
subg. Basanacantha (J. D. Hooker) L. O. Williams, comb. n ov.]
YAMAZAKI, T. A revision of the genus Randia L. in eastern eh Jour. Jap. Bot. 45:
BTA, 1970.
176 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
7. Casasia A. Richard in Sagra, Hist. Fis. Cuba. ed. 2. 11: 9. 1850.
Dioecious shrubs or small trees with thick, glabrous twigs covered with light-
colored flaking bark. Leaves clustered toward branch tips, glabrous except for
axillary tufts of trichomes abaxially, petiolate, obovate or oblanceolate, truncate
or emarginate to obtuse or rounded at the apex, cuneate to caudate at the base;
stipules with the single lobe centered between adjacent petioles, oblong to
deltoid or ovate, acute or acuminate and sometimes apiculate apically, fre-
quently denticulate along the margins, the adaxial side with colleters. Flowers
fragrant, on tapered pedicels, tending to blacken upon drying, imperfect with
the nonfunctional organs (gynoecium or stamens) developing and with sta-
minate and carpellate flowers superficially fairly similar. Staminate flowers in
terminal, compound, monochasial or partly dichasial inflorescences: bracts
scalelike, highly irregular in shape. Carpellate flowers solitary and terminal,
often overtopped and thereby left in lateral position. Calyx made up of a cup-
shaped tube topped with 5 finger-shaped to filiform [to deltoid] lobes about as
long as to twice as long as the tube, the lobes frequently hooked or curled at
the tips when dry. Corolla much longer than calyx, white [or yellow], salverform,
with 5 lanceolate or narrowly deltoid lobes as long (or nearly as long) as the
slender corolla tube, imbricate-contorted in bud, often hispid-serrulate along
apical margin. Stamens inserted in throat of corolla on very short filaments;
anthers linear. Ovary unilocular, with 2 (or 3) intrusive, parietal placentae;
style rising to throat of corolla tube, expanded apically and divided into 2 (or
3) lobes. Fruit ovoid or ellipsoid, roughly the size of a hen’s egg or more nearly
globose, tapered at base, spotted on the surface, crowned with the thickened
calyx tube, the sclerified endocarp covered by a tough exo- and mesocarp, the
large internal cavity filled with the fleshy placentae in which are embedded
numerous black (dry), compressed seeds stacked horizontally or obliquely in
the fleshy matrix and having pebbled testae. Type species: C. calophylla A.
Richard, the only species known when the genus was established. (Named for
Sr. D. Luis de las Casas, Captain General of Cuba.)— SEVEN-YEAR-APPLE,
A genus of perhaps 11 species in Florida, the West Indies, and Mexico:
Casasia Acunae Fernandez & Borhidi (Cuba); C. calophylla A. Rich. (Cuba);
C. chiapensis Miranda (Chiapas, Mexico); C. clusiifolia (Jacq.) Urban (Ber-
muda, Bahamas, Florida, Cuba); C. domingensis Urban (Hispaniola), C. Ek-
mani Urban (Hispaniola); C. haitiensis Urban & Ekman (Hispaniola); C. jac-
quinioides (Griseb.) Standley (Cuba; C. parviflora Britton, synonymy fide
Standley); C. /ongipes Urban (Jamaica; C. piricarpa Urban, synonymy fide
Adams); C. nigrescens (Griseb.) C. Wright ex Urban (Cuba); and C. Samuels-
sonit Urban & Ekman (Hispaniola). (It should be noted that this list comes
from an uncritical examination of the literature and from the Gray Herbarium
Card Index. The only herbarium materials that I have studied, except for the
survey of stipules mentioned below and extralimital specimens of C. clustifolia,
originated in the area of the sreneie Flora.) Our C. clusiifolia (Randia clusiifolia
(Jacq.) Chapman, Genipa clusiifolia Jacq.) is by far the most widespread species,
occurring in our area ae in the Florida Keys, but also as far north along
the coast as Lee County, Florida.
1987] ROGERS, CINCHONOIDEAE 177
FiGure 2. Casasia. a-l, C. clusiifolia: a, branch from staminate plant, showing par-
tially cymose inflorescence, x '4; b, staminate flower, x 1; .c, opened corolla of staminate
ovules, x 1; e, branchlet from carpellate plant with single floral bud and fruit, x 4; f,
carpellate flower, x 1; g, opened corolla of carpellate flower with nonfunctional stamens,
x 1; h, tricarpellate gynoecium, ovary in longitudinal section to show | of 3 placentae,
x 1:1, view from axis of portion of spongy placenta showing partially embedded ovules,
x 5; j, semidiagrammatic cross section of tricarpellate ovary with 3 parietal placentae,
x 2;k, longitudinal section of bicarpellate fruit, 1 placenta sectioned to show embedded
seeds, x '4; 1, longitudinal section of seed with embryo embedded in abundant endo-
sperm, x 2.
Casasia clusiifolia tolerates high salinity and lives in coastal scrub and ham-
mocks in our area. Flowers form throughout the year, but mostly during spring
and summer. In Florida Tuskes observed that the moth Aellopos tantalus uses
this species as a larval food plant, evidently along with at least Annona glabra
L. Almost every plant that he examined showed signs of the moth.
As a whole, the genus Casasia is made up of small trees or shrubs with
terminal cymose inflorescences (or solitary carpellate flowers), conspicuous
white or yellow flowers that blacken upon drying, cupular calyces with subulate
to deltoid lobes, salverform corollas with the lobes twisted in bud, stamens on
short filaments in the corolla throat, included or nearly included anthers, in-
cluded stigmas, intrusive parietal placentae bearing numerous embedded ovules,
arge berries with tough pericarps containing numerous more or less horizontal
178 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
seeds in a fleshy matrix, corneous endosperm, and foliaceous cotyledons. Ad-
ditional useful characters for our species are its usually obovate or oblanceolate,
coriaceous leaves clustered toward the tips of thick twigs, staminate flowers in
compound monochasia, solitary carpellate flowers, and large, mottled fruits
crowned with a much-thickened calyx cup. The parietal placentae filling the
ovary make it appear bi- or sometimes trilocular. Most of the published illus-
trations show either staminate inflorescences or the fruit, seldom the solitary
carpellate flowers.
Probably the most closely related genus is Genipa L., which throughout the
literature is held to differ from Casasia in having lateral (vs. terminal or mostly
terminal) inflorescences. Urban (1908) further separated Genipa by its inter-
nally sericeous (vs. glabrous) calyx limb, this entire or with obtuse lobes (vs.
lobes filiform to acute), pubescent corolla, exserted anthers and style, and thick
(vs. linear) stigmas. A modern reevaluation of these differences is desirable.
Despite indications to the contrary in the literature, herbarium material at the
Missouri Botanical Garden showed no difference between Casasia and Genipa
in the position of the stipules. One lobe of the interpetiolar stipules is centered
between adjacent petiole bases in both, as it is in most Rubiaceae.
e genus needs a full revision. Schumann’s treatment in the Nattirlichen
Pflanzenfamilien is based on only one (or perhaps two) species. The principal
accounts are those by Standley (North American Flora, 1918), Fernandez Ze-
queira & Borhidi, and Urban (1908, 1927). In addition, Miranda’s surprising
report of the only continental species should not be overlooked.
REFERENCES:
Under subfamily references see ADAMS; ALAIN; BARKER & DARDEAU; CORRELL &
CoRRELL; LoNG & LAKELA; SCHUMANN; STANDLEY (1918); and TOMLINSON
FERNANDEZ ZEQUEIRA, M., & A. Boruipi. Rubiaceas cubanas IJ-HI. I. El género Casasia
A. Rich. en Cuba. Acta Bot. Acad. Sci. Hungar. 28: 81-85. 1982.
MARIE- ee Fr. [C. KrrouAc], & Fr. LEON [J. 8. SAuGET]. Itinéraires botaniques
dans l’ile de Cuba. Contr. Inst. Bot. Univ. Montréal 50: 1-410. 1944. [C. clusiifolia,
105 (photo of fruiting plant), 106.]
MIRANDA, F. Plantas nuevas de Chiapas. Ceiba 4: 126-145. 1953. [C. chiapensis, 142-
145
Tuskes, P. M. The life history of Ael/opos tantalus (Sphingidae). Jour. Lepidopt. Soc.
34: 327-329. 1980. [C. clusiifolia in Florida a larval food plant for this moth, 328.]
UrBan, I. Casasia. Symbolae Antillanae 5: 504-507. 1908. [Includes generic description
and five species.]
Plantae Haitienses novae vel rariores IV. a cl. E. L. Ekman 1924-26 lectae.
Ark. Bot. 21A(5): 1-97. 1927. [Three new species of Casasia, 73-78; also see bid.
24A(4): 45. pl. 2. 1931.]
8. Hamelia Jacquin, Enum. Syst. Pl. Carib. 2. 1760.
Shrubs with raphide bundles often conspicuous in several organs, pilose to
puberulent throughout (except sometimes becoming glabrate with age). Leaves
opposite or ternate, petiolate, (oblanceolate to) elliptic (to ovate-lanceolate),
with several pairs of pinnate nerves, usually acute or acuminate at both ends;
1987] ROGERS, CINCHONOIDEAE 172
stipular lobes single between adjacent petioles, narrowly deltoid to subulate.
Inflorescence terminal, roughly pyramidal or somewhat flat topped, usually
consisting of long, uncrowded cincinni (or occasionally dichasia) in cymose
clusters, these not infrequently in thyrsiform arrangements and often with
multiple orders of branching. Flowers pentamerous. Calyx lobes low, deltoid,
inconspicuous. Corolla red or orange, slender and nearly cylindrical but con-
stricted near the base, the lobes deltoid, only a small fraction of the length of
the tube. Stamens inserted on the corolla tube near its base; anthers linear and
very long (over half the length of the corolla tube and somewhat longer than
the filaments), partly exserted (or sometimes included?), sagittate at base. Style
filiform, expanded and papillose in the upper 4 of its length at the mid-level
of the anthers. Ovary topped with a conical disc around the base of the style,
usually 5-loculate, each locule containing numerous anatropous ovules on axile
placentae. Fruit a berry, red before becoming black, ellipsoid, conspicuously
crowned with a disc (this sometimes taking the form ofa beak) and the persistent
calyx. Seeds numerous, small, longer than broad, irregularly shaped, usually
angular, coarsely reticulate. LecroTYPE species: Hamelia erecta Jacq. (= H.
patens Jacq., the only other species included in the protologue; see Wernham,
London Jour. Bot. 49: 206. 1911; Britton & Millspaugh, Bahama FI. 411. 1920;
and Elias, Mem. New York Bot. Gard. 26: 112. 1976 for lectotypification and
for choice of epithets when the two species are merged). (Named for Henri
Louis Duhamel du Monceau, botanist, 1700-1 782.)— FIREBUSH.
A genus of about 16 woody species in two sections distributed in tropical
and subtropical America and concentrated in Mexico and Central America. A
representative of section HAMELIA, Hamelia patens, 2n = 24, is the only species
indigenous to the continental United States. The range of H. patens var. patens
extends from Lake County, Florida, southward through the West Indies, much
of Mexico, Central America, and (mostly western) South America to northern
Argentina and Chile. A second variety, H. patens var. glabra Oersted, 1s limited
to Central America and northern South America.
In Florida Hamelia patens var. patens most frequently grows in coastal
hammocks, although it sometimes occurs inland and has weedy tendencies,
turning up in sunny, disturbed places. In tropical America it is common, a
pioneer in clearings and a weed, and is cultivated ornamentally. It is also
cultivated in the Old World, no doubt escaping there as well. Flowering takes
place throughout the year in our area. Bawa & Beach found the flowers to be
monomorphic, and they found selfing to yield reduced fruit set, with fruits
aborting.
Hamelias are recognized as shrubs or small trees with often secund, red to
yellow, frequently angular, tubular flowers with imbricate aestivation and long,
linear anthers. The typically five-locular ovary is topped with a persistent, often
beaklike disc. The berries contain numerous flattened seeds. Hamelia patens
var. patens is easily separated from all other Rubiaceae in our area by its long,
narrow, tubular, orange or red flowers with an inconspicuous calyx and short
corolla lobes.
Schumann placed Hamelia in his large tribe Gardenieae within subfam.
180 JOURNAL OF THE ARNOLD ARBORETUM
FiGurE 3. Hamelia. a-j, H. pate
x Ib: b, me with bases of petioles of3 leaves and interpetiolar ae 2 axillary buds
visible, x 3; c, portion of inflorescence, x 2; d, flower in longitudinal section—note
epipetalous stamens, ae anthers, and axile placentation, < 3; e, adaxial side of anther
and portion of filament, x 4; f, style with stigmas, x 4; g, diagrammatic cross section
of ovary, showing ae placentae with numerous ovules, x 6; h, fruit, a berry, x
seed, x 25; j, seed in te eae section, seed coats unshaded and hatched, eae
stippled, ian unshaded, 0.
Cinchonoideae, a subfamilial and tribal position not generally accepted by
subsequent authors. Stressing the presence or absence of raphides in distin-
guishing the Rubioideae from the Cinchonoideae, Bremekamp (1966), Verd-
court (1958), and Elias positioned Hamelia in the Rubioideae, where they all
acknowledged, however, that it is anomalous in having imbricate, rather than
valvate, aestivation.
At the tribal level, Bremekamp (1966) paired Hamelia with Hoffmannia Sw.
1987] ROGERS, CINCHONOIDEAE 181
as the tribe Hamelieae, which Elias adopted in his revision of Hamelia, as did
Standley & Williams. According to Elias, Hamelia and Hoffmannia are linked
by their woody habit, raphides, imbricate aestivation, ovarian discs, two- to
five-locular ovaries, numerous ovules per locule, and baccate fruits. Except for
multilocular ovaries, these features are fairly generalized in the Rubiaceae;
however, Elias also noted without elaboration similarities in their pollen and
seeds. He distinguished Hamelia from Hoffmannia by the former’s occupying
lower altitudes and by its having terminal (vs. axillary), usually monopodial,
more often paniculate inflorescences generally containing more flowers, usually
unribbed and secund corolla tubes, pentamerous (vs. usually tetramerous) flow-
ers, most often 5 (4) locules (vs. usually (4) 3 or 2 locules) in the ovary,
stamens inserted lower in the tube, and sagittate anthers. With only a small
number of chromosome counts in hand so far, Hamelia appears to have 2n =
24, while only 2” = 48 is known in Hoffmannia.
Steyermark (1974) accepted the tribe Hamelieae but differed from Breme-
kamp and Elias by including the genus Bertiera Aublet, which—unlike Hoff-
mannia and Hamed has contorted aestivation and lacks raphides. Dwyer,
too, associated H. in the Hamelieae but with Yerococcus
Oersted, which stands apart in having valvate aestivation.
Hamelia has been revised twice in this century. Wernham recognized 28
species in 1911; Elias accepted 12 of these in 1976, changing the name of one,
which was a later homonym. Most of the remainder fell into synonymy, a large
cluster under the two varieties of H. patens. Elias added three species discovered
since Wernham’s study, bringing the total number in his revision to 16, sorted
into two sections of eight species each.
The pollen of Hamelia patens is tricolporate, with circular ora and with an
areolate, tegillate sexine (Anand & Bhandari).
Beyond being ornamental, Hamelia patens has edible berries used in Mexico
for preparing a fermented beverage (Standley). Having a high tannin content,
the bark has been used in tanning leather (Morton, Standley). As Morton
documented, this species has multiple applications in folk remedies, mostly to
counter dysentery and to treat skin wounds and irritations.
REFERENCES:
Under subfamily references see BREMEKAMP (1966); Dwyer; LONG & LAKELA; MorTON;
SCHUMANN; STANDLEY (1926); a & WILLIAMS; STEYERMARK (1974); TOMLINSON;
VERDCOURT (1958); and WUNDER
ANAND, S. K., & M. M. BHANDARI. Pollen morphology of Rubiaceae from Mount Abu
(Rajasthan). Jour. Econ. Taxon. Bot. 4: 335-342. 1983. [H. patens, 336, 338, 339;
cultivated or escaped, if accurately identified.]
Bawa, K.S., & J. H. BEAcH. Self-incompatibility systems in the Rubiaceae of a tropical
lowland wet forest. Am. Jour. Bot. 70: 1281-1288. 1983. [H. patens, 1282, 1283.]
Borcers, J., & A. RUMBERO. Two new oxindole alkaloids isolated from Hamelia patens
Jacq. Tetrahedron Lett. 20: 3197-3204. 1979.*
BORGES DEL CASTILLO, J., J. L. dee Ramon, L. F. RoDRIGUEZ, P. VAZQUEZ BUENO,
. MANRESA FERRERO. Two more new oxindole alkaloids of Hamelia patens.
Ann. Quim. Ser. C. 76: 294, 295. *1980. * [Title given here probably translated from
Spanish. ]
182 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
Britton, N.L. The genus Hamelia Jacq. Torreya 12: 30-32. 1912. [Includes ca aoa
of H. scabrida Britton and comments on WERNHAM’S revision of fameli
Euias, T. S. A monograph of the genus Hamelia (Rubiaceae). Mem. New York Bot.
Gard. 26: 81-144. 1976.
Rippercer, H. Isolation of isopteropodine from Hamelia patens. Pharmazie 32: 415—
441. 1977.*
SCURFIELD, G., A. J. MICHELL, & S. R. Si-va. Crystals in woody stems. Bot. Jour. Linn.
oc. 66: 277-289. pls. 1-7. 1973. [H. patens, 278, 280, 286; includes scanning
electron micrograph of raphides, p/. /c.]
SHARMA, M. A comparative study of sclereids in some members of the Rubiaceae. Proc.
Indian Natl. Sci. Acad. B. 36: 289-296. 1970. [H. patens, see especially p. 290;
sclereids absent; bast fibers and sclerotic pith present.]
SUBRAHMANYAM, K., J. M. RAo, & K. V. J. Rao. Chemical examination of Hamelia
patens (Rubiaceae). Curr. Sci. Bangalore | 42: 841. 1973. [Malvidin, betunidin, ae .
tosterol, ursolic acid, and 8-sitosterol-D
WERNHAM, H. F. A revision of the genus Hamelia. Jour. Bot. London 49: 206- a
1911. [Also see /bid. 346 for addendum, and see ELIAs.]
9. Catesbaea Linnaeus, Sp. Pl. 1: 109. 1753; Gen. Pl. ed. 5. 48. 1754.
Spiny shrubs [small trees or scandent shrubs] with puberulous branches often
inserted at oblique angles. Leaves opposite or fascicled on short-shoots, gla-
brous, sessile or on short petioles, small (mostly under | cm long in our species)
[sometimes virtually absent by reduction]. Spines stiff, sharp, frequently longer
than leaves, paired, generally arising at oblique angles. Stipular lobes initially
solitary between adjacent petiole bases, quickly cleft into 2 lobes, disappearing
during expansion of twig. Flowers borne singly among leaves, on short pedicels,
small and inconspicuous [or large and showy], tetramerous. Calyx lobes per-
sistent, subulate, longer than ovary. Corolla white, the tube narrowing toward
ase, the valvate and deltoid lobes much shorter than tube. Stamens inserted
at base of corolla tube, rising to level of lobes. Ovary bilocular, with ovules
on faces of septum [or on placentae arising from septum]; stigma bifid. Berries
globose, white (or black), containing a small number of compressed seeds with
rugose surfaces. TyPE SPECIES: C. spinosa L., this the only species in the generic
protologue. (Named for Mark Catesby, 1683-1749, British naturalist, known
in part for his The natural history of Carolina, Florida, and the Bahama Is-
lands.)
A genus of approximately 15 species in the West Indies, one of them reaching
the Florida Keys. Most are known from only a single island each, although
Catesbaea spinosa L., 2n = 24, C. melanocarpa Urban, and C. parviflora Sw.
occur on a number of islands. Cuba has the greatest number of species—about
seven endemics, in addition to two more widespread species. There are about
six endemics on Hispaniola. On all other islands where it occurs, Catesbaea 1s
limited to one or two species. Catesbaea parviflora, the most broadly distributed
species, grows on the Florida Keys, the Bahamas, Cuba, Jamaica, Puerto Rico,
Antigua, the Cayman Islands, and undoubtedly other islands. In Florida C.
parviflora is encountered in dry, open areas. Its habitats include pine woods,
edges of hammocks, and sand dunes.
With its conspicuous paired thorns and small, clustered leaves widest above
1987] ROGERS, CINCHONOIDEAE 183
the middle, Catesbaea is easily recognized among shrubs in our area, although
it might be confused with Randia. Catesbaea usually has tetramerous flowers
(vs. pentamerous ones in Randia), valvate (vs. contorted) aestivation, and
stamens inserted basally in the corolla (vs. in the throat in Randia and Hoff-
mannia). As Proctor pointed out, our species of Catesbaea has smaller fruits
than our species of Randia (4 mm vs. 8-12 mm in diameter). Additional
distinguishing features of Catesbaea include bilocular ovaries (vs. five-locular
in Hamelia), perfect flowers (vs. imperfect ones in Bertiera and Randia acu-
leata), and solitary, axillary flowers
erdcourt diverged from Schumann in placing Catesbaea outside of the
Gardenieae in the segregate tribe Catesbaeeae J. D. Hooker, which he regarded
as close to the Gardenieae. According to him, distinguishing features of the
Catesbaeeae are valvate aestivation (vs. contorted or imbricate in the Garde-
nieae), usually spiny branches, and fleshy fruits containing rugose seeds ad-
hering in a mass.
Catesbaea is in need of revision. The only comprehensive treatment is Stand-
ley’s (1918). Taxonomy of the genus rests on this, coupled with regional floristic
orks.
Catesbaea spinosa, which has large, showy flowers, is cultivated as an or-
namental shrub.
REFERENCES:
Under subfamily references see ADAMS; ALAIN; ay er CORRELL & CORRELL;
PROcTOR; SCHUMANN; STANDLEY (1918); and VERDcouRT (195
Gituis, W. T. Phantoms in the flora of the Bahamas. Phytologia 29: 154-166. 1974.
[Catesbaea, 161, 162; C. campanulata, C. parviflora var. septentrionalis, C. fasci-
culata, and C. foliosa (but see CoRRELL & CorReLL) all in synonymy under C.
parviflora; also see Rhodora 76: 67-138. 1974.]
PANDEY, D.S. Notes on teratology of certain angiosperms. Bull. Bot. Survey India 21:
121-124. 1979 [1981]. [C. spinosa, 121-123; some flowers with parts in threes, some
with petaloid sepals.]
Raman, V. S., & P. C. KesAvAN. Chromosome numbers of some dicotyledons. Sci.
Cult. 29: 413, 414. 1963.
AL-SHEHBAZ, ALYSSEAE 185
THE GENERA OF ALYSSEAE
(CRUCIFERAE; BRASSICACEAE) IN THE
SOUTHEASTERN UNITED STATES!”
IHSAN A. AL-SHEHBAZ?
Tribe Alysseae A. P. de Candolle, Syst. Nat. 2: 147, 280. 1821, “Alyssineae.”
Annual, biennial, or perennial herbs [sometimes subshrubs, shrubs, or even
trees]; usually with stellate, dendritic, cruciform, or furcate trichomes, rarely
strongly differentiated into blade and claw. Nectar glands distinct or connate.
Stamens usually 6, often tetradynamous; filaments with or without wings, teeth,
or appendages. Fruits usually less than 3 times as long as wide, dehiscent [or
rarely indehiscent], inflated or most commonly flattened parallel to the septum
(latiseptate), sessile or long stipitate; valves usually 1-nerved, glabrous or with
1 or more types of trichome; septum present or absent; styles long to obsolete;
stigmas entire to 2-lobed. Seeds | to numerous, usually biseriately arranged in
each locule, mucilaginous or not when wet, broadly winged to wingless; funicles
| Cl £4}
'Prepared for S United States, a long-term project made possible
by grants from the National Science Foundation and currently supported by BSR-8415769 (C. E.
Wood, Jr., principal investigator), under which this research was done, and BSR-8415637 (N. G.
of this area, with information about extraregional members of a family or genus in brackets. The
references that I have not verified are marked vain asterisks.
I am most indebted to Carroll Wood for d advice d th aration
of this paper, and especially for his critical review of the manuscript. I am grateful io Red a as
for allowing study his manuscripts on the genera Draba and Lesquerella for his forthcoming
book on the Cruciferae of North rica. | am variously indebted to Norton iller, George K
Rogers, R HONES gpd meee oli as eel to Barbara Nimblett, who typed the
manuscript. I ongberg for their editorial advice
p
Some of the wstratons (Figures i Jj, k; 2f, 1, j) were mate by Karen Stoutsenberger (KS) under
earlier pew Carroll Wood prepared the material and supervised the illustrations. The remaining
illustrations were drawn by me (IAS). The fruits and seed fi herbari i in the Gray
H rium and Arnold Arboretum.
*For an account aoe family and its tribes, see Al-Shehbaz, The tribes of Cruciferae (Brassicaceae)
in ube southeastern United States. Jour. Arnold Arb. 65: 343-373. 1984.
Arnold Arboretum, Harvard University, 22 Divinity Avenue, Canpages Massachusetts 02138.
© President and Fellows of Harvard College, 1987
Journal of the Arnold Arboretum 68: 185-240. ae 1987.
186 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Figure 1. Selected representatives of tribe Alysseae. a—c, Berteroa incana: a, portion
of plant with eae nd fruits, x ‘4; b, fruit with rectangular portion of valve showing
trichomes, x 5; c, petal, x 5. d, e, Camelina microcarpa: d, portion of infructescence,
x |: e, fruit, x 5—note beaklike apex of valve. f, g, A/yssum Alyssoides: f, fruit with |
on sepal removed, x 5—note filiform nectar glands; g, fruit with | valve and all
pals remov ae x 5—note subapical placentae. h, i, Lumaria annua: h, septum and
sre x |—note gynophore and adnation of funicles to septum; i, seed, x 3. J, k,
Lobularia ana j, fruit, x 6; k, replum and septum, x 6
free or adnate to the septum, apically or laterally attached to the replum;
cotyledons accumbent. (Including Camelineae DC., Drabeae O. E. Schulz,
Lunarieae O. E. Schulz.) Type Genus: A/yssum L.
A poorly defined tribe with some 40 genera (15 monotypic) and about 650
species (excluding Lesquerella S. Watson) distributed primarily in the Irano-
Turanian (ca. 210 species) and Mediterranean (ca. 110 species) regions. The
majority of species belong to two genera: Draba L. (350) and Alyssum (170).
Except for Draba, the Alysseae are poorly represented in Siberia, eastern Asia,
and North America and are absent in the Southern Hemisphere and in the
arctic and Himalayan regions. The tribe is represented in the southeastern
United States by seven genera and 20 species, of which 13 are indigenous.
The limits of the Alysseae adopted here closely follow Janchen’s classifica-
tion, which unites the tribes Lunarieae, Drabeae, and Alysseae of Schulz. Jan-
1987] AL-SHEHBAZ, ALYSSEAE 187
chen followed Von Hayek in treating the first two as subtribes of the last. Both
Selenia Nutt. and Armoracia Gaertner, Meyer, & Scherb., which were placed
by Schulz in the Lunarieae and the Drabeae, respectively, will be treated in
the Arabideae DC., where their nearest relatives are usually placed. The South
African Schlechtera Bolus, treated in the Lunarieae by Schulz, has diplecolobal
embryos (with cotyledons twice transversely folded) and should therefore be
included with its allies of the Heliophileae DC. Lesquerella was placed by
Schulz in the Drabeae, but as is clearly shown below, it should be placed with
its nearest generic relatives in the Lepidieae. However, it is treated here in the
Alysseae, as shown in the outline adopted by Al-Shehbaz (1984).
Schulz separated the Lunarieae from the Alysseae mainly on the basis of
simple vs. branched or stellate trichomes. Both Ricotia L. and Peltaria Jacq.
(including Leptoplax O. E. Schulz), which he placed in the former tribe, have
members with simple or branched trichomes, as do numerous other genera of
the Cruciferae. Therefore, the type of pubescence alone cannot be used as the
basis for tribal delimitation. Similarly, the cellular pattern of the fruit septum,
considered by Schulz to be the main difference between the Alysseae and the
Drabeae, is an unreliable feature and should not be overemphasized. Many
authors (e.g., De Candolle, 1821, 1824; Von Hayek; Janchen) placed the core
genera Draba, Alyssum, and Lunaria L., as well as their immediate relatives,
in the tribe Alysseae, a disposition I presently support. It is clear, however,
that the tribal classification of the Cruciferae is inadequate, and further studies
may alter the boundaries of the Alysseae. Knights & Berrie found that data
from sterols support the placement of Lunaria but not Draba in the Alysseae.
Chromosome numbers are known for some 275 species (ca. 43 percent of
the tribe) and 28 genera (excluding Lesquere/la). Nearly 80 percent of the species
surveyed have chromosome numbers based on eight, and only about seven
percent have numbers based on seven (author’s compilation). About 50 percent
of the species are diploid, and nearly 38 percent are exclusively polyploid.
Polyploidy occurs in nearly 60 percent of the species of Draba. Aneuploidy
and polyploidy probably played important roles in the evolution of Lobularia
Desv. and Hormathophylla Cullen & T. R. Dudley.
The Alysseae are almost exclusively herbaceous; only a few species in three
genera are woody. Some species of A/yssum and Hormathophylla, particularly
those growing in the eastern Pyrenees, southern France, and eastern Spain, are
subshrubs or shrubs to 50 cm high. Farsetia Turra has the most diversified
habit of any genus of the Cruciferae. It includes several annual and perennial
herbs, as well as subshrubs, shrubs, and even small trees. Farsetia somalensis
(Pax) Gilg & C. Benedict (Somalia, Kenya, and Ethiopia) is a large shrub or
small tree with hard wood and glossy, gray to red-brown bark, while F. un-
dulicarpa Jonsell (Kenya and Tanzania) is a shrub to 2 m high (Jonsell, 1986).
The majority of the Alysseae have rather small seeds dispersed either by
strong winds in open habitats or by rain wash. Wind dispersal is common in
many genera with broadly winged seeds (e.g., Farsetia, Fibigia Medicus, Lu-
naria). It is restricted, however, to a few genera with samaroid (Neotchihat-
chewia Rauschert, Peltaria) or inflated (Physoptychis Boiss.) indehiscent fruits.
Seeds that produce abundant mucilage when wet may be dispersed by adhering
188 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
to animals. The fruits of two species of Clypeola L. (C. lappacea Boiss. and C.
aspera (Grauer) Turrill) and of the monotypic Asperuginoides Rauschert (for-
merly Buchingera Boiss. & Hohen.; see Rauschert) are covered with glochidiate
trichomes or deflexed barbellate spines and are dispersed by clinging to the fur
of mammals.
Genera of the Alysseae in the southeastern United States are either noxious
weeds or have members with weedy tendencies. Except for a few species of
Camelina Crantz that are cultivated for their seed oils in parts of the Soviet
Union and Europe, the tribe has no food value. Several species of A/yssum,
Aurinia (L.) Desv. (golden-tuft alyssum), Draba, and Lunaria (honesty or mon-
ey plant) are ornamentals cultivated on a limited scale. On the other hand,
Lobularia maritima (L.) Desv. (sweet alyssum) is probably the most widely
cultivated ornamental of the family Cruciferae.
REFERENCES:
Under family Seer in AL-SHEHBAZ (Jour. Arnold Arb. 65: 343-373. 1984), see
BENTHAM & HOOKER; BuscH; DE CANDOLLE (1821, 1824); Von HAYEK; HEDGE; HEDGE
RECHINGER; ane eee JONSELL (1982); KNIGHTS & BERRIE; MANTON;
Ro uins (1981); ScHULz; and SMAL
AL-SHEHBAZ, I. A. The tribes of Cruciferae A eae in the southeastern United
States. Jour. Arnold Arb. 65: 343-373. 1984.
BaiLey, L. H. Manual of cultivated plants. eae pp. New York. 1949. [Alyssum, Draba,
Lobularia, Lunaria.]
BOLKHOVSKIKH, Z., V. GRIF, T. pane oie ben ZAKHARYEVA. Chromosome numbers
of flowering plants. A. A. Feporov, ed. (Russian and English prefaces.) 926 pp.
Leningrad. 1969. Alsou Berteroa, cao Draba, Lesquerella, Lobularia, Lu-
naria, 162-175.]
Burtt, B. L. The genus Ricotia. Kew Bull. 6: 123-132. pls. J, 2. 1952.
CHAYTOR, D. A., & W. B. TurRILL. The genus C/ypeola and its ae variation.
Bull. Misc. Inf. Kew 1935: 1-24. 1936.
CONTANDRIOPOULOS, J. Contribution a l’étude cytotaxinomique des Alysseae Adams
d ce. Bull. Soc. Bot. Suisse 79: 313-334. 1970. [Al/yssoides, Alyssum, Aurinia,
Berteroa, Bornmuellera, Fibigia.]
Davis, P. H., ed. Cruciferae. Fl. Turkey 1: 248-495. 1965. [Lunarieae, Alysseae, Dra-
2
Dub _ey, T. R., & J. CULLEN. Studies in the Old World Alysseae Hayek. Feddes Repert.
71: 218-228. 1965. [Tribal limits, key to eee ng species, evaluation of Ptilo-
trichum, Hormathophylla, gen. nov., key to spec
Duncan, W. H., . T. Kartesz. Vascular flora of Cone An annotated checklist.
ix + 143 pp. Athens, Georgia. 1981. [Camelina, Draba, Lunaria.
GATTINGER, A. The flora of Tennessee and a philosophy of botany. 296 pp. Nashville.
1901. [Berteroa, Camelina, Draba, Lesquerella, Lobularia
GoLpBLATT, P., ed. Index to plant chromosome numbers 1975-1978, Monogr. Syst.
ot. 5. vii + 553 pp. 1981. [4/vssum, Berteroa, Draba, Lesquerella, Lobularia,
Lunaria, 151- hee
———., ed. Index to plant chromosome numbers 1979-1981. bid. 8. viii + 427 p
1984, alas, en Camelina, Draba, Lesquerella, Lobularia, 115-123.]
Index to plant chromosome numbers 1982-1983. /bid. 13. vii + 224 pp.
98 iim, Berteroa, Camelina, Draba, Lesquerella, 63-66.
GREUTER, W. Some notes on Bornmuellera in ieeee and an interspecific hybrid in
the Nips (Cruciferae) Candollea 30: 13-20. 1975.
1987] AL-SHEHBAZ, ALYSSEAE 189
Hepce, I. C. Elburzia: a new genus of the Cruciferae from Iran. Notes Bot. Gard.
Edinburgh 29: 181-184. 1969. [Distributions and comparative morphology of the
monotypic Elburzia, Petrocallis, and Pseudovesicaria.]
JoNsELL, B. A monograph of Farsetia (Cruciferae). Symb. Bot. Upsal. 25(3): 1-107 +
ae als 1986.
Kumar, P. R., & S. TsuNopA. Variation in oil content and ae acid composition
among oa from the Cruciferae. Pp. 235-252 in S. TsuNopA, K. HINATA, & C.
Gomez-Campo, eds., Brassica crops and wild allies. Aen 1980. Dipti Ber-
teroa, Camelina, Lesquerella, Lobularia, Lunaria.]
Kuprer, P. Recherches sur les liens de parenté entre la flore orophile des Alpes et celle
des Pyrénées. (English summary.) Boissiera 23: 1-322. pls. 1-10. 1974. [Alyssum,
suena sk aaa Alyssoides, Aurinia, Lobularia, 192-228, pl. 10.]
LITCHFIELD, C. e a, B-distribution of oleic, linoleic, and linolenic acids in Cruciferae
seed ence Jour. Am. Oil Chem. Soc. 48: 467-472. 1971. [Alyssum, Ca-
melina, Lobularia, Lunaria.]
MacRoserts, D. T. The vascular plants of Louisiana. An annotated checklist and bib-
liography of the vascular plants reported to grow without cultivation in Louisiana.
ull. Mus. Life Sci. Louisiana State Univ. Shreveport 6: 1-165. 1984. [A/yssum,
Camelina, Draba, Lobularia.]
Moors, R. J., ed. Index to plant chromosome numbers 1967-1971. Regnum Veg. 90:
1-539. 1973, [Alyssum, Berteroa, Camelina, Draba, Lesquerella, Lobularia, Lunar-
ia, 201-210.]
Pou ter, B. A. The genus Grael/sia. Notes Bot. Gard. Edinburgh 22: 85-93. p/. 4. 1956.
[Six species, occurrence of latiseptate and angustiseptate fruits within the genus, key,
ma
p.
Princen, L. H., & J. A. RotHFus. Development of new crops f d materials.
Jour. Am. Oil Chem. Soc. 61: 281-289. 1984. ee Lunaria, 285, 286.]
RAUSCHERT, S. Nomina nova generica et combinationes novae Spermatophytorum et
Pteridophytorum. Taxon 31: 554-563. 1982. [Asperuginoides replaces Buchingera
and Neotchihatchewia replaces Tchihatchewia, 558.]
Reeves, R. D., R. R. Brooks, & T. R. DupLey. Uptake of nickel by species of A/yssum,
Bornmuellera, and other related genera of Old World tribus Alysseae. Taxon 32:
184-192. 1983. [Survey of 15 genera.]
SMALL, J. K. Flora of the southeastern United States. xii + 1370 pp. New York. 1903.
[Draba, Camelina, Lesquerella, Lobularia (as Koniga).]
Key TO THE GENERA OF ALYSSEAE IN THE SOUTHEASTERN UNITED STATES*
A. Fruits more than | cm wide, gynophores 1-5 cm long, funicles completely adnate to
the septum; trichomes simple or lacking. .....................24.. 20. Lunaria.
A. Fruits less than 1 cm wide, gynophores absent or to 3 mm long, sae ni from
the septum or adnate only at base; trichomes branched 1 simple
re)
nes
B. All trichomes uniformly bifurcate, medifixed, sessile, appressed. .............
speaisac geting atte gach In airs aie ea a a go Sauces ease dy aa ce ne deeaieys 22. Lobularia.
. Trichomes furcate, branched, or stellate, a mixed with es ones,
stalked or sessile, usually appressed when s
C. Fruits inflated, not compressed, era or eibos to subdidymous.
ee)
‘The genera are numbered as in the treatment of the tribes of the Cruciferae in the southeastern
United States (Jour. Arnold Arb. 65: 343-373. 1984). Genera | and 2 (Thelypodieae) appeared in
ibid. 66: 95-111. 1985; genera 3-13 (Brassiceae) in ibid. 279-351; and genera 14-19 (Lepidieae) in
ibid. 67: 265-311. 1986.
190 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
D. Fruits pyriform, keeled at the replum, the valves acuminate, ending abruptly
in a Stylelike beak, the septum nerveless; seeds usually oblong, cotyledons
Incumbent. ....0....00.0.00 0.0 eee eee eee.. 26. Camelina.
D. Fruits globose to subdidymous, not keeled, the valves rounded or obtuse
at apex, the septum (when present) with a midnerve extending from its
center to the base of style; seeds nearly orbicular, — accumbent.
Diner teat A oe dantena koa aoe Grad ae ea enc aae ae Lesquerella.
C. Fruits not peraee vee parallel to the septum, peer to oblong or
lanceolate to lin
E. Seeds | per locule. borne on an apical placenta, copiously ee
Wiel Wele outdo oti ben Sessa eicewen de hate eet aue! 21. Alyssum.
E. Seeds 2 to many per locule, borne on 2 parietal placentae, not or only
slightly mucilaginous when wet.
F. Cauline leaves Gon auriculate; fruits with bulbous-based tri-
chomes, septum with a midnerve extending from its center to the base
Ol StVlOo ees rai s ae A ae tae een 25. Lesquerella.
F. Cauline leaves rained not auriculate; fruits without bulbous-based
trichomes, septum nerveless
G. Petals deeply 2- lobed: filaments of lateral stamens a
seeds winged or margined. .................... 23. Berteroa.
G. Petals entire or sometimes emarginate, if 2-lobed (Draba oo
then plants scapose; filaments unappendaged; seeds neither winged
nor margined. .......0... 0.00.0 c cece eee eee 4. Draba.
20. Lunaria Linnaeus, Sp. Pl. 2: 653. 1753; Gen. Pl. ed. 5. 294. 1754.
Annual(?), biennial uy perennial] herbs with simple trichomes. Stems erect,
branching above. Basal and lower cauline leaves opposite or rarely alternate,
long petiolate, large, ovate-cordate, undivided, coarsely dentate [or spinulose-
dentate]; upper leaves alternate, sessile or subsessile [or distinctly petiolate].
Inflorescences corymbose racemes or panicles, greatly elongated in fruit; low-
ermost branches bracteate; flowers ebracteate, large, showy. Sepals erect, cu-
cullate; outer pair linear, not saccate at base; inner pair broadly oblong-elliptic,
strongly saccate. Petals violet or purple, rarely lavender or white, obovate, long
clawed, usually twice as long as the sepals or longer. Lateral nectar glands large,
annular, 2-lobed on the outer side, 3-lobed on the inner [sometimes divided
pairs) or strongly curved (lateral pair); anthers large, linear or oblong, obtuse.
Ovary stipitate, 4- to 8-ovulate, glabrous or ciliate; style filiform; stigma 2-lobed,
the lobes decurrent [or not], opposite the replum. Fruits dehiscent, very large
(2-9 x 1-3.5 cm), strongly flattened parallel to the septum, usually pendulous,
op/one to suborbicular [or lanceolate-elliptic], obtuse [or acute] at both ends;
es glabrous, flat, papery, finely or obscurely net veined, without a midnerve;
styles long [or short usually flattened near the base; replum ciliate or glabrous:
septum persistent, shining, membranaceous, nerveless, very broad, with nar-
rowly linear epidermal cells perpendicular to the long axis of fruit; funicles
long, almost completely adnate to the septum; gynophores slender, 1-5 cm
long [rarely obsolete or to | mm]. Seeds few, large, biseriately arranged in each
locule, reniform or rarely suborbicular, flattened, slightly biconvex, minutely
1987] AL-SHEHBAZ, ALYSSEAE 191
reticulate, brown, uniformly broad winged all around except at the wingless
area of hilum, nonmucilaginous when wet; cotyledons accumbent, large. Base
chromosome number 15. Lectotype species: L. annua L.; see Britton & Brown,
Illus. Fl. No. U. S. & Canada, ed. 2. 2: 190. 1913; see also Green and Maire
for a later lectotypification (based on L. rediviva L.) that contradicts article 8
of ICBN. (Name from Latin, /una, moon, which the large, persistent, silvery
septum of the fruit superficially resembles.)— HONESTY, MONEY PLANT, SATIN
FLOWER, MOONWORT.
A genus of three species native to southern, central, and eastern Europe. Two
species are grown as ornamentals, and these sometimes escape from cultivation.
The third, Lunaria Telekiana Jav., is a narrow endemic of northeastern Al-
bania. It differs from the other species in having very short (to ca. 1 mm)
gynophores, densely ciliate valve margins, and lateral sepals with longer (to ca.
2.5 mm) saccate bases. Both L. annua (L. biennis Moench, L. inodora Lam.),
honesty, bolbonac, silver-dollar, penny flower, money plant, 2m = 30, and L.
rediviva (L. odorata Lam., L. alpina Berg.), money plant, 2n = 30, are grown
in North America. Lunaria annua has been reported as an escape from cul-
tivation, but apparently not a naturalized one, in many states (including Ar-
kansas and Georgia). It is easily distinguished by its biennial habit, oblong to
suborbicular fruits with both apex and base obtuse, and subsessile or sessile
upper cauline leaves. In the perennial L. rediviva the upper cauline leaves are
petiolate and the fruits are usually elliptic-lanceolate with both apex and base
acute. Of the two subspecies of L. annua, only the biennial subsp. annua is
present in our area. Subspecies pachyrrhiza (Borbas) Hayek, a perennial with
fusiform tubers, is distributed in Romania, the Balkan peninsula, and southern
Italy.
Lunaria is most closely related to the eastern Mediterranean Ricotia (nine
species), from which it differs in its coarser habit (stems to 16 dm high),
undivided leaves, and stipitate fruits (1—-)1.5-3.5 cm wide with a well-developed
septum. Species of Ricotia are smaller plants to 4 dm high having pinnate or
trifoliolate (very rarely undivided) leaves and sessile fruits 0.5-1(-1.5) cm wide
with a very delicate septum that is sometimes lacking. Both genera were main-
tained in the Alysseae by De Candolle (1821, 1824), Bentham & Hooker, Von
Hayek, and Janchen, but the last two placed them in subtribe Lunariinae Hayek.
In Schulz’s classification Lunaria and Ricotia, along with six other genera, are
placed in the tribe Lunarieae, which was distinguished from the Alysseae only
by the presence of simple instead of branched trichomes. Both types of tri-
chome, however, are found in several genera of the Cruciferae, notably Arabis
L., Draba, and Sisymbrium L. Dvorak (1971) suggested that the Lunarieae
sensu Schulz, particularly Lunaria, represent an evolutionary line derived from
an ancestor not very different from meee se ea Scat Pe-
trop. of the Thelypodieae Prantl. The Lunarieae a
in which some genera (e.g., Se/enia Nutt. and inten W. J. Hooker)
are clearly unrelated to Lunaria. Von Hayek’s derivation of Lunaria from
Ricotia needs careful evaluation, but it is evident that the two are more closely
related to each other than to other genera of the Cruciferae.
192 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
The erect sepals, long claws of the petals, and flattened bases of the median
staminal filaments of Lunaria form a long tube that makes the abundant nectar
usually accessible to insects with proboscises longer than 1 cm. The butterflies
Vanessa (Nymphalidae) and Pieris (Pieridae), the bees Bombus (Bombidae)
and Andrena (Andrenidae), and the honeybee Apis mellifera (Apidae) are among
the most common visitors of Lunaria flowers (Knuth). Self-pollination can be
brought about effectively by small pollen-collecting insects because of the close
proximity of the stigma to the median anthers. Insects with short proboscises
can reach the nectar by poking holes through the base of the calyx.
Most chromosome counts for Lunaria annua and L. rediviva indicate 2n =
30, but Dvorak & Dadakova and Polatschek reported 2n = 28 for these species.
The last author suggested that Lunaria is based on x = 7, while Dvorak (1971)
speculated that the genus evolved through allopolyploidy from unknown ances-
tors with x = 7 and 8. The karyotype of Lunaria consists of small chromosomes,
of which two (at least in L. rediviva) are believed to be B chromosomes (Manton).
Failure to observe this pair may have led to deviant counts. Diploid and
tetraploid counts based on x = 15 have been found in L. rediviva (Jankun).
Lunaria is unusual in the Cruciferae for its high concentrations of unique
or very rare secondary compounds. It 1s rich in alkaloids, of which some are
known only in this genus and at least six (lunarine, lunaridine, lunariamine,
numismine, tetrahydrolunarine, and tetrahydrolunaridine) have been charac-
terized. Isopropyl, 2-butyl, and 5-methylthiopenty!l glucosinolates have been
found in L. annua, and the last compound occurs in L. rediviva (Kjaer). The
green parts of plants of the former species also contain 3-methylthiopropyl-
glucosinolate (Cole). It has been suggested that the high concentrations of
alkaloids in Lunaria may have evolved as an escape from crucifer-adapted
pathogens or herbivores. The seed extract of Lunaria is the first reported source
of m-carboxy-substituted aromatic amino acids among higher plants (Olesen
Larsen). The unhydrolyzed seed extract of L. annua contains four amino acids
and y-glutamyl derivatives not discovered previously in nature.
Lunaria annua is an excellent source of long-chain monounsaturated acids,
which constitute 90 percent of the total fatty-acid content. The seed oil is a
potential source of erucic acid (42 percent) and contains 21 to 25 percent
nervonic acid. The content of the latter acid is the highest reported for any
seed oil (Wilson et a/., Mukherjee & Kiewitt).
Because the funicles are adnate to the septum, the seeds of Lunaria usually
remain attached to the septum after the valves fall off. They are eventually
detached as a result of the vibration of the septum and may glide away from
the plant because of the presence of a broad wing. However, they sometimes
adhere to the valves and can be carried away with them.
Lunaria annua has an absolute requirement of cold treatment (vernalization)
for flowering. Stem elongation in rosette plants can be induced by the appli-
cation of the gibberellic acids GA3 and GA7. However, the gibberellin treat-
ment fails to induce flowering in nonvernalized plants (Zeevaart). Likewise,
sprouts developed on callus or on petioles grown in sterile cultures do not
flower unless vernalized (Pierik, 1967). Annual plants of L. annua, which is
otherwise a biennial, have been obtained recently (Wellensiek, 1973).
—_—
1987] AL-SHEHBAZ, ALYSSEAE 193
Both Lunaria annua and L. rediviva are grown for their attractive flowers
and particularly for their infructescences, which are used in dry bouquets after
the removal of valves and seeds. Crisp stated that the seeds are occasionally
used as condiments and the roots are eaten as a salad or cooked as a vegetable.
The seeds of L. annua contain high levels of long-chain fatty acids, but the
species has not been used as a source of industrial oils. Although both species
may escape from cultivation, neither is a successful weed in the New World.
REFERENCES:
Under family references in AL-SHEHBAz (Jour. Arnold Arb. 65: 343-373. 1984), see
BATEMAN (1955a); BENTHAM & HOOKER; BERGGREN; BOUMAN; BRITTON & BROWN; BUSCH;
De CANDOLLE (1821, 1824); CoLE (1976); Crisp; ae et al.; EIGNER; FERNALD;
Von Hayek; JANCHEN; KJAER (1960); KNIGHTS & BERRIE; KNUTH; LA Porte; MAIRE;
MANTON; MARKGRAF; MEDVE; MURLEY; PONZI; rages Ro .uins (1981); ScHuLz; E. B.
SMITH; and VAUGHAN & WHITEHOUSE.
Under tribal references see BAILEY; BOLKHOVSKIKH et a/l.; DUNCAN & KARTESZ;
GOLDBLATT (1981); KUMAR & TsUNODA; LITCHFIELD; Moore; and PRINCEN & ROTHFUS.
Asupy, J. Ww. & A. D. THomson. New plant disease record in New Zealand: turnip
BALL, P. W. Lunaria. es T. TuTin et al., eds., Fl. Europaea 1: 295, 296. 1964.
BLapDon, P., R. IKAN, F. S. SprRING, & A. D. Tait. The chemistry of Lunaria alkaloids.
I. Tetrahedron Lett. 1959(9): ae 23. 1959.
Bort, H.-G. Uber die Alkaloide von Lunaria biennis. Chem. Ber. 87: 1082, 1083. 1954.
Cimin1, M. Sopra un caso di fillomania nella Lunaria annua L. Bull. Soc. Bot. Ital.
1921: 58-61. 1921. [Teratology.]
CRANFILL, R., & J. W. THIERET. Thirty additions to the vascular flora of Kentucky. Sida
: 55- 58. 1981. [L. annua, 57.
Dosxotcu, R. W., E. H. FAIRCHILD, & W. KuBeLKA. A revision of the structures of the
Lunaria alkaloids LBX and LBZ. Experientia 28: 382, 383. 1972.
DvorAk, F. Pyispévek ke studiu variability Lunaria rediviva a Biologia (Bratislava)
22: 451-457. 1967. [A related paper in ibid. 23: 549-553.
—. On the evolutionary relationship in the family ae Feddes Repert. 82:
357-372. 1971. [L. annua, L. rediviva; relationship to Macropodium.|
ADAKOVA. Chromosome counts and chromosome morphology of some
selected species. Folia Geobot. Phytotax. 19: 41-70. 1984. [L. annua, 58, 59, 2n
8.
2
Green, M. L. Pp. 111-195 in A. S. Hircucock & M. L. GREEN, Standard-species of
Linnean genera of Phanerogamae (1753-1754). Internatl. Bot. Congr. Cambridge
(England), Nomenclature. Proposals by British botanists. 1929. [L. rediviva as the
lectotype species, 171.
HAGEMANN, P. Histochemical patterns in pith lignification in the fruit stalk of Lunaria
annua L. (In German; English summary.) Beitr. Biol. Pflanz. 51: 81-97. 1976a
. Pith lignification in the pedicel of Lunaria annua (Cruciferae); example of a
histochemical investigation. Mikrokosmos 65(3): 86-91. 1976b.*
Hansen, O. R. Lunarine, an alkaloid from Lunaria biennis. Acta Chem. Scand. 1: 656-
658. 1947. [Isolation and purification. ]
HarrRIMAN, N. A. Jn: A. Love, ed., IOPB chromosome number reports LX. Taxon 27:
223-231. 1978. [L. annua, 228, 2n = 30.]
Huneck, S. Uber die Alkaloide von Lunaria rediviva L. Naturwissenschaften 49: 233.
62
1962.
Husson, H.-P., C. Poupat, B. Ropricuez, & P. Potier. Alkaloids of Lunaria biennis
194 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
(Cruciferae): oe of (+)-tetrahydrolunaridine. (In French.) Tetrahedron Lett.
1971: 2697-2 971.
JANKUN, A. /n: ar SKALINSKA & E. PoGAn, Further studies in chromosome numbers
of Polish angiosperms, ninth contribution. Acta Biol. Cracov. Bot. 14: 199-213. pl.
50, 1971, rediviva, 201, 202, 2m = 30, 60.
JANOoT, M.-M., & J. LEMEN. Sur les alcaloides de Lunaria biennis Mnch. (Cruciféres).
Bull. Soc. Chim. France 56: 1840-1842. 1956.
JAvorka, 8, Lunaria Telekiana Jav. n. sp. Magyar Bot. Lapok 19: 1, 2.
Lanpl, M. Osservazioni e ricerche sulla Lunaria pachyrrhiza Borbas. rere re Flori
9: 104-117. 1933.
Miwa, T. K. Gas chromatograms of synthetic liquid waxes prepared from seed triglyc-
erides of Limnanthes, Crambe and Lunaria. Jour. Am. Oil Chem. Soc. 49: 673,
674. 1972,
& I. A. Worf. Fatty acids, fatty alcohols, wax esters, and methyl esters from
Crambe abyssinica and Lunaria annua seed oils. Jour. Am. Oil Chem. Soc. 40:
3
MUKHERJEE, K. D., & I. Kiewitr. Lipids containing very long chain monounsaturated
acyl moieties in seeds of Lumaria annua. Phytochemistry 25: 401-404. 1986. [Ma-
ture gan contain 24 percent oleic, 42.7 percent erucic, and 24.7 percent nervonic
fatty acids.]
OLESEN ee P. Amino acids and y-glutamyl derivatives in seeds of Lunaria annua
L. Part III. Acta Chem. Scand. 21: 1592-1604. 1967. [Part I in ‘bid. 16: 1511-1518.
1962; part II in ‘bid. 19: 1071-1078. 1967.]
Prerik, R. L. M. Regulation of morphogenesis by growth regulators and temperature
treatment in isolated tissues of Lunaria annua L. Proc. Nederl. Akad. Wet. C. 68:
324-332. 1965.
he induction and initiation of flowerbuds in vitro in tissues of Lunaria annua
L. Naturwissenschaften 53: 45. 1966. [Petioles 3 cm long produced flowers at 26°C
sien placed in culture and later vernalized for 16 weeks.]
—. Regeneration, vernalization and flowering in Lunaria annua L. in vivo and in
vitro. Meded. Landb. Wageningen 67(6): 1-71. figs. 1-15. 1967.
ventitious root formation in oe ae segments of Lunaria annua L.
Zeitschr. Pflanzenphysiol. 66: 343-351.
POLATSCHEK, A. Cytotaxonomische Beitrage zur Flora der Ostalpenlinder, I. Osterr.
Bot. Zeitschr. 113: 1-46. 1966. [L. rediviva, 24, 2m = 28+
Portier, P., & J. LEMEN. Alcaloide du Lunaria ne Moench (Cruciféres). Bull. Soc.
Chim. ees 1959: 456-459. nie ae oe cay in ibid. 201, 202.]
PoupatT, C., H.-P. Husson, B. RopriGugz, A. ON, P. Potier, & M.-M. JANOT.
Recent pies of the alkaloids ae ae er Moench, Cruciferae—I. Lu-
narine and derivatives: structure of four secondary alkaloids. (In French; English
summary.) Tetrahedron 28: 3087-3101. 1972. [Part II in ibid. 3103-3111.]
RopriGuez, H.-P. Husson, P. Potier, & M.-M. JANoT. New alkaloids iso-
lated from the seeds of the dollar-plant Lunaria biennis Moench (Cruciferae). (In
French.) Compt. Rend. Acad. Sci. Paris C. 269: 335-338. 1969. [A related paper
on the biogenesis of lunarine in ibid. 273: 433-436. 1971.]
PRINCcEN, L. H. New oilseed crops on the horizon. Econ. Bot. 37: 478-492. 1983.
ee 484]
Rees, E. Lunaria annua a $ active principle. res Abstr. 5: 3123. 1911. [Discovery
gait alkaloid eae jehal to frogs and rabbits.
Ropionova, G. B. On the embryogenesis of Eumanis annua L. (In Russian.) Bot. Zhur.
51: 1506-1511. 1966.
UnrikovA, A. In: J. MAsovsxy et al., Index of chromosome numbers of Slovakian flora.
Acta Fac. Nat. Comen. Bot. 23: 1-23. 1974. [L. rediviva, 13, 2n = 30; 2n = 30
reported for L. annua in ibid. 25: 9. 1976.]
1987] AL-SHEHBAZ, ALYSSEAE 195
Uxricn, R. Observations biométriques sur la croissance des fruits de lunaire (Lunaria
biennis Moench). Bull. Soc. Bot. France 84: 645-654. 1938.
Usner, B. F., & P. FEENY. Atypical secondary compounds in the family Cruciferae:
tests for toxicity to Pieris rapae, and adapted crucifer- ee insect. Entomol. Exp
Appl. 34: 257-265. 1985. [Toxicity of lunarine, L. annua.]
WELLENSIEK, S. J. en and age in Lunaria Fetes Proc. Nederl. Akad. Wet.
. 61: 561-571. 8.
. Genetics me flower formation of annual Lunaria. Nether]. Jour. Agr. Sci. 21:
163-166. 1973.
. K. Hicazy. The juvenile see for flowering in Lunaria biennis. Proc.
Nederl. Akad. Wet. C. 64: 458-463.
WILLAMAN, J. J., & H.-L. Li. Alkaloid- eee plants and their contained alkaloids,
1957-1968. Lloydia 33(3A, supplement): i-vil, 1-286. 1970. [Lunaria, 86
. SCHUBERT. Alkaloid-bearing plants and their contained alkaloids. U. S.
Dep. Agr. Tech. Bull. 1234. 287 pp. 1961. [Lunaria, 79.]
Witson, T. L., C. R. Smirn, Jr., & I. A. WoLrr. Lunaria seed oil—a rich source of C,,
fatty acids. Jour. Am. Oil Chem. Soc. 39: 104, 105. 1962.
oe J. A.D. Vernalization and gibberellins in Lunaria annua L. Pp. 1357-1370
n F. WIGHTMAN & G. SETTERFIELD, eds., Biochemistry and physiology of plant
iicer: en Ottawa. 1968.
21. Alyssum Linnaeus, Sp. Pl. 2: 650. 1753; Gen. Pl. ed. 5. 293. 1754.
Annual [biennial or perennial] herbs [rarely subshrubs]. Stems erect to de-
cumbent, usually branched at base [sometimes with sterile shoots and winter
rosettes]. Indumentum of appressed, stellate trichomes with few [or many]
branched [or unbranched] rays [or sometimes of lepidote trichomes]; simple
or furcate trichomes present [or absent]. Leaves undivided, entire, attenuate,
neither swollen nor persistent at base. Inflorescence an ebracteate, corymbose
raceme [or panicle], elongated [or not] in fruit; fruiting pedicels divaricate
[ascending, or reflexed]. Sepals equal [or unequal], free [or sometimes appearing
connate because of interlocking trichomes at adjacent margins of sepals], per-
sistent [or caducous], [inflated] or not, equal, not saccate at base, pubescent on
outside, glabrous [or pubescent] on inside. Petals yellow [white, or rarely pink
or lavender], obovate [or spatulate], emarginate [or entire], gradually [or abrupt-
ly] narrowed into claws, glabrous or sparsely [to densely] pubescent on outside;
claws without [or rarely with] a basal appendage. Nectar glands 4, 1 on each
side of the lateral stamens, filiform [globose, or triangular], median glands
always absent. Stamens 6, somewhat tetradynamous; filaments wingless [or
unilaterally or bilaterally winged], toothless and unappendaged [or variously
toothed and/or appendaged], free [or rarely connate]; anthers small, introrse,
acute or obtuse at apex. Ovary sessile, 2 [1 or 4-8]-ovulate; placentation api-
cal [or rarely parietal]; stigmas capitate. Fruits dehiscent [rarely indehiscent],
orbicular [oblong, elliptic, ovate, obovate, or obcordate], almost always flat-
tened parallel to the septum, inflated in the middle [or throughout, or not
inflated], emarginate or truncate [acute, or retuse] at apex, entire [rarely cren-
ulate or undulate] at margin, pubescent [or glabrous]; valves nerveless; styles
persistent, pubescent [or glabrous]. Seeds compressed, narrowly [to broadly]
winged [or wingless], mucilaginous [or not] when wet; cotyledons accumbent
[or incumbent]. Base chromosome number 8. (Including Gamosepalum Hausskn.
196 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
non Schlechter, Meniocus Desv., Moenchia Roth, Odontarrhena C. A. Meyer,
Psilonema C. A. Meyer, Ptilotrichum C. A. Meyer, Triplopetalum E. J. Ny-
arady.) LECTOTYPE SPECIES: A. montanum L.; see Britton & Brown, Illus. Fl.
No. U. S. & Canada, ed. 2. 2: 154. 1913. (Name from Greek, a, not or privative,
and /yssa, rabies or madness; the name was used for plants reputed in ancient
times as a remedy for hydrophobia, as a cure for madness, and as a calmative
for anger.)— MADworrt.
A well-defined, taxonomically difficult genus of at least 170 (probably to
190) species primarily centered in Turkey (90 species, 50 endemic), with a rich
representation in the Balkan peninsula (45 species, 20 endemic) and in the
Caucasus and adjacent parts of the Middle East (63 species, 25 endemic). The
genus 1s poorly developed in central and eastern Asia (seven species endemic)
and in North Africa and the Iberian peninsula (eight endemic). With the ex-
ception of A/yssum americanum Greene (Alaska and Yukon Territory, Can-
ada), which may be conspecific with the Siberian 4. obovatum (C. A. Meyer)
Turcz. (Dudley, 1964b), the genus is almost exclusively Eurasian and is mostly
confined south of the 50th parallel. The great majority of taxa are narrowly
endemic, and only about ten species are widely distributed weeds. A/yssum is
represented in North America by one indigenous and six naturalized species,
and in the southeastern United States by one weedy species.
Alyssum 1s divided into at least six or seven well-marked sections previously
recognized as distinct genera. Section PstLONEMA (C. A. Meyer) J. D. Hooker
(Psilonema, Alyssum subg. Tetratrichia Gay) (annuals; filaments slender, eden-
tate, unappendaged, wingless; fruits dehiscent, valves equally inflated; seeds 2
per locule, winged or wingless, mucilaginous when wet), containing five species
indigenous to southwestern Asia and the Mediterranean region, is represented
in our area by a single species. A/yssum Alyssoides (L.) L. (Clypeola Alyssoides
L., C. campestris L., A. calycinum L., A. campestre (L.) L., Psilonema Alys-
soides (L.) C. A. Meyer), pale alyssum, 2” = 32, a native of northern Africa
and western Europe eastward to India, is naturalized in Canada, the United
States, and Argentina. It grows on disturbed gravelly or sandy banks, waste
grounds, and dry hillsides, in meadows, and along roadsides. It is rare in the
Southeastern States and occurs in Cumberland County, Tennessee (R. Simmers,
pers. comm.), Marion County, Arkansas (Smith), and Lincoln Parish, Louisiana
(Logan). According to MacRoberts, the record from Louisiana needs verifi-
cation.
Of the two varieties recognized by Dudley (1965a) in A/yssum Alyssoides,
only var. A/yssoides 1s naturalized in the New World. The other, var. depressum
(Schur) T. R. Dudley, is endemic to the Balkan peninsula. The species is
distinguished from the other alyssums in North America by its persistent sepals;
the compressed margin and inflated center of its fruits; its filiform, persistent
nectar glands; and its unappendaged, toothless, and wingless staminal filaments.
It may be confused with 4. desertorum Stapf, but this has dentate filaments,
deciduous sepals, and glabrous fruits.
Section ALyssuM (annuals, biennials, or perennials; filaments winged, ap-
pendaged, or toothed; fruits dehiscent; seeds 2 per locule, winged or wingless,
1987] AL-SHEHBAZ, ALYSSEAE 197
mucilaginous when wet), contains more than 70 species and is represented in
North America by the Eurasian A. desertorum, A. minus (L.) Rothm. var.
micranthum (C. A. Meyer) T. R. Dudley, A. strigosum Banks & Solander, and
A, Szowitsianum Fischer & Meyer. These are naturalized in Manitoba and
Alberta southward into the Mountain and Pacific states and Nebraska.
Section ODONTARRHENA (C. A. Meyer) W. D. Koch (perennials; filaments
winged, dentate, or appendaged; fruits dehiscent or indehiscent, 1-seeded; seeds
winged or wingless, rarely mucilaginous when wet) contains more than 70
species, of which only the native A/yssum americanum (= A. obovatum?) and
the European A. murale Waldst. & Kit. grow in North America. The latter is
an occasional escape from cultivation and is known from a few localities in
Colorado, Michigan, and Québec.
The remaining sections of A/yssum (sects. MENIOcUS (Desv.) J. D. Hooker
(seven species), GAMOSEPALUM (Hausskn.) T. R. Dudley (ten species), and
TETRADENIA (Spach) T. R. Dudley (three species)) are not represented in North
America. Krasnoborov has recently proposed the monotypic sect. STEVENIOI-
DES, which resembles sects. PSILONEMA and ODONTARRHENA in Its edentate
staminal filaments and uniovulate locules, respectively.
Both Aurinia saxatilis (L.) Desv. (Alyssum saxatile L.), golden-tuft alyssum,
basket-of-gold, gold-dust, rock madwort, and Au. petraea (P. Ard.) Schur (A.
petraeum P. Ard.) are occasional escapes from cultivation in the United States,
and the former has been reported from Mississippi (Jones). Although Aurinia
Desv. has been treated as a section of A/yssum by numerous authors (e.g.,
Busch, Schulz, Ball & Dudley, Maire, Markgraf), Dudley (1964c) recognized
it as a genus remotely related to A/yssum and most closely allied to Berteroa
DC. or possibly to Alyssoides Miller. Dudley separated Aurinia from Alyssum
mainly on the basis of leaf characters. Aurinia was said to have repand-sinuate
or dentate rosette leaves 2-10 cm long, deeply grooved petioles with swollen
and persistent bases, and cauline leaves about half (or less) the size of the rosette
ones. On the other hand, A/yssum has entire rosette leaves 0.5—2 cm long, flat
petioles neither swollen nor persistent at the base, and cauline leaves subequal
in size to the basal ones. These alleged differences, however, are inconsistent
within each of the two genera. For example, the basal leaves of Au. corymbosa
Griseb. and Au. halimifolia (Boiss.) Cullen & T. R. Dudley are usually entire
and have petioles neither swollen nor persistent, while several species of A/ys-
sum (e.g., A. aizoides Boiss.) have persistent and swollen petiole bases, and
many others (e.g., A. argenteum All. and A. Bertolonii Desv.) have deeply
grooved petioles. The other differences listed by Dudley, particularly the shape
of floral buds and the lobing of immature stigmas, are not sharply defined and
are therefore unreliable. All species of Aurinia have spreading sepals, while all
except a few species of A/yssum (e.g., A. spinosum L.) have erect ones. Aurinia
consists of closely related species that are difficult to separate from Alyssum
on the basis of fruit and floral characters alone. The differences in leaf characters
between these genera may not justify the recognition of Aurinia as an inde-
pendent genus remotely related to A/yssum. The lack of reliable differences
between these genera has led to the reduction of the former to a section of the
latter, as was done by numerous authors including Ball & Dudley.
198 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Alyssum is easily distinguished from other members of the Alysseae by its
nonsaccate sepals, entire or inconspicuously lobed stigmas, appressed stellate
trichomes occasionally mixed with furcate (but never medifixed and bifid) ones,
and usually dehiscent and latiseptate fruits without barbulate trichomes. The
genus 1s often confused with Lobularia, but this always has bifid, medifixed,
appressed trichomes.
Very high rates of selfing have been observed in several species of A/yssum
(Persson). Cleistogamy is often associated with damp weather. Giinthart sug-
gested that the basal wings, teeth, or appendages of staminal filaments guide
the proboscis of a visiting insect to the nectar glands. Dudley (1963) and
Bergdolt, on the other hand, claimed that these staminal structures are not
involved in pollination, and the latter maintained that they are vestiges of
ancestral petaloid structures from which the filaments evolved. It is highly
unlikely, however, that these floral structures, which are present in all except
five species of A/yssum and in several other genera of the Alysseae, do not
participate in pollination and do not have adaptive value. It should be noted
that the distinctions between certain sections of A/yssum and between certain
genera of the Alysseae rely primarily on the presence vs. absence of the staminal
appendages or teeth (Dudley, 1964b; Dudley & Cullen).
Chromosome numbers are known for about 90 species of A/yssum, and all
except a few are based on eight. Species with deviating base numbers (e.g., A.
hirsutum Bieb., 2n = 46) most likely evolved through aneuploidy from ancestors
with x = 8. Section TETRADENIA sensu Dudley (1964b) is the most cytologically
heterogeneous of all sections of A4/yssum. On the basis of chromosome numbers,
morphology, and geographic distributions, Kiipfer has transferred its three
species, A. spinosum (2n = 16, 32), A. cochleatum Cosson & Durand (2n =
22), and A. Lapeyrousianum Jordan (2n = 30), to Hormathophylla. Diploid
and tetraploid counts based on eight are known for 4. obovatum from Siberia
(Goldblatt, 1981, 1984, 1985). Recent counts of 2” = 30 for A. americanum
from Alaska (Dawe & Murray) may support its recognition as a distinct species,
instead of its reduction to a synonym of A. obovatum, as was suggested by
Dudley (1964b). At least 50 species are diploid, 20 are polyploid, and 20 have
both diploid and polyploid populations. Polyploidy played an important role
in the evolution of A/yssum, as is evidenced by its occurrence in about 45
percent of the species for which counts are known. Dudley (1963), however,
found polyploidy in only two of the 21 species he compiled and suggested that
it was insignificant in the evolution of the genus.
Persson studied the karyotypes of several species of A/yssum and noted that
members of sect. ALyssuM have rod-shaped chromosomes, while A. Al/yssoides
has elliptic ones. He suggested that 4. sicu/um Jordan (2n = 48) is an inter-
sectional allopolyploid hybrid, the parental species of which are A. A/yssoides
and A. minus (2n = 16). Interspecific hybridization is apparently very rare in
the genus.
Little is known about the chemistry of 4/yssum; only eight species have been
surveyed for fatty-acid composition, and eight others for glucosinolates. The
limited data indicate that linolenic acid is the primary seed-oil constituent (39-
66 percent), that oleic and linoleic acids are secondary (9-24 percent each),
1987] AL-SHEHBAZ, ALYSSEAE 199
and that erucic acid is lacking (Kumar & Tsunoda). Methionine-derived glu-
cosinolates, particularly 5-methylthiopentyl, 5-methylsulfinylpentyl, 3-meth-
ylsulfinylpropyl, and 3-butenyl glucosinolates, are the dominant compounds
(Hasapis et a/., Kjaer). The distribution of seed glucosinolates and fatty acids
does not support the maintenance of Aurinia as a genus distinct from A/yssum,
but that of seed sterols apparently does (Knights & Berrie).
Vaughan & Whitehouse indicated that seed-coat anatomy supports the sec-
tional classification of A/yssum. They found that in Aurinia (treated as a section)
the epidermal cells have no central columns, the subepidermis is present, and
the palisade cells have thickened radial and inner tangential walls. In A/yssum
the epidermis contains large and hollow central columns, the subepidermis is
lacking, and the palisade cells either have only the inner tangential walls thick-
ened or have all walls evenly thickened. However, they surveyed only five
percent of the species of A/yssum, and it is not known whether their obser-
vations hold for the rest of the genus. According to Metcalfe & Chalk, the
stems of A. spinosum are composed of alternating concentric rings of small,
unlignified, spirally thickened vessels and large, lignified ones with horizontal
bordered pits. It appears that the vascular cambium periodically produces the
“juvenile” form of xylem.
Most species of A/yssum have dehiscent fruits with small, usually mucilag-
inous, and often winged seeds. The seeds are dispersed either by wind or (when
wet) by adhering to animals and equipment. In sect. ODONTARRHENA subsect.
Samarifera T. R. Dudley (nine species; Turkey, northern Syria, and Lesbos
Island, Greece) the fruits are modified into indehiscent, thin-walled, one-seeded
samaras borne on slender, brittle, usually deflexed pedicels and are therefore
dispersed by wind. The evolution of this type of dispersal was accompanied
by an increase of fruit size.
Several species of Al/yssum (e.g., A. Szowitsianum) have conical infructes-
censes, the lowermost pedicels of which are two to three times longer than the
upper ones. The pedicels are closely appressed to the rachis, but soon after
their exposure to rain, they spread horizontally, displaying the concave valves
upward. The impact of raindrops eventually leads to the detachment of the
valves and the release of mucilaginous seeds. The anatomical basis for this
hygrochastic movement of the fruiting pedicels was studied by Zohary & Fahn.
They showed that the adaxial side of the swollen bases of the pedicels consists
of thick-walled fibers with transversely arranged pores, while the abaxial side
has thin-walled fibers with diagonally arranged pores. Due to water absorption
by the thick-walled fibers, the bases of the pedicels swell further and conse-
quently spread in a purely mechanical way.
Species of A/yssum occupy diverse habitats, but the majority are distributed
especially chalks, and rarely on gypsum. Most species of sect. ODONTARRHENA
are endemic to serpentine and other ultrabasic substrates, and at least 46 (66
percent) are hyperaccumulators of nickel. Nickel levels in these species are
often higher than 1000 ug/g of dry weight. The physiology of tolerance and
200 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
hyperaccumulation of nickel is directly related to the presence of high levels
of malic and malonic acids. Nickel is accumulated in the cell vacuoles, and its
presence in the mitochondria is believed to block the citric-acid cycle by deac-
tivating malic acid dehydrogenase. This deactivation leads to the buildup of
malic acid in the vacuoles, enabling them to absorb more nickel (Brooks ef al.,
198la). Seeds of the hyperaccumulators of the A. serpyllifolium Desv. complex
(Iberian peninsula) can germinate on soils with nickel concentrations up to
centrations below 60 ug/g. These physiological differences support the treat-
ment of each of the three subspecies of 4. serpyllifolium as a distinct species.
Except for a few weedy species, the genus has little economic importance.
Alyssum murale, silver alyssum, 1s cultivated as an ornamental in parts of
Europe and North America. The ancients used an infusion prepared from the
flowers and leaves of some species as a sedative for anger and a cure for rabies.
REFERENCES:
Under family references in AL-SHEHBAZ (Jour. Arnold Arb. 65: 343-373. 1984), see
AL-SHEHBAZ & AL-OMAR; BENTHAM & HOOKER; BERGGREN; BUSCH; DE CANDOLLE (1821,
1824); CoLe (1976); FERNALD; GUNTHART (1902); Hasapis et al.; Won HAYEK; JONES;
KJAER (1960); KNIGHTS & BERRIE; MAIRE; MANTON; MARKGRAF; METCALFE & CHALK;
POLATSCHEK; ROLLINS (1981); SCHULZ; E. B. SMitH; and VAUGHAN & WHITEHOUSE.
Under tribal references see BAILEY; BOLKHOVSKIKH et al.; CONTANDRIOPOULOS; DUDLEY
& CULLEN; GOLDBLATT (1981, 1984, 1985); KUMAR & TSUNODA; KUPFER; LITCHFIELD;
MacRoserts; Moore; and REEVES ef al.
Ancev, M. E. Karyological characteristics of Alyssum umbellatum Desv. and Alyssum
hirsutum M. B. (Brassicaceae). [Proc.] 3rd Natl. Conf. Cytogenetics, Bulgaria. Pp.
428-431. 1
& T. Dupe Ley. Jn: A. L6ve, ed., Chromosome number reports LX XIII. Taxon
30: 829-861. 1981. [Counts re i species, 856.]
AVETISIAN, V. Synopsis specierum generis A/yssum L. (Brassicaceae) e Caucaso. Novit.
Syst. Pl. Vasc. 20: 115-120. 1983. [Recognizes 19 species in three sections; a related
paper in ‘bid. 18: 199-204. 1981.]
BaKANOoVA, V. V. Study of biological and morphological characteristics of dwarf sem1-
shrubs of the genus A/yssum under cultivation. (In Ukrainian.) Intr. Aklim. Rosl.
Ukr. Akad. Nauk USSR. 11: 15-20. 1977.
BALL, P. W., & T. R. DuDLEy (with the assistance of E. NyArApy). Al/yssum. In: T. G.
TUTIN ef a eds., Fl. Europaea 1: 297-304. 1964. [Recognized 64 species.]
Baskin, J. M., & C. C. BAskin. Germination and survival in a population of the winter
annual A/yssum eens Canad. Jour. Bot. 52: 2439-2445, 1974a. [Seeds ger-
minate during summer and autumn, but most plants from summer-germinating
seeds are killed by aroun in July and August. ]
—— & ——.. Effect of vernalization on flowering of the winter annual A/yssum
Alyssoides. Bull. Torrey Bot. Club 101: 210-213. 1974b. [Vernalization is not an
abso ses requirement i Sudan: Life cycles of vernalized plants are shorter than
those d before the onset of summer drought.]
Sec en. J. Die ausdauernden Arten der Sectio Eualyssum aus der Gattung
Alyssum, I. Beil. Jahresb. Nied.-Ost. Land-Lehrers., Wiener-Neustadt 34: i-xiv +
1-35. 1907.* [Part II in ibid. 35: 1-58. 1908*; part tl in ibid. 36: 1-38. 1909*: part
IV in Jahresb. Kaiser Franz Josef-Land.-Gymn. Oberreals., Baden 48: 1-18. 1911.*
According to DupDLEy (1963, p. 36), Baumgartner’s work, which deals with the
1987] AL-SHEHBAZ, ALYSSEAE 201
perennial species of sect. A/yssum, is valuable for its accurate diagnoses of species
and for its detailed discussions. It has received little attention, however, because it
was published in obscure annual reports of “high schools.”
BERGDOLT, E. Uber die Bliitenbiologie von ca montanum und ihre Zweckmas-
sigkeitsdeutungen. Flora 125: 217-231.
BGcueEr, T. W., & K. LARSEN. i etary oe cytological studies on plant species.
IV. Further studies in short-lived herbs. Biol. Skr. Dan. Vid. Selsk. 10(2): 1-24.
1958. [A. Alyssoides, A. montanum, 14-16.
Bonnet, A. L. M. Contribution a l'étude caryologique du genre Alyssum L. (s. lat.).
Nat. Monspel. Bot. 15: 41-52. 1963. [Aurinia, Alyssum, Lobularia.
Brooks, R. R., R. S. Morrison, R. D. Reeves, T. R. DUDLEY, & Y. AKMAN. Hyper-
accumulation of nickel by A/yssum L. (Cruciferae). Proc. Roy. Soc. London B. 203:
387-403. 1979. [Analysis of 167 species for nickel content. Correlation between
species diversity and endemism and high nickel concentrations. The chemical data
support raising sect. Odontarrhena to generic rank.
oRD. Nickel accumulation by European species of the genus A/ys-
sum. Proc. Roy. Soc. London B. 200: 217-224. 1978. [Analysis of 64 species for
nickel and cobalt content. All except one of the 14 hyperaccumulators belong to
sect. Odontarrhena.|
. SHAW, & A. Asenst MarFIL. The chemical form and physiological function
of nickel i in some Iberian A/yssum species. Physiol. Pl. 51: 167-170. 1981a. [Phys-
iology of 2 aera pO of nickel in two subspecies of A. serpyllifolium.
: ce) bservations on the ecology, metal uptake and nickel
tolerance of eee serpyllifolium ie from the Ae peninsula. Vegetatio
45: 183-188.
CAYOUETTE, R. aie ons a la flore adventice du ee Nat. Canad. 99: 135, 136.
1972. [First record a murale for North Ameri
CONTANDRIOPOULOS, J., & Z. AFZAL-RAFII. ae ihre a l’étude cytotaxinomique des
lyssum de oe rite summary.) Bull. Soc. Bot. Suisse 83: 14-29. 1973.
[Chromosome counts for 24 species, role of polyploidy in the evolution of the genus,
geographic distributions of diploid and polyploid races ie certain species.]
Dawe, J. C., & D. F. Murray. Jn: A. Léve, ed., Chro ome number reports LXX.
Taxon 30: 68-80. 1981. [A. americanum, 71, 2n ~ 30
Dupb_ey, T. R. Some new Alyssa from the Near Fast. Notes Bot. Gard. Edinburgh 24:
157-163. pls. 6, 7. 1962. [Six new taxa
. Taxonomic studies in the Cruciferae of the Near East with particular reference
to the systematics of the genus A/yssum in Turkey. 687 pp. + 8 figs. + +
31 maps. Unpubl. Ph.D. dissertation, Univ. Edinburgh, U. K
Studies in A/yssum: Near Eastern representatives and their allies, 1. Jour. Arnold
Arb. 45: 57-100. 1964a. [Numerous new taxa and new combinations, nomenclature
of A. Alyssoides and A. minus. ]
. Synopsis of the genus A/yssum. Ibid. 358-373. 1964b. [Recognized 164 species
n six sections, three subsections, and four series; notes on 16 species of doubtful
status; distributions
ynopsis of the genus Aurinia in Turkey. Ibid. 390-400. 1964c. [Differences
neti Alyssum and Aurinia, seven species, key, descriptions, distributions
—. Alyssum turgidum: a new species from Iran. Great Basin Nat. 24: 7-12. 1964d.
— . Studies in A/yssum: Near Eastern representatives and their allies, II. Section
Meniocus and section Psilonema. Jour. Arnold Arb. 46: 181-217. 1965a. [Keys,
descriptions, distributions, nee habitats of 12 species; diversity of petals, stamens,
fruits, and trichomes in A/yssu
Alyssum. In: P. H. Davis, a Fl. Turkey 1: 362-409. 1965b. [Recognized 89
species in five sections; figs. 17, 18, ma
—. Ornamental madworts (A/yssum) and the correct name of the goldentuft alys-
202 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
sum. Arnoldia 26: 33-45. 1966. [Differences between A/yssum and Aurinia; de-
scriptive list of 20 species of A/yssum available ee with notes on habit,
leaves, flower color, fruits, flowering, and native range.
Alyssum (Cruciferae) introduced in North America. Rhodora 70: 298-300. 1968.
(A. Alyssoides, A. desertorum, A. strigosum, A. minus var. micranthum, A. Szowit-
sianum. |
A new nickelphilous species of A/yssum (Cruciferae) from Portugal: Alyssum
Pintodasilvae T. R. Dudley. Feddes Repert. 97: 135-138. 1986. [A related paper in
ibid. 139-141.)
GABBRIELLI, R., R. BrRTOLO, & O. VERGNANO GAmai. Evaluation of nickel tolerance in
Alyssum. Atti Soc. Tosc. Sci. Nat. Mem. B. 88: 143-153. 1981 [1982]. [Nickel
tolerance in four species is measured by root elongation and protoplasmic resistance
of epidermis. ]
GrREUTER, W. Note on two Greek varieties of A/yssum Doerfleri (Cruciferae) and on the
classification of some perennial species of the genus. (In French; English summary.)
Candollea 29: 135-146. 1974. [Suggested that ser. Libera of sect. Gamosepalum be
transferred to sect. A/yssum.]
Hicains, R. S. What’s in a name? A/yssum. Garden 4(4): 22. 1980. [Derivation of
generic name, early medicinal uses.
ILytnskA, A. F. Chromosomal numbers of certain Ukrainian species of the genus Alys-
sum L. (In Ukrainian; English summary.) Ukr. Bot. Zhur. 32: 371, 372. 1975.
[Counts for four species.
Kyaer, A., & R. GMELIN. Isothiocyanates XIX. L(-)-5-Methylsulphinylpentyl isothio-
cyanate, the aglucone of a new naturally occurring glucoside (glucoalyssin). Acta
Chem. Scand. 10: 1100-1110. 1956. [Distributions of four glucosinolates in 11 tax
now placed in Alyssoides, Alyssum, Aurinia, and Lobularia.]
KRASNOBOROV, I. M. New species of the genus A/yssum L. from Tuva A.S.S.R
Zhur. 60: 663, 664. 1975. [A new species and the new monotypic section Stevenioides
are described.
Locan, L. A. A list of seed plants of Lincoln Parish, Louisiana. Proc. Louisiana Acad.
Sci. 26: 18-32. 1963. [A. Alyssoides, 23.]
Morrison, R. S., R. R. BRooxs, & R. D. Reeves. Nickel uptake by Al/yssum species.
Pl. Sci. Lett. 17: 451-457. 1980.*
Mozinco, H. N. Two European invaders. The source of two alyssums is a puzzle.
Mentzelia 3: 32, 33. 1978. [4. Alyssoides and A. strigosum in Nevada.]
NyArApy, E. J. Vorstudium iiber einige Arten der Section Odontarrhena der Gattung
Alyssum. Bull. Grad. Bot. Cluj 7: 3-51, 65-160. pls. 1-10. 1927; 8: 152-156. 1928:
9: 1-68. 1929.]
. Uber einige westmediterrane A/yssum-Arten. Bul. Soc. Stiinte Cluj 6: 446-460.
1932."
———.. Synopsis speciecum, variatonum et formarum sectionis Odontarrhenae. Generis
Aivssum: Anal. Acad. Repub. Pop. Romane A. 3, 1(separate): 1-130. pls. 1-6. 1949.*
Pancaro, L., M. INNAMORATI, O. VERGNANO GAmBI, & S. OCCHIOCHIUSO. Effects of
cobalt, nickel, and chromium on germination of A/yssum during afterripening and
aging. (In Italian; English summary.) Giorn. Bot. Ital. 115: 265-284, 1981. [The
serpentine endemics 4. argenteum and A. Bertolonti are more tolerant than the
limestone inhabitant 4. nebrodense to nickel and chrom J
, P. PELost, O. VERGNANO GAMBI, & C. GALOPPINI. Seite contribution on the
relationship between nickel and malic and malonic acids in A/yssum. (In Italian;
English summary.) Giorn. Bot. Ital. 112: 141-146. 1978.
Persson, J. Studies in the Aegean flora XIX. Notes on A/yssum and some other genera
of Cruciferae. Bot. Not. 124: 399-418. 1971. [Alyssoides, Alyssum, Aurinia, Car-
damine, Iberis, Ricotia.]
1987] AL-SHEHBAZ, ALYSSEAE 203
Reeves, R. D., & R. R. Brooks. Hyperaccumulation of lead and zinc by two metal-
lophytes from mine areas of central Europe, Phase) rotundifolium, Alyssum Wul-
fenianum. Environ. Poll. A. 31: 227-285. 1983.
Rotuns, R. C. Some new or noteworthy North American crucifers. Contr. Dudley
Herb. 3: 174-183. 1941. ae report of A. desertorum, 183.
ScHuLz, O. E. Uber die Gattung Gamosepalum Hausskn. Notizbl. Bot. Gart. Berlin 10:
109-111. 1927. [Recognized three species; genus is reduced to a section of A/yssum,
see DuDLEY (1964b).]
SUPAVARN, P., F. W. Knapp, & R. SiGAFus. Investigations of mucilaginous seeds as
potential biological control agents against mosquito larvae. Mosq. News. 36: 177-
182. 1976. [Mucilage from seeds of 11 species of Alyssum caused up to 85 percent
mortality among larvae of Aédes aegypti. ]
Toma, C. The morphological-anatomical features of Alyssum Borzaeanum Nyar. (In
French.) Feddes Repert. 88: 477-489. 1977. [Anatomy of root, stem, and leaf.]
TurrILL, W. B. Alyssum campestre. Jour. Bot. London 73: 261, 262. 1935. [4. Alys-
soides. |
VERGNANO GamplI, O. First data on the histological localization of nickel in Alyssum
Bertolonii Desv. (In Italian; English summary.) Giorn. Bot. Ital. 101: 59, 60. 1967.
[Nickel is accumulated in the epidermis and the sclerenchyma between the vascular
bundles of ie stem.]
Ro REE
: . RADFORD. Nickel een by Italian species of the
genus ee. (in Italian; English summary.) W a 33: 269-277. 1979. [Tests
for accumulation in 13 species of A/yssum and in ee now placed in Al/yssoides,
Aurinia, Lobularia, and Berteroa; a related paper in ibid. 32: 175-188. 1977.]
ZoHARY, M. Carpological notes on A/yssum. sa Jour. Bot. Jerusalem Ser. 4: 239,
240. 1949. [Seed dispersal in the annual spec
. Anatomical-carpological on aoa in some eae plants
of the oriental flora. Palestine Jour. Bot. Jerusalem Ser. 2: 125-131.
damascenum, A. marginatum, A. pyramidatum, A. Szowitsianum, 129- re
22. Lobularia Desvaux, Jour. Bot. II. 3: 162. 1815, nom. cons.°
Annual or perennial canescent herbs [rarely subshrubs], densely to sparsely
covered with a uniform indumentum of sessile, appressed, bifid, medifixed
trichomes. Stems erect to prostrate, branched from the base or above. Leaves
entire, short petiolate, linear, oblong, lanceolate, or spatulate, always attenuate
at base. Inflorescences terminal, usually ebracteate (or the lowermost flowers
subtended by leaflike bracts), densely flowered, corymbose racemes, usually
greatly elongated in fruit. Sepals oblong or ovate, obtuse, always spreading,
equal, not saccate at base, densely pubescent. Petals white or rarely pink or
purple, clawed, suborbicular to spatulate or obovate, entire, about twice as
long as the sepals. Nectar glands 8, filiform to subclavate; median glands 4,1
outside each median stamen; lateral glands smaller, | on each side of lateral
stamens. Stamens 6, tetradynamous; filaments free, strongly dilated at base,
toothless, neither appendaged nor winged; anthers ovate. Ovary pubescent,
2- [to 12-]ovulate; ovules on subapical [or parietal] placentae. Fruits dehiscent,
‘The year of publication has been wrongly given in all floras as 1814. According to ean &
Cowan’s Taxonomic Literature (Regnum Veg. 94: 634. 1976), the year of publication of the above
page of Desvaux’s Journal was 1815. Lobularia is conserved, and the earlier generic names ye
and Konig we Adanson (Fam. Pl. 2: 420. 1763) are rejected because the last name was not Latinized
and Aduseton was spelled in two ways by Adanson, who added further confusion in his prefatory errata
(p. 23) by acon these names.
204 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
flattened parallel to the septum, sessile or short stipitate, elliptic, ovate, orbic-
ular, [oblong, or obovate]; valves obscurely nerved, glabrous or pubescent:
styles persistent, short; stigmas capitate. Seeds | [2-6] per locule, narrowly [to
broadly] winged, compressed, minutely reticulate, mucilaginous when wet;
cotyledons accumbent. Base chromosome numbers 11, 12. (Including Konig
Adanson, Aduseton Adanson, Koniga R. Br., Glyce Lindley.) Type SPECIES:
Clypeola maritima L. = L. maritima (L.) Desv. (Name from Latin lobulus, a
little lobe, referring to the small fruit, but some authors (e.g., Fernald) maintain
that the name probably refers to the 2-lobed (bifid) trichomes.) — SWEET ALYSSUM.
A genus of four species distributed primarily in the Mediterranean region
and the Macaronesian archipelago (Azores, and the Salvage, Canary, and Cape
Verde islands). One species, Lobularia maritima (L.) Desv. (Clypeola maritima
L., Alyssum maritimum (L.) Lam., Koniga maritima (L.) R. Br., A. minimum
L.), sweet alyssum or alison, 2” = 24, is an ornamental widely cultivated
throughout the world, an escape from cultivation, and a naturalized weed in
the southeastern United States. It grows in waste places and lawns and on
cultivated grounds in the Carolinas, Florida, Tennessee, Mississippi, and Lou-
isiana. Lobularia maritima is an annual or perennial herb under cultivation,
but in its native habitat in the Mediterranean region, Madeira, and the Canary
Islands, where it occupies sea cliffs or sandy areas at sea level, it is always a
perennial with a woody base and is sometimes a subshrub.
Earlier authors (e.g., De Candolle (1821, 1824), Bentham & Hooker, Baillon)
treated Lobularia as a subordinate (often as a section) of the closely related
Alyssum. There are, however, several morphological differences that support
its treatment as a distinct genus. Lobularia has bifid trichomes, eight nectaries
characteristically arranged (see above), spreading sepals, and toothless and
unappendaged staminal filaments. 4/yssum always has stellate trichomes, a
different arrangement of the nectaries, toothed or appendaged filaments (except
in five species of sect. PsLonema), and erect sepals (except in a few species).
Two other genera of the Alysseae, Farsetia and Bornmuellera Hausskn., have
trichomes similar to those of Lobularia, but they are easily separated by their
dentate staminal filaments and strongly 2-lobed stigmas, respectively.
The identification of species of Lobularia relies heavily on the number of
seeds per locule and on habit. Fragmentary specimens that lack mature fruits
are often difficult to identify. As in several other genera of the Cruciferae,
woody habit may have evolved in connection with insular isolation. All of the
five taxa occurring in the Macaronesian archipelago are suffruticose perennials
and under favorable conditions often become subshrubs. Annual habit, which
is considered by Borgen (1984) to be derived in the genus, is found in two
desert species, L. arabica (Boiss.) Muschler (Egypt, Israel) and L. libyca (Viv.)
Meisner (Canary Islands, southern Spain, all of North Africa, Israel, and south-
ern Iran). Lobularia libyca, the most widely distributed species in the genus,
has the largest fruits, with up to six seeds per locule. Lobularia maritima, on
the other hand, has the smallest fruits, with only one seed per locule. The fourth
species, L. intermedia Webb & Berth., is intermediate between L. maritima
and L. libyca in fruit size and in the number of seeds per locule. It is highly
1987] AL-SHEHBAZ, ALYSSEAE vA)
polymorphic, particularly in leaf morphology, fruit shape, and seed number.
It was subdivided into several poorly defined varieties, the identities of which
need critical evaluation. Lobularia spathulata (J. Schmidt) O. E. Schulz (Cape
Verde Islands), L. marginata Webb & Berth. (high crests of the Anti Atlas
Mountains, Morocco, and of Lanzarote and Fuerteventura, Canary Islands),
and L. palmensis Webb ex Christ (eastern Canary Islands) are separated from
L. intermedia on the basis of minor characters. They can be hybridized easily
with each other and with L. intermedia, and the first generation hybrids show
pollen fertility higher than 80 percent (Borgen, 1984). Therefore, they should
be recognized as infraspecific taxa of L. intermedia.
Consistent counts of 2n = 24 were reported for Lobularia maritima from
many Mediterranean countries. However, Borgen (1984) also recorded 2” =
22 and observed meiotic irregularities such as univalent and multivalent for-
mations, chromosomal bridges, and lagging chromosomes. On the basis of
these meiotic irregularities, particularly the frequent occurrence of univalents,
Borgen (1984) suggested that L. maritima is probably an allopolyploid, but he
did not indicate what its ancestral species were. He considered L. maritima to
be the most primitive member of the genus despite its 1-seeded fruits that are
the smallest in Lobularia. This species and the L. intermedia complex are self-
incompatible, large-flowered, suffruticose perennials, whereas L. /ibyca and L.
arabica are autogamous, small-flowered annuals. Uniform counts of 2” = 22
have been reported for members of the L. intermedia complex, as well as for
L. libyca (Borgen, 1970, 1974, 1984; Larsen). The count of n = 6 for L. libyca
by Negodi may be an error. Snogerup and Borgen (1984) reported 2n = 46
and 2n = 42, respectively, for L. arabica.
Seed-coat anatomy of Lobularia maritima differs from that of A/yssum in
the cell-wall thickening of the palisade layer. In Lobularia the cells are thin
walled, while in A/yssum they are either evenly thickened throughout or the
radial and/or inner tangential walls are thickened (Vaughan & Whitehouse).
Only Lobularia maritima has been surveyed for seed glucosinolates and fatty
acids. High concentrations of 3-butenylglucosinolate, smaller amounts of
6-methylthiohexyl and 4-pentenyl glucosinolates, and traces of allyl, benzyl,
and 2- phenylethy! glucosinolates were identified (Hasapis et a/.). Kjaer & Gme-
lin found 5 late to be the major component of
the species. Although the fatty- acid composition of L. maritima resembles that
of Alyssum in lacking erucic acid, it is markedly different in its high concen-
trations (42 percent) of eicosenoic acid and small amounts (10 percent) of
linolenic acid. A/yssum contains only traces (0.4 percent or less) of the former
acid and 36-66 percent of the latter (Kumar & Tsunoda). These observations,
however, are based on an incomplete sampling of both genera.
Lobularia maritima growing in its natural habitats is highly variable in habit,
leaf succulence, and resistance to salinity. Seashore populations are low-grow-
ing, bushy plants with broad, thick leaves and are resistant to salinity; inland
ones are taller and generally erect plants with thin, linear leaves and are in-
tolerant of salinity (Catarino ef a/.). Leaf succulence can be induced experi-
mentally by prolonged treatment with 0.2 M sodium chloride. Such treatment
increases the cell volume, nucleus size, and DNA content (often accompanied
206 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 68
a oe in both palisade and spongy parenchyma (Capesius &
en).
oie literature indicates that Lobu/aria maritima was used as an astringent,
an antiscorbutic, a diuretic, and a febrifuge. The species is the most widely
cultivated of any ornamental crucifer. It is grown as a border plant and has
sweet-smelling, white or purple flowers. It is also a naturalized weed in many
parts of the world.
REFERENCES:
Under family references in AL-SHEHBAz (Jour. Arnold Arb. 65: 343-373. 1984), see
BAILLON; BENTHAM & HOOKER; BERGGREN; CAIUS; DE CANDOLLE (1821, 1824);
DAXENBICHLER et al/.; FERNALD; Hasapis et al.; VoN HAYEK; JONES; LA PorTE; MAIRE;
MepveE; PANT & Kipwal; QuUEIROS; ROLLINS (1981); ScHULz; and VAUGHAN & WHITE-
HOUSE
Under tribal references see BAILEY; BOLKHOVSKIKH et al.; GATTINGER; GOLDBLATT
(1981, 1984); KUMAR & TSUNODA; KUPFER; LITCHFIELD; MACROBERTS; Moore; REEVES
et al., and SMALL. Under references to A/yssum see KJAER & GMELIN.
Bau, P. N., & S. L. TANDON. A preliminary study of colchicine-induced polyploids of
Alyssum maritimum Lam. Curr. Sci. Bangalore 27: 407, 408. 1958a.
sis in A/yssum maritimum. Indian Jour. Hort. 15: 22-25. 1958b.*
& Mo srphological and cytological studies of the induced polyploids in
Alyssum maritimum Lam. Genetica 30: 129-139. 1959. [Colchicine-induced poly-
ploids showed increase in the size of pollen, stomata, fruits, flowers, and seeds;
meiotic irregularities; n = 12.
BARTOLO, G., S. BRULLO, & P. Pavone. Numeri cromosomici per la flora italiana: 617-
631. Inf. Bot. Ital. 11: 149-159. 1979. [L. poe 149, 2n = 24; L. libyca, 149,
150, 2n = 22; figs. 1
BorcGen, L. Chromosome numbers of vascular plants from the Canary ee with
special reference to the occurrence of polyploidy. Nytt. Mag. Bot. 16: 81-121. 1969.
[L. maritima, 95, 97, 2n = 22, fig. 68; suggested x = 11 as the base pei
number = eas ]
. Chro e numbers of Macaronesian flowering age | 17: 145-161.
1970. [L. ome and L. libyca, 154, 155, 2n = 22, figs. 5
r me numbers of Macaronesian pace II. one Jour. Bot. 21:
195- 210. 1974. [L. marginata, 198, 2n = 22, fig. |
Chromosome numbers and fertility acne in Lobularia, Cruciferae. A
preliminary report. Webbia 38: 645-653. 1984. [Seven taxa in six species
Capesius, I., & S. LoeBeN. Changes of nuclear DNA composition after induction of
succulence i in Lobularia maritima. Zeitschr. Pflanzenphysiol. 110: 259-266. 1983.
CaATARINO, F. M. Endopolyploidy and differentiation. Experimental induction of en-
dopolyploidy in Lobularia maritima (L.) Desv. and Bryophyllum crenatum Bak. (In
Portuguese; English summary.) Portug. Acta Biol. A. 11: 1-218. 1968. [Natural and
induced pee re chromosome numbers, ultrastructure, effects of sodium
chloride, figs. 1-67.]
& I. Capestus. Changes in uptake of labelled precursors into DNA ae de-
velopment of salt succulence in Lobularia. Portug. Acta Biol. A. 15: 59-74. 1979.
. Martins, & C. MEDEIRA. Ecotypic variation in Lobularia ann (L.)
es sv. Bol. Soc. Brot. 47(Suppl.): 339. 1974.
CHoprA, R. N., & S. P. RATNAMBA. Morphogenic studies on stem segments of Lobularia
maritima Desv. Phytomorphology 25: 490-492. 1976. [Indole acetic acid induced
callus and root formation, kinetin-induced shoot formation.]
1987] AL-SHEHBAZ, ALYSSEAE 207
CRUTCHFIELD, P. J. Taxa collected from Roanoke Island new to the flora of North
Carolina. Castanea 29: 129-137. 1964. [L. maritima, 133.
Gort, S., A. K. Mupaa, & S. C. Gupta. In vitro induction of divisions in pollen,
callus formation and plantlet a in anthers of Lobularia maritima. Zeitschr.
Pflanzenphysiol. 104: 187-191. 19
KHANNA, R., & R. N. CHopra. Regula - of shoot-bud and root formation from stem
explants of mane maritima. Phytomorphology 27: 267-274. 1978. [Auxin and
cytokinin treatmen
LarsEN, K. Cytological a experimental studies on the flowering plants of the Canary
Islands. Biol. Skr. Dan. oe ee 11(3): 1-60. pls. 1-6. 1960. [L. intermedia, 6,
7,n = 11, 2n = 22, figs. 2 A
Les, K., & C. HOLZAPFEL. ae o the Canary Islands: the Cruciferae, the eae
and the ferns and their allies. Anal. Inst. Nac. Invest. Agrar., Ser. Prod. Veg. 4
273. 1974. [Lobularia, 183, 184, 198.]
MARTINS- Lou¢Ao o, M. A., & F. M. CaTarino. Nuclear changes associated with callus
induction in Lobularia maritima. Bol. Soc. Brot. 53: 1211-1221. 1981. [Callus
originates from cambial cells of the vascular bundles of leaves, endopolyploidy.]
Necop1, G. Contributo alla cariologia dei generi /satis L. e Lobularia Desv. (Cruciferae).
Atti Mem. Accad. Naz. Sci. Let Peek Modena VI. 7: 45-52. 1965. [Lobularia, 49-
52; L. libyca, n = 6; L. rie = 12.]
PRABHAKAR, K., & M. R. VAYARAGHAVAN. Endothelium in /beris amara and Alyssum
maritimum—its histochemistry and ultrastructure. Phytomorphology 32: 28- 36.
1983a
& mbryo sac ee in Iberis amara and Alyssum maritimum. Phyton
Austria 23: 31 a pls. 1-3. b.
—. Histoche saree ae ultrastructure of suspensor cells in A/yssum mari-
timum. Cytologia 48: 389-402. 1983c
Ronbet, P. Organogenesis in the course of embryogenesis in Alyssum maritimum Lamk.
(In French.) Compt. Rend. Acad. Sci. Paris 255: 2278-2280. figs. 1-13. 1962.
SIKKA, K. Chromosome pee OF two species of Lobularia (Cruciferae). Curr. Sci.
Bangalore 46: 681-683. 1977
Snocerup, B. In: A. Love, ed., Chromosome number reports LXXXIX. Taxon 34:
727-730. 1985. [L. arabica, 727, 2n = 46.]
VUAYARAGHAVAN, M. R., & K. PRABHAKAR. Ontogenetical and histochemical studies
on chalazal proliferating tissue in /beris amara and Alyssum maritimum. Beitr. Biol.
Pflanzen 56: 7-17. 1982.
,&
Histochemical, structural and ultrastructural features of
endosperm in Alyssum maritimum Lam. Acta Bot. Neerl. 33: 111-122. 1984.
23. Berteroa A. P. de Candolle, Syst. Nat. 2: 290. 1821.
Annual or perennial herbs, densely pubescent with stellate trichomes mixed
with fewer simple or bifid ones. Basal leaves petiolate, entire or occasionally
repand or sinuate; upper cauline leaves sessile, entire. Inflorescences ebracteate,
densely flowered, corymbose racemes, greatly elongated in fruit; fruiting ped-
icels erect-ascending [or divaricate], straight or curved. Sepals ascending to
spreading, oblong, not saccate at base, densely pubescent, with or without a
subapical tuft of simple trichomes. Petals white [or yellow], attenuate into a
clawlike base, deeply emarginate, the sinus extending to nearly half the length
of blade. Lateral nectar glands 4, | on each side of each lateral stamen; median
glands absent. Stamens 6, tetradynamous; lateral filaments with a basal, adaxial
appendage; median filaments dilated at base, neither appendaged nor winged;
208 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
anthers oblong, slightly exserted. Fruits sessile, 1-3 times as long as broad,
elliptic, oblong, ovate [or orbicular], compressed parallel to the septum; valves
with obscure midvein, inflated [or not], densely pubescent with appressed
stellate trichomes [or glabrous]; styles persistent; stigmas capitate, obscurely
2-lobed, wider than the style. Seeds 2-6 per locule, compressed, suborbicular
to obovate, margined [or conspicuously winged], nonmucilaginous when wet;
cotyledons accumbent. Base chromosome number 8. (Including Myopteron
Sprengel.) LecroryPe species: A/yssum incanum L. = B. incana (L.) DC.; see
Britton & Brown, Illus. Fl. No. U. S. & Canada, ed. 2. 2: 153. 1913. (Name
honoring Carlo Giuseppe Bertero, Oct. 14, 1789-April 9, 1831, Italian phy-
sician and botanist of Piedmont, who traveled in the West Indies (1816-
1821), settled in Chile in 1827, and died in a shipwreck in the southern Pa-
cific.) — HOARY ALYSSUM.
A genus of five species centered in the Balkan peninsula and distributed from
central Europe eastward into Turkey and the Caucasus. A few authors expand
the limits of the genus to include Berteroa spathulata (Stephan ex Willd.) C. A.
Meyer (central Asia and western Siberia), B. Potaninii Maxim. (Mongolia),
and B. ieee Ikonn.-Galitz. (Mongolia and central Asia). However, these
are m g y different and geographically disjunct from the remaining
species ae Berteroa. They have been transferred recently to a new segregate,
Galitzkya V. Bocz., which differs from Berteroa in its subscapose habit, un-
appendaged filaments, and uniform pubescence.
Berteroa is represented in North America by two naturalized weeds, one of
which is sporadic in the southeastern United States. Berteroa incana (L.) DC.
(Alyssum incanum L., Farsetia incana (L.) R. Br., Draba cheiranthifolia Lam.),
hoary alyssum, 2 = 16, usually grows on dry sandy or gravelly soils in meadows,
pastures, waste places, and fields, as well as along roadsides, railroad tracks,
streams, and riverbanks. It was probably introduced into North America with
either grass or clover seeds or in ballast (Martindale). Although B. incana was
recorded from Tennessee as early as 1901 (Gattinger), it has been reported only
recently from Arkansas, Kentucky, and Virginia. It is most abundant in the
northeastern United States and is noxious in Minnesota and Michigan.
Berteroa mutabilis (Vent.) DC. is sporadically distributed in the United
States and is naturalized in parts of Massachusetts, New York, and Kansas.
Brooks’s record of B. obliqua (Sibth. & Sm.) DC. from the Catskill region, New
York, is based on a misidentified plant (True 78, nys!) of B. incana. The former
grows only as a native in Italy and the Balkan peninsula. The remaining species
of the genus, B. Gintlii Rohlena and B. orbiculata DC., are endemic to Yu-
goslavia and the Balkan peninsula, respectively.
Although some earlier authors (e.g., Bentham & Hooker, Baillon) reduced
Berteroa to a section of Alyssum, the two genera are not closely related. Von
Hayek suggested that Berteroa is directly derived from Fibigia, while Schulz
placed it between Lobularia and Lepidotrichum Velen. & Bornm. (= Aurinia).
Obviously, the relationships between these and several other genera of the
Alysseae have not been fully established. Berteroa is distinguished by its deeply
bifid petals, appendaged lateral staminal filaments, mixed indumentum of stel-
late and bifid trichomes, and two to six seeds in each locule.
1987] AL-SHEHBAZ, ALYSSEAE 209
Very little is known about the floral biology of the genus. Knuth indicated
that Berteroa incana is protogynous. Autogamy occurs as a result of contact
between the median anthers and the stigma. In Europe the species is pollinated
by several species of flies, particularly of the genera Eristalis, Rhingia, Syritta,
and Syrphus, as well as by species of the butterfly genus Vanessa and the bee
Halictus. Bateman listed one species of Berteroa (without name) as self-incom-
patible.
Chromosome numbers are known for all species except Berteroa Gintlii. The
genus is uniformly based on x = 8, and all species are diploid. No interspecific
hybridization has been reported.
The seeds of Berteroa incana contain very high concentrations (89 percent)
of C,, fatty acids, of which linolenic acid is the major constituent (48 percent),
and no traces of erucic acid (Appelqvist). Goering and colleagues considered
the species to be agronomically acceptable and a good source of drying oils.
The seedlings have large and small amounts of benzyl and isopropyl glucosi-
nolates, respectively (Cole), while the seeds contain 5-methylsulfinylpentyl,
5-methylthiopentyl, 4-pentenyl, and 2-hydroxy-4-pentenyl glucosinolates
(Daxenbichler et al, Kjaer). The remaining species of Berteroa have not been
surveyed for fatty acids or glucosinolates.
Seed-coat anatomy of Berteroa incana is indistinguishable from that of B.
obliqua. The epidermis in both has large columns with markedly flattened tops
and hollow centers, while the palisade layer has isodiametric cells with strongly
thickened radial and inner tangential walls (Vaughan & Whitehouse).
Except for the weedy Berteroa incana and B. mutabilis, the genus has very
little economic importance. The leaves of Berteroa are said to be eaten as a
salad (Crisp)
REFERENCES:
Under family references in AL-SHEHBAZ one Arnold Arb. 65: nome 1984), see
APPELQVIST (1971); BAILLON; Seuregn ge NTHAM & HOOKER; BERGGREN; BRITTON
Brown; BuscH; DE CANDOLLE (1821, 1824). CoLe (1976); Crisp; oes
HAYEK; KGAER: KNUTH; Nie Soon MARKGRAF; MUENSCHER; MULLIGAN (1957); ROuNS
(1981); ScHuLz; E. B. SmirH; and VAUGHAN & WHITEHOUSE.
Under tribal references see BOLKHOVSKIKH ef al.; CONTANDRIOPOULOS,; DUDLEY &
CULLEN; GATTINGER; GOLDBLATT (1981, 1984, 1985); KuMaAR & TSUNODA; Moore; and
REEVES ef al
AncEv, M.E. Jn: A. Love, ed., Chromosome number ae LX XIII. Taxon 30: 829-
861. 1981. a incana, 2n = 16, B. mutabilis, 2n = 55.
BALL, P. W. Berteroa. In: T. G. Tutin et al., eds., oe Burondes 1: 305. 1964. [Five
species recognized. ]
BELYAEVA, L. E., & N.S. Fursa. Formation of male and female structures of Berteroa
incana (L.) DC. flower. (In Russian; English summary.) Ukrain. Bot. Zhur. 36: 574-
577. 1979.
BoczANTZEVA, V. The new genus Galitzkya V. Boczantzeva (Cruciferae). (In Russian.)
ot. Zhur. 64: 1440-1442. 1979. [Transfer of three central Asiatic and Mongolian
species of Berteroa to Galitzkya; see IKONNIKOV-GALITZKY. |
Brooks, K. L. A Catskill flora and economic botany. IV. (Part 1.) Polypetalae—Cheno-
podiaceae through Capparidaceae. New York State Mus. Bull. 453. xiii + 358 pp.
1983. [Berteroa, 210, 211, 336, 337.
210 JOURNAL OF THE ARNOLD ARBORETUM [voL. 68
CRANFILL, R., & J. W. THIERET. Thirty additions to the vascular flora of Kentucky. Sida
9: 55-58. 1981. [B. incana, 57.]
DAXENBICHLER, M. E., W. P. SCHROEDER, & G. F. SPENCER. (+)-5-Allyloxazolidine-2-
ione, an enantiomer of turnip antithyroid factor isolated from Berteroa incana
(L.) DC. Jour. Agr. Food Chem. 30: 1248-1250. 1982.
Dewey, L. H. Three new weeds of the mustard family. Circ. Bot. we S. Dep. Agr. 10.
6 pp. 1897. [B. incana, the average plant produces ca. 5000 see
GokrRING, K. F., R. Estick, & D. L. BRELSFORD. The composition of rs oil of Berteroa
incana and the potential value of its seed as a cash crop for Montana. Econ. Bot.
19: 44, 45. 1965.
Hastinas, R. E., & C. A. Kust. Reserve carbohydrate storage oa era ie yellow
rocket, white cockle, and hoary alyssum. Weed Sci. 18: 148. 1970.* [B. incana.]
IKONNIKOV-GALITzKY, N. P. A new species of the family see ee in the Moneoliae
flora. (In Russian; English summary.) Acta Inst. Bot. Acad. Sci. URSS. I. 3: 189-
193. 1937. [B. macrocarpa, sp. nov.; key to three central Asiatic species; see
BOCZANTZEVA. |
KgAer, A., I. LARSEN, & R. GMELIN. Isothiocyanates XIV. 5-Methylthiopentyl isothio-
cyanate, a new mustard oil present in nature as a glucoside (glucoberteroin). Acta
Chem. Scand. 9: 1311-1316. 1955. [B. incana
Kust, C. A. Selecti ntrol of hoary ee in alfalfa. Weed Sci. 17: 99-101. 1969.*
Loon, J. C. VAN, & H. DE Jonc. Jn: A. Léve, ed., IOPB eo aaa number reports
LIX. Taxon 27: 53-61. 1978. [B. ae 57, 2n = 16.]
MARTINDALE, I. C. The introduction of foreign plants. te Gaz. 2: 55-58. 1876. [B.
incana (as Alyssum) on ballast near Philadelphia, 57.]
Mucina, L., & D. BRANDES. Communities of Berteroa incana in Europe and their
pa aea ta differentiation. Vegettio 59; 125-136. 1985. Acne study;
two geographic races recognized.]
TSELINKO, S. A., Y. K. ie as K, & N. S. Fursa. Flavonoids of Berteroa incana L.
Chem. Nat. Comp. 9: 765. (English transl.) 1975. [Rhamnocitrin, kaempferol, and
quercetin. }
24. Draba Linnaeus, Sp. Pl. 2: 642. 1753; Gen. Pl. ed. 5. 291. 1754.
Annual, biennial, or most commonly perennial herbs, usually with much-
branched caudices. Stems simple or branched, scapose or foliose. Trichomes
simple, furcate, cruciform [malpighiaceous, pectinate, stellate, or dendritically
branched], usually more than one kind present. Basal leaves petiolate or rarely
sessile, entire or toothed to laciniate [rarely pinnately lobed], usually forming
distinct rosettes in the perennials but rarely so in the annuals. Cauline leaves
(when present) sessile [or petiolate], cuneate [or amplexicaul]. Inflorescences
ebracteate [or bracteate], few- to many-flowered, corymbose racemes, slightly
to greatly elongated in fruit; fruiting pedicels ascending to divaricate [or erect].
Sepals erect to spreading, oblong to elliptic or ovate, not saccate or only slightly
so at base, usually membranaceous at margin, caducous [or persistent], glabrous
or pubescent. Petals present, reduced or absent in some autogamous annuals,
white [yellow, rarely lilac, violet, orange, or red], obovate to spatulate [orbicular
or linear], obscurely to distinctly clawed, the apex obtuse or rounded to truncate,
or shallowly to deeply emarginate, or bifid. Nectar glands tooth- or ringlike,
usually subtending the bases of filaments, median glands sometimes absent.
Stamens 6 [very rarely 4], usually tetrad laments free, unappendaged,
linear, slender or sometimes dilated at base; anthers oblong to ovate, pollinif-
1987] AL-SHEHBAZ, ALYSSEAE 244
erous [or pollen aborted or absent in agamospermous taxa]. Ovary glabrous or
pubescent, [2-] to 80-ovulate. Fruits dehiscent, ovate, lanceolate, elliptic, ob-
long, linear [or orbicular], sessile, flat or spirally twisted, flattened parallel to
the septum, sometimes slightly inflated; valves glabrous or pubescent, usually
with a distinct midnerve and with obscurely to prominently anastomosing
lateral nerves; septum membranaceous, complete, usually not veined; styles
persistent, long to short or obsolete; stigmas capitate, entire or 2-lobed. Seeds
[1-] 3-40 per locule, ovate to ellipsoid [or orbicular], usually flattened, light
to dark brown, reticulate, nonmucilaginous when wet, wingless [or very rarely
broadly winged], weakly to strongly biseriately arranged in each locule, pen-
dulous on slender funicles; cotyledons accumbent. Base chromosome numbers
6-12. (Including Abdra Greene, Aizodraba Fourr., Dolichostylis Turez., Dra-
bella Fourr., Drabella Nabélek, Erophila DC., Holargidium Turcz., Leptonema
W. J. Hooker, Nesodraba Greene, Odontocyclus Turcz., Pseudobraya Korsh..,
Stenonema W. J. Hooker, Thylacodraba O. E. Schulz, Tomostima Raf.)
LECTOTYPE sPEcIES: D. incana L.; see M. L. Green, Bull. Misc. Inf. Kew 1925:
51. 1925. Britton & Brown (Illus. Fl. No. U. S. & Canada, ed. 2. 2: 148. 1913)
chose D. verna L. as the lectotype species of Draba. This species, however, 1s
the conserved type of Erophila. (Name from Greek drabe, acrid, used by
Dioscorides to describe the taste of the leaves of certain cruciferous plants
thought by some authors to have been hoary cress, Cardaria Draba (L.) Desv.)
— WHITLOW GRASS.
A natural genus and the largest of the Cruciferae, with some 350 species
distributed primarily in the Northern Hemisphere, particularly in the arctic
and subarctic regions, as well as in the alpine and mountainous portions of the
temperate regions. There are about 65 species in South America distributed at
higher elevations from Colombia and Venezuela southward along the Andes
into Patagonia. Draba is poorly represented in Mexico and Central America
(11 species, six endemic; Rollins, 1984) and in Africa (five species, two endemic;
Atlas Mountains of Morocco and Algeria) and is absent in Australia. More
than 100 species are found in North America and Greenland, and the ranges
of about 20 of these extend into the arctic and subarctic regions of Europe and/
or Asia. The genus is well developed in the Himalayan and Irano-Turanian
regions (ca. 50 and 40 species, respectively), as well as in China and Japan (ca.
35 species), Siberia and central Asia (ca. 30 species), central and northern
Europe (ca. 35 species), and the Mediterranean area (18 species). Draba is
represented in the southeastern United States by seven species, one of which
is naturalized.
The sectional classification of Draba is controversial. Schulz (1927, 1936),
who treated the genus on a worldwide basis, recognized 17 sections, while
Tolmachev (1939) assigned the 91 species occurring in the U.S.S.R. to 29 series
without recognizing sections. Although some of the infrageneric groups rec-
ognized by these authors represent natural assemblages of closely related species,
the boundaries between the majority of them are artificially drawn and clearly
unsatisfactory. Fernald (1934), who was the first to point out weaknesses in
Schulz’s (1927) sectional classification, indicated that his keys to the sections
22 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
and to the species are misleading and impractical. It is beyond the scope of
this flora to present a comprehensive sectional treatment for Draba. The genus
is poorly represented in our area, and I prefer not to recognize any sections
here.
Draba brachycarpa Nutt. (Abdra brachycarpa (Nutt.) Greene), 2n = 16, 24,
is the most widely distributed species in the Southeast. It grows on open clay
soil in lawns, pastures, fields, disturbed areas, waste grounds, and cedar glades,
on limestone rubble, and along roadsides in all of the Southeastern States. It
appears to have restricted distribution in portions of the Florida Panhandle
(Leon, Gadsden, Liberty, and Jackson counties) and in northern Louisiana. Its
range extends west into Texas, north into Kansas, and east into Missouri,
Illinois, Ohio, and Virginia. It is adventive in some of the Mountain and Pacific
states. Draba brachycarpa is easily distinguished from the other annual drabas
in our area by its glabrous, elliptic to oblong-lanceolate fruits 2-6 mm long,
and by its cruciform, sessile trichomes. Diploid and triploid populations based
on x = 8 have been found in Arkansas (Smith, 1969) and Texas (Rollins &
Rtidenberg), respectively.
Draba aprica Beadle (D. brachycarpa var. fastigiata Nutt. ex Torrey & Gray)
is a very close relative of D. brachycarpa. It grows on granite outcrops and
in shallow sandy soils over siliceous rock. It is locally common in open knolls,
woods, and alluvial areas near streams in South Carolina (Lancaster County),
Georgia (Piedmont; Towns, Richmond, Oglethorpe, Cobb, and De Kalb coun-
ties), Arkansas (Drew, Faulkner, Cleburne, Washington, Montgomery, and Polk
counties), eastern Oklahoma (McCurtain and Cherokee counties), and south-
eastern Missouri (Madison and Iron counties). Hitchcock suggested that D.
aprica should be regarded as a variety of D. brachycarpa, but Fernald (1934)
and Rollins (1961) clearly demonstrated that they are sufficiently different to
be treated as distinct species. They do not hybridize in areas of sympatry, and
according to Kral, D. brachycarpa flowers early and is usually in full fruit when
plants of D. aprica start to bloom. Both species are white-flowered annuals
with cruciform trichomes and small fruits to 6 mm long. Draba aprica differs
from D. brachycarpa in its pubescent fruits, stalked trichomes, larger seeds (1-
1.2 mm instead of 0.5—0.8 mm long), and corymbiform lateral branches of the
infructescence (FIGURE 2g, J).
raba ramosissima Desv. (Alyssum dentatum Nutt., D. dentata (Nutt.) W. J.
Hooker & Arnott, D. ramosissima var. glabrifolia E. L. Braun), 2n = 16, is
a mat-forming perennial with much-branched, long caudices covered with
remnants of old leaves and terminated by rosettes of laciniate to subpectinate
leaves. It differs from its relatives with spirally twisted fruits in its paniculate
infructescences with strongly divergent branches and in its styles |-3 mm long.
Draba ramosissima grows primarily on open shale banks, dolomitic bluffs, and
limestone cliffs in North Carolina (Madison and Buncombe counties), Ten-
nessee (Blount and Cocke counties), Kentucky, West Virginia, Virginia, and
Maryland. Gattinger reported it from Polk County, Tennessee, but subsequent
botanists have not confirmed this record. Plants with glabrous to sparsely
pubescent stems and leaves were recognized by Fernald (1934, 1950) as var.
glabrifolia. As shown by both Nye (1961, 1969a) and Reed, however, trichome
1987] AL-SHEHBAZ, ALYSSEAE 213
Ficure 2. Selected species of Draba. a-c, D. platycarpa: a, infructescence, x %; b,
fruit, x 5; c, fruit with | valve removed, x 5. d, D. cuneifolia, infructescence, x %. e,
D. reptans, infructescence, x 4. f, D. ramosissima, infructescence, x 2. g, h, D. bra-
chycarpa: g, fruiting plant, x 1; h, fruit, x 12.1, j, D. aprica: i, fruiting plant, x 1; j,
infructescence, x
214 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
density is highly variable in the species, and both glabrous and pubescent forms
are found within a given population. Schulz (1927) placed D. ramosissima in
sect. PHYLLODRABA O. E. Schulz and assigned its nearest relative, D. arabisans
Michx. (Maine to Newfoundland and westward to Minnesota and Ontario) to
sect. LEUCODRABA DC. He separated these sections mainly on the basis of the
many-leaved stems and yellow flowers in the former vs. the few-leaved stems
and white flowers in the latter. Neither set of characters, however, was carefully
observed or evaluated in either of the species or in the sections to which they
were assigned.
The remaining species of Draba indigenous to the Southeastern States are
very closely related. They were placed by Schulz (1927) in sect. TOMOSTIMA
(Raf.) O. E. Schulz, which also included the South American D. araboides
Wedd. and D. australis R. Br. All are subscapose annuals with subsessile basal
leaves, obsolete styles, and heteromorphic flowers (some with broad, white
petals, others apetalous and cleistogamous). Draba reptans (Lam.) Fern. (Arabis
reptans Lam., D. caroliniana Walter, D. micrantha Nutt., D. coloradensis Rydb.,
D. reptans var. stellifera (O. E. Schulz) C. L. Hitche.; see Hitchcock and Fernald
(1934) for 15 additional synonyms), 2” = 16, 30, 32, grows in open sandy
areas, rock crevices, pastures, prairies, and disturbed sites, as well as along
roadsides and railroad tracks. It is distributed from Massachusetts southward
into North Carolina (Lincoln County), South Carolina (Darlington County),
Georgia (Kenesau Mtn.), Tennessee (Nashville Basin), Alabama (Lee and
Montgomery counties), Arkansas (Washington and Sebastian counties), and
westward into the Pacific States, as well as in Manitoba, Ontario, and Sas-
katchewan. Draba reptans is easily distinguished from its nearest relatives by
its entire or subentire leaves with simple or sometimes forked trichomes on
the upper surface and stellate ones on the lower, and by its subumbellate
infructescences with glabrous rachises and pedicels. Smith (1965) reported 27 =
16 from plants of Kansas, but Léve & Léve found tetraploid populations
(2n = 32) in Manitoba, and Mulligan (1966) counted ” = 15 in plants from
Saskatchewan and South Dakota.
Draba cuneifolia Nutt. ex Torrey & Gray is a variable and widely distributed
species in which Hartman and colleagues recognized three varieties. Variety
cuneifolia (D. Helleri Small, D. ammophila Heller, D. cuneifolia var. leiocarpa
O. E. Schulz, D. cuneifolia var. Helleri (Small) O. E. Schulz, D. cuneifolia var.
foliosa Mohlenbrock & Voigt), 2” = 32, 1s widely distributed in northern and
southern Arkansas, southeastern Kansas, Missouri, Oklahoma, the South-
western States, central and western Colorado, western Utah, southern Nevada,
and adjacent southeastern California. It is sporadic and probably introduced
in North Carolina (New Hanover County), Florida (Duval, St. Johns, and
Jackson counties), Alabama (Sumter County), Tennessee (Decatur County),
Mississippi (Oktebbeha County), Louisiana (Grant, Rapides, and Caddo par-
ishes), and Ohio. It is indigenous but apparently uncommon in Chihuahua,
Coahuila, Baja California, and Zacatecas, Mexico. Mohr stated that D. cunei-
folia is found in Georgia, but I have not seen any specimens from this state,
and Hartman and colleagues did not list it from there. The species grows on
limestone ledges, rocky slopes, and disturbed sandy soils in prairie pastures,
1987] AL-SHEHBAZ, ALYSSEAE 215
lawns, grassy plains, fallow fields, cedar glades, and waste places. The other
varieties of D. cuneifolia, var. integrifolia S. Watson and var. sonorae (Greene)
S. B. Parish, do not occur in our area and are primarily distributed in the
southern parts of California, Nevada, Arizona, and Texas and in adjacent
northern Mexico. They differ from var. cuneifolia in their fruits with stellate
instead of simple trichomes. Variety cuneifolia sometimes has glabrous fruits.
Draba platycarpa Torrey & Gray (D. cuneifolia var. platycarpa (Torrey &
Gray) S. Watson, D. viperensis St. John), 2n = ca. 16, 32, differs from D.
cuneifolia in its obovate to broadly elliptic, rounded fruits 2.5-3.7 mm wide
and in its scapes pubescent with a mixture of long, simple trichomes and short,
branched ones. The latter species has oblong to lanceolate or narrowly elliptic,
acute fruits 1.8—2.8 mm wide and scapes with short, branched trichomes only.
Draba platycarpa is sporadic in Louisiana (Lincoln Parish), Arkansas (Hemp-
stead and Garland counties), and Oklahoma but is widespread in Texas and
central and southern Arizona. It is disjunct and probably introduced in Idaho,
Oregon, and Washington. Several authors (e.g., Watson, Hitchcock) reduced
D. platycarpa to a variety of D. cuneifolia, but Hartman and colleagues have
clearly shown that they should be treated as closely related species. They are
morphologically distinct, and their profiles of flavonoid glucosides and volatile
components are very different. They do not hybridize in areas of sympatry,
and despite the numerous attempts to make artificial crosses between the two
species, no hybrids were obtained (Hartman et a/.). Diploid and tetraploid
populations of D. platycarpa were found in Texas in Tarrant (Hartman et al.)
and Kinney (Rollins & Riidenberg) counties, respectively.
Draba verna L. (Erophila verna (L.) Chev., £. vulgaris DC.; see Schulz (1927)
for more than 200 additional synonyms listed as species, varieties, or forms),
whitlow grass, whitlow wort, 27 = 14, 16, 24, 30, 32, 34, 36, 38, 40, 52, 54,
58, 60, 64, is a Eurasian plant naturalized throughout the New World. It has
been reported from all of the Southeastern States except Louisiana and Florida.
It is one of the earliest annuals to bloom in late winter and early spring (the
generic name Erophila, under which D. verna is often placed, is derived from
Greek er, spring, and phileo, to love, referring to its early appearance in spring).
The species grows in lawns, fields, waste places, pastures, cedar glades, and
open rangeland, on grassy hillsides, and along roadsides. It was well established
in North America as early as the first half of the eighteenth century (Benson).
Draba verna isa highly variable and taxonomically difficult complex in which
numerous extremes have been recognized as species, subspecies, or varieties.
It consists of self-pollinating, morphologically distinct, uniform, local popu-
lations with different chromosome numbers. Crosses between such populations
often produce hybrids that are sterile because of meiotic abnormalities. Autog-
amy played a major role in the formation and stabilization of a very large
number of easily separable populations. Nearly 200 such populations were
recognized by the nineteenth-century French botanist Alex Jordan as distinct
“species,” sometimes called “Jordanons.” Schulz (1927) reduced these to eight
species and some 60 varieties, but subsequent workers (e.g., Winge, 1940)
questioned the taxonomic status of most of them. There is no correlation
between the morphological, cytological, genetic, geographic, and ecological data
216 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
on this complex, which is best recognized as a single polymorphic species with
several subspecies. No attempt is made here to determine the subspecies of D.
verna naturalized in the Southeastern United States.
Draba is a well-defined genus easily recognized by its latiseptate (flattened
parallel to the septum), ovate to orbicular or oblong to linear fruits, usually
wingless seeds biseriately arranged in each locule, unappendaged staminal fil-
aments, accumbent cotyledons, and usually branched trichomes. The limits of
the genus have not been altered during the past two centuries, and only one of
its segregates is controversial. Erophila, which is united with Draba by North
American botanists and retained as an independent genus by those elsewhere,
differs from Draba only in its bifid instead of entire to deeply emarginate petals.
The two are indistinguishable in every other morphological character. In my
opinion this difference is not important; certain genera of the Cruciferae (e.g.,
Megacarpaea DC. and Alyssoides) have species with either entire or bifid petals.
Petal apex (bifid vs. entire) may be controlled by a few genes or by a single
pleiotropic gene and could therefore be insignificant for generic delimitations
within the Cruciferae.
Perhaps the major taxonomic complexity in Draba, other than its sectional
classification, lies in its species limits. Rollins (1966) suggested that apom1xis
together with polyploidy and interspecific hybridization are responsible for this
complexity. Many species have been described on the basis of minor differences
in characters of which the variation was poorly understood. For example,
presence vs. absence of trichomes on fruits 1s insignificant in certain complexes,
and numerous species (e.g., D. reptans and D. cuneifolia) have plants with
either glabrous or pubescent fruits within the same population. On the other
hand, the type of trichome (simple, furcate, cruciform, stellate, or dendritic) is
very important in separating species.
Self-compatibility is apparently very common in Draba, and only a few
species are self-incompatible (Bateman; Mulligan, 1976; Mulligan & Findlay).
Protogyny occurs in a few species such as D. aizoides L. and D. alpina L. (AI-
Shehbaz, 1977; Kay & Harrison), while autogamy is widespread in the genus.
Species such as D. reptans, D. cuneifolia, and D. aprica produce heteromorphic
flowers: some have sizeable petals, others have reduced ones, and still others
are apetalous and cleistogamous (Fernald, 1934). They apparently produce
apetalous flowers toward the end of the growing season (Kral), but D. tenerrima
O. E. Schulz (Kashmir, Pakistan) is always apetalous and has only four stamens.
Agamospermy occurs in the North American D. densifolia Nutt. ex Torrey &
Gray, D. Paysonti Macbr., D. ventosa A. Gray, D. exunguiculata (O. E. Schulz)
C. L. Hitche., D. Grayana (Rydb.) C. L. Hitche., D. oligosperma W. J. Hooker,
and D. streptobrachia R. A. Price (Mulligan, 1976; Mulligan & Findlay; Price,
1979, 1980). In these species pollen fertility is zero or nearly so and the anthers
do not dehisce. Some apomicts are triploids with highly irregular meiosis, but
all produce abundant viable seeds without the need for pollen stimulation of
seed production. Earlier claims of apomixis in D. verna (see Lotsy) were based
on misinterpreted observations.
Chromosome numbers are known for some 115 species, the majority of
which (nearly 60 percent) are polyploid; only a few (about 5 percent) have
1987] AL-SHEHBAZ, ALYSSEAE 217
diploid and polyploid populations. Although base chromosome numbers in
Draba range from six to 12, those of nearly 85 percent of the species are based
on eight. Mulligan (1966) suggested that the North American species probably
evolved through aneuploidy at the polyploid level. The lowest chromosome
number in the genus (2 = 12) is found in D. Olgae Regel & Schmalh. (central
Asia), while the highest counts (27 = 128, 144) are found in the North American
D. corymbosa R. Br. ex DC. (including D. macrocarpa J. M. F. Adams and D.
Bellii T. Holm), which consists of 16- and 18-ploid populations based on x =
8 (Bécher, 1966; Mulligan, 1974a). As shown above, D. verna is the most
cytologically complex species in the genus. It contains many chromosomal
races ranging from diploid to octoploid, as well as intermediate aneuploid
derivatives.
Boécher (1966) indicated that the majority of the alpine species are diploid
while the arctic ones are polyploid. He speculated that the mountains south of
the arctic areas are probably the centers of origin and that Draba may be
polyphyletic. He observed that polyvalent formations are very rare in drabas
with high ploidy levels and suggested that allopolyploidy may have played an
important evolutionary role at the hexaploid and decaploid levels.
Despite claims by many authors (e.g., Ekman (1932b), Schulz (1927), Wein-
gerl) that interspecific hybridization is widespread in Draba, very little exper-
imental work supports this. Fernald (1934) suggested that most of the alleged
interspecific hybrids represent variations within poorly circumscribed, poly-
morphic species, while Knaben seriously questioned the validity of several
hybrids listed by Ekman (1932b). There are strong sterility barriers between
pairs of many closely related species. Mulligan (1974b, 1975, 1976) showed
that artificial hybridization between many sexual species produces offspring
with zero or very low pollen fertility and with aborted fruits. He concluded
that interspecific hybridization is very rare in nature. Some members of the
D. nivalis Liljeblad group produce sterile natural interspecific hybrids. Viable
seeds were obtained from a few successful artificial crosses, but the second-
generation hybrids did not reach maturity (Mulligan, 1975).
The chemistry of Draba is poorly studied, and only a few species have been
surveyed for secondary constituents. Isopropyl, 2-butyl, allyl, 3-butenyl, and
benzyl glucosinolates are found in four unrelated species (Kjaer; Rodman &
Chew; Hartman et a/.). The fatty-acid content of only six species has been
determined (Jart).
Roots of certain rock-dwelling, perennial species of Draba have peculiar
secondary growth characterized by the formation of armed periderm, abundant
soft tissue, and secondary xylem structurally resembling the primary. These
anatomical specializations, which are believed to be adaptations to rocky hab-
itats, are also found in genera outside the Cruciferae (Pirogov).
Many species of Draba are cultivated as rock-garden or wall plants (Irving).
Very few are weeds or show weedy tendencies. The fruiting stalks and seeds
of D. nemorosa L. are used in China and Japan as diuretics and are prescribed
to treat coughs, dropsy, nausea, and pleurisy (Perry; Kung & Huang). Draba
verna (whitlow grass) was believed to cure whitlow, inflammation around the
nails.
218 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
REFERENCES:
Under family references in AL-SHEHBAZ (Jour. Arnold Arb. 65: 343-373. 1984), see
AL-SHEHBAZ (1977); BATEMAN (1955a); BENTHAM & HOOKER; BRITTON & BROWN; DE
CANDOLLE (1821, ae FERNALD; VON HAYEK; Beau & RECHINGER; JAFRI; JART;
JONES; ie (1960); KNUTH; MAIRE; MARKGRAF; PERRY; RADFORD et al.; RICKETT;
RODMAN & CHEW; ROLLins (1966, 1981); SCHULZ; oe E. B. SmitH; and VAUGHAN
& Wurrenouse.
Under tribal references see BAILEY; BOLKHOVSKIKH ef al/.; DUNCAN & en
GATTINGER; GOLDBLATT (1981, 1984, 1985); MACRoBERTS; Moore; and SMAL
Arwipsson, T. Notizen iiber Arten der Gattungen Draba, Erophila und Hutchinsia.
Bot. Not. 1929: 169-174. 1929.
AveTISIAN, V. E. Role of high mountain areas of the Caucasus in speciation of Draba
L. and some aspects of the genesis of this genus. (In Russian.) Probl. Bot. 14(1): 59-
62. 1979. [Sections, chromosome numbers. ]
BaLpaccl, A. Monografia della sezione “Aizopsis DC.” del genere Draba L. Nuovo
Giorn. Bot. Ital. If. 1: 103-121. 1894. [Treatment of 12 species.]
Baskin, J. M., & C. C. BAskin. Germination eco-physiology of Draba verna. Bull. Torrey
Bot. Club 97: 209-216. 1970. [Light, temperature, moisture. ]
& . The light factor in the germination ecology of Draba verna. Am. Jour.
Bot. 59: 756-759. 1972.
Effect of relative humidity on afterripening and viability in seeds of
the winter annual Draba verna. Bot. Gaz. 140: 284-287. 1979.
BENSON, A. B., ed. Peter Kalm’s travels in North America. Vol. 1. xvii + 380 pp. New
York. 1937. [D. verna was abundant in 1749 near Philadelphia, 257.]
BERKUTENKO, A. Notulae systematicae de genere Draba L. in parte boreali-orientale
URSS. (In Russian.) Novit. Syst. Pl. Vasc. 16: 119-125. 1979. [Four North American
and eastern Asiatic species
BocHER, T. W. Experimental and cytological studies on plant species. [X. Some arctic
and montane crucifers. Biol. Skr. Dan. Vid. Selsk. 14(7): 1-74. pls. I-10. 1966.
[Draba, 5-39, 63-67, figs. 1-10, tables I-6, pls. 1-8; chromosome numbers, distri-
butions, relationships, hybridization, speciation, polyploidy.]
. Variation and distribution pattern in Draba sibirioa (Pall.) Thell. Bot. Not. 127:
317-327. 1974. [Chromosome numbers, taxonomy, leaf anatomy, ecology, a new
subspecies. ]
BuTTLer, K. P. Zytotaxonomische Untersuchungen an mittel- und siideuropdischen
Draba-Arten. Mitt. Bot. Staatssam. Miinchen 6: 275-362. nae [Comparative mor-
phology, — and distributions of 16 species, key, maps.]
EKMAN, E. Nomenclature of some North-European Drabae. Atk. Bot. II. 12(7): 1-17.
pl. I. 1912.
. Zur Kenntnis der nordischen Hochgebirgs-Drabae [I]. Kungl. Sv. Vetensk. Handl.
57(3): 1-68. pls. I-3. 1917. [Part II in ibid. ser. 3. 2(7): 1-56. maps 1-5, pls. 1-3.
26.]
—. Studies in the genus Draba. Sv. Bot. Tidsk. 23: 476-495. 1929. [Interspecific
hybridization; contribution to the Draba flora of Greenland I.]
Contribution to the Draba flora of Greenland. II. /bid. 24: 280-297. pi. 3. 1930.
art II in ibid. 25: 465-494. pi. 5. 1931; part IV in ibid. 26: 431-447. 1932a; part
in ibid. 27: 97-103. 1933; part VI in ibid. 339-346; part VII in ibid. 28: 66-83.
1934; part VIII in hid. 29: 348-364. 1935.]
. Some notes on the hybridization in the genus Draba. Ibid. 26: 198-200. 1932b.
—. Notes on the genus Draba. A posthumous, unfinished fragment. /bid. 35: 133-
141. 1941. [Key to the species of Greenland; infraspecific taxonomy of D. rupestris.]
ELveN, R., & A. AARHuS. A study of Draba cacuminum (Brassicaceae). Nordic Jour.
Bot. 4: 425-441. 1984. [Numerical analysis, infraspecific taxa, ecology, distribution. ]
<3
1987] AL-SHEHBAZ, ALYSSEAE Z19
FERNALD, M.L. Draba in temperate northeastern America. Rhodora 36: 241-261, 285-
305, 314-344, 353-371, 392-404. pls. 290-310. 1934. (Reprinted in Contr. Gray
Herb. 105. 1934.) [Treatment of 25 species, descriptions, distributions, key, maps
1-24.]
& H. KNow.ton. Draha incana and its allies in northeastern America.
Rhodora 7: 61-67. pl. 60. 1905.
Gita, E. Uber die ee und die ee der amerikanischen
Arten der Gattung Draba. Bot. Jahrb. 40(Beibl. 90, 1): 35-46. 1907.
GRIESINGER, R. Zytologische tad experimentelle Uniesucnaeecn an Erophila verna.
Flora 129: 363-379. pls. 1, 2. 1935. [Chromosome numbers, micro- and megaspo-
rogenesis, crosses. ]
HarTMAN, R. L., J. D. BAcon, & C. F. BoHNSTEDT. Biosystematics of Draba cuneifolia
and D. platycarpa (Cruciferae) with emphasis on volatile and flavonoid constituents.
Brittonia 27: 317-327. 1975. [Distributions, descriptions, eee numbers,
key, flavonoid profiles, nitrites, ea te sesquiterpen
Hence, I. C. The status of Thylacodraba O. E. Schulz. Notes Bot, rere Edinburgh 23:
173, 174. pl. 13. 1960. [Reduced to synonymy of Draba.]
HErLBorn, O. Chromosome numbers in Draba. Hereditas 9: 59-68. 1927.
Some chromosome numbers in Draba. Sv. Bot. Tidsk. 35: 141, 142. 1941.
HircHcock, C.L. A revision of the drabas of western North America. Uae Washington
Publ. Biol. 11; 3-132. table 1, pls. 1-8. 1941. [Treatment of 64 species, tabulation
of 21 characters for all species, illustrations of fruits and leaves; D. acaba D.
cuneifolia, D. platycarpa (as a variety), D. reptans, D. verna.
Hooker, W. J. Draba dentata. Hooker’s Ic. Pl. 1: pi. 31. 1837. [D. ramosissima.]
Des W.] The whitlow grasses. Garden (London) 87: 657-659. 1923.
J. sg Biological flora of the British Isles. Draba aizoides L.
ae Ecol. 58: 877-888. 1970.
KNABEN, G. Cytological oe in some Draba species. Bot. Not. 119: 427-444. pl. J.
6. [D. fladnizensis, D. lactea, D. nivalis, D. norvegica; chromosome numbers,
ee variation, hybridization.]
KRAL, R. Are on some rare, threatened, or endangered forest-related vascular plants
of the Sante oy S. Dep. Agr. Forest Serv. South Reg. Tech. Publ. R8-TP2. Vol. 1.
x + 718 pp. 1983. [D. aprica, 476-479, description, toe map. |
Kuna, H. P., & W.-Y. HuanG. Chemical investigation of Draba lee L. The
isolation eee nauine iodide. Jour. Am. Chem. Soc. 71: 1836, 1837.
LeseGue, A. Le développement de l’embryon et la eas del’ eee chez
le Draba verna L. Bull. Soc. Bot. France 97: 103-1 0
Licuvar, R. W. Evaluation of Draba oligosperma, D. Bi her and D. juniperina
complex (Cruciferae). Great Basin Nat. 43: 441-444. 1983. [Comparative mor-
phology, SEM of leaf trichomes, key; see ROLLINs, 1953.]
Locan, L. A. A list of seed plants of Lincoln Parish, Poieana: Proc. Louisiana Acad.
Sci. 26: 18-32. 1963. [D. brachycarpa, 24.]
Lotsy, P. Has Winge proved that Erophila is not apogamous? Genetica 8: 335-344.
1926. [No; crosses within D. verna complex.
LOVE, ns & D. Love. In; A. Léve, ed., IOPB chromosome number reports LXXIV.
Taxon 31: 119- 128. 1982. [Draba, 125, 126; counts for 11 species.]
MERXMULLER, H., & K. P. BUTTLER. Die Chromosomenzahlen der a eee ica
aie os Draben. Ber. Deutsch. Bot. Ges. 77: 411-415. 1965. [Counts for 20
species. }
Monr, c. Plant life of Alabama. xii + 921 pp. aca Alabama. 1901. (Reprinted
from Contr. U. S. Natl. Herb. 6.) [Draba, 527.]
MUuLLIGAN, G. A. Chromosome numbers of family Cruciferae. IJ. Canad. Jour.
Bot. 44: 309-319. 1966. [Thirteen species of Draba, evaluation of Erophila.]
ytotaxonomic studies of Draba glabella and its close allies in Canada and
Alaska. Ibid. 48: 1431-1437. pls. 1, 2. 1970. [D. glabella, D. arabisans, D. borealis,
220 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
D. longipes, D. incana, D. fabian distributions, key, pollen of 20 species, SEMs
of leaf trichomes, chromosome n
. Cytotaxonomic studies of hen closely allied Draba cana, D. cinerea, and D.
groenlandica in Canada and Alaska. Ibid. 49: 89-93. p/. 1. 1971a. [Chromosome
numbers, Saeeeen key, map, SEMs of trichomes. ]
nomic studies of Draba species in Canada and Alaska: D. ventosa, D.
ruaxes, aD Paysonii. [bid. 49: 1455-1460. pl. 1. 1971b. [Chromosome numbers,
taxonomy, distributions, reproductive biology, key, maps, SEMs of trichomes.]
Cytotaxonomic studies of Draba ae : Canada and Alaska: D. oligosperma
and D. incerta. Ibid. 50: 1763-1766. pl. J
Confusion in the names of three cr et species of the arctic: D. Adamsii, D.
oblongata, and D. corymbosa. Ibid. 52: 791-793. pls. I, 2. 1974a. [Synonymy, SEMs
of trichomes, photos of types
Cytotaxonomic studies of Draba nivalis and its close allies in Canada and Alaska.
Ibid. 52: 1793-1801. 1974b. [Taxonomy, distributions, chromosome numbers, nat-
ural and artificial hybridization, key, figs. /-30; D. Porsildii, sp. nov.
. Draba crassifolia, D. Albertina, D. nemorosa, and D. stenoloba in Canada a
Alaska. [bid. 53: 745-751. 1975. [Chromosome numbers, taxonomy, re or ae
artificial interspecific Ae sa ]
The genus Draba in Canada and Alaska: key and summary. [bid. 54: 1386-
1393. 1976. [Forty sere chromosome numbers, breeding systems, distributions,
interspecific hybridization, SEMs of trichomes.]
Four new species of Draba in northwestern North faa Ibid. 57: 1873-
1875. 1979. [D. Hatchiae, D. Murrayi, D. kluanei, D. Sco
. N. Finpiay. Sexual reproduction and a in the genus Draba.
Canad. Jour. Bot. 48: 269, 270. p/. 7. 1970. [Agamospermy in D. oligosperma, self-
incompatibility in D. Helleriana, self-compatibility in 15 species.]
Nye, T. G. An ecological study of Draba ramosissima Desv. with notes on the intra-
specific taxonomy and leaf morphology of the species. vi + 28 pp. Unpubl. M.S.
dissert asus Univ. Kentucky, Lexington. 19
on the intraspecific taxonomy and leaf morphology of Draba ramosissima
Desv. eae 34; 210-217. wae ili variation in leaf morphology
— four populations from Kent
: ecological study of the ae Draba ramosissima Desv. Ibid. 409-413.
1969b.,
Payson, E. B. The perennial scapose drabas of North America. Am. Jour. Bot. 4: 253-
267. 1917. [Distributions of 26 species, key, 14 new species.
. ST. Joun. The Washington species of Draba. Proc. Biol. Soc. Washington
43: 97-122 1930. [Fifteen species, descriptions, distributions, key, evaluation of
PiroGcov, V. S. The specialization of the roots of lithophytes belonging to the genus
Draba and of some nee rock-dwelling plants. (In Russian; English summary.) Bot.
Zhur. 53: 350-357.
PoHLe, R. Drabae ee ee 'Systematik und Geographie nord- und Seer aeate
ng Repert. Sp. Nov. Beih. 32: 1-225. 1925. [Fifty-five species in nine group
key
PRICE, R. ‘A. The Draba crassa complex loa aaa ever and geography. ili +
88 pp. Unpubl. M.S. dissertation, Univ. W sin, Madison. 1979.
. Draba streptobrachia (Brassicaceae), a new eee from Caleealo Brittonia 32:
160-169. 1980. [Numerical analysis, chromosome numbers, agamospermy, com-
parison with D. spectabilis, SEMs of trichomes, map.]
RATCLIFFE, D. Biological flora of the British Isles. Draba muralis L. Jour. Ecol. 48: 737-
744. 1960.
Reep, C. F. Contribution to the flora of Maryland, 3. Draba ramosissima in Maryland,
1987] AL-SHEHBAZ, ALYSSEAE 221
with notes on the general distribution. Castanea 22: 113-119. 1957. [Variation in
leaf shape and pubescence; map
Roiums, R. C. Draba on Clay Butte, Wyoming. Rhodora 55: 229-235. 1953. [Notes
on |1 species, new taxa, chromosome numbers
raba aprica in Oklahoma. Ibid. 63: 223- 225, 1961. [New state record; differ-
ences tenet D. aprica and D. brachycarpa.|
Species of Draba, Lesquerella, and Sibara (Cruciferae). Contr. Gray Herb. 211:
107- 113. 1982. [Notes on species of Draba with winged seeds; D. carnosula, sp.
nov.]
. Draba (Cruciferae) in Mexico and Guatemala. /bid. 213: 1-10. 1984. [Eleven
species and six varieties, key, distributions, four new taxa.
& ENBERG. Chromosome numbers of Cruciferae II]. Contr. Gray Herb.
201: 117- 133. 1971. [D. Cie D. cuneifolia, D. platycarpa, 122.]
SALMON, C. E., & E. G. BAKER. Notes on the British species and forms of Erophila.
Jour. Bot. ieondon 66: 234-241. 1928.
Scuutz, O. E. Cruciferae—Draha et Erophila. In: A. ENGLER, Pflanzenr. IV. 105(Heft
89): 1-396. 1927. oe monograph; 258 species of Draba in 17 sections,
eight species of Erop
Uber Mee ice Bitten der Erophila verna (L.) E. Meyer. Verh. Bot. Ver.
Brandenb. 72: 76. 1930.
SHARSMITH, C. W. Notes on Draba in the Sierra Nevada. Madrofio 5: 147-151. 1940.
Smitu, E. B. Chromosome numbers of Kansas flowering plants. I. Trans. Kansas Acad.
Sci. 67: 818, ee ae [D. reptans, 2n = 16.]
—. In: A. Lév , IOPB orene ene number reports XXIV. Taxon 18: 683,
684. 1969. [D. mae 684, n J
STEYERMARK, J. A. Draba aprica in the Onis of southeastern Missouri. Rhodora 42:
33
Sutter, R. D., L. MANSBERG, & J. Moore. Endangered, threatened, and rare plant
species of North Carolina. ASB Bull. 30: 153-163. 1983. [D. ramosissima, D. rep-
tans, 159.]
Totmacuev, A. I. Draba. In: V. L. Komarov & N. A. Buscu, eds., Fl. USSR 8: 371-
454, 649, 650. 1939. ee translation by R. Lavoortr, 8: Gs 337, 486, 487.
1970. [Ninety-one species in 29 series, keys.]
. Acontribution to the history and geographical distribution of the genus Draba
L. (In Russian.) Bot. Zhur. 42: 1446-1456. 1957.
Watters, S. M. Draba and Erophila. In: T. G. TuTin et al., eds., Fl. Europaea 1: 307-
313. 1964. [Forty-two species of Draba in four sections; two species of Erophila.]
Watson, S. Contributions to American botany. 1. Some new species of plants of the
United States, with revisions of Lesquerella (Vesicaria) and of the North American
species of Draba. Proc. Am. Acad. Arts Sci. 23: 249-267. 1888. [Five sections and
32 species; D. brachycarpa, D. cuneifolia, D. platycarpa, D. ramosissima, D. reptans
(as D. caroliniana), D. verna.
WEINGERL, H. Beitraége zu einer Monographie der europdisch-asiatischen Arten aus der
Gattung Draba, sect. Leucodraba. Bot. Arch. 4: 9-109. 1923. [Nineteen species in
four series, extensive synonymy, distributions, hybridization, keys.
WincE, O. Das Problem der Jordan-Rosen’schen Erophila-Kleinarten. Beitr. Biol. Pflan-
zen 14: 313-334. pi. 6. 1926. [Chromosome numbers, crosses between four mor-
phological extremes of the D. verna complex.]
A case of amphidiploidy within the collective species Erophila verna. Hereditas
18: 181-191. 1933. [Stable hybrids with = 47 were obtained from crossing two
ees with n = 15 and n = 32.]
: onomic and evolutionary studies in Erophila based on cytogenetic inves-
tigations Compt. Rend. Lab. Carlsb. Physiol. 23: 41-74. figs. 1-92. 1940. [Taxo-
nomic history, chromosome numbers, hybridization, stabilization of hybrids; four
PP, JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
species recognized; eee series of photographs exhibiting the continuous vari-
ation in rosettes fru
ZHUKOVA, P. G., & V ee A cytotaxonomical study of some species of the
family Brassicaceae in northern Asia. (In Russian.) Bot. Zhur. 69: 236-240. 1984.
[Counts for 23 species of Draba
25. Lesquerella S. Watson, Proc. Am. Acad. Arts Sci. 23: 249. 1888.
Annual, biennial, or perennial herbs, usually densely pubescent with stellate
[or lepidote] trichomes, sometimes pubescent with a mixture of simple, bifur-
cate, and dendritic ones. Stellate trichomes with few [to numerous] rays; rays
smooth or tuberculate, simple or once [or twice] forked; webbing absent [or
present only between the bases of rays, or progressively developed to their
tips]. Stems decumbent to erect, several to numerous or rarely solitary, usually
arising laterally from the basal rosette. Basal leaves petiolate, entire or dentate
to sinuate or lyrately to pinnately lobed. Cauline leaves petiolate or sessile,
usually cuneate at base, sometimes auriculate or amplexicaul, entire to dentate
[rarely sinuate or incised]. Inflorescences ebracteate, few- to many-flowered,
corymbose racemes; infructescences lax [or congested]; fruiting pedicels per-
sistent, ascending to horizontal [or reflexed], straight or curved [sometimes
sigmoid]. Sepals pubescent, narrowly oblong or elliptic to broadly ovate [oc-
casionally linear or obovate], ascending to spreading [or erect], almost always
ng
purple veins], glabrous, broadly obovate [to narrowly spatulate], undifferen-
tiated or slightly differentiated into claw and blade, obtuse or retuse to emar-
ginate at apex. Nectar glands usually forming a ring [or a hexagon] subtending
the bases of median filaments and surrounding those of lateral stamens. Sta-
mens 6, tetradynamous; filaments linear, unappendaged, not dilated or some-
times strongly dilated at base; anthers linear [to oblong or ovate], usually
sagittate at base. Fruits globose to subdidymous, sometimes obovoid to sub-
pyriform [or ovoid to oblong], inflated or rarely strongly flattened parallel [or
at right angles] to the septum, sessile or stipitate; valves glabrous or variously
pubescent on the exterior or on both outer and inner surfaces, obscurely or
rarely strongly nerved, thick or sometimes papery or membranaceous, rounded
[or rarely strongly keeled] on the back; replum glabrous or pubescent; septum
complete or occasionally with a central perforation, rarely reduced to a narrow
band around the inner margin of replum, usually with a conspicuous nerve
extending from the base of style to about or slightly beyond the middle, trans-
lucent or opaque; styles slender, persistent, glabrous or pubescent; stigmas
capitate, entire or slightly 2-lobed, often much greater in diameter than the tip
of style; ovules 2—14[-20] per locule: base of funicles usually adnate to septum.
Seeds reticulate, orbicular or suborbicular, rarely hemispherical [or oblong to
oval], flattened or rarely plump, with or without a narrow margin or wing,
nonmucilaginous [or copiously mucilaginous] when wet; cotyledons accumbent
[or rarely obliquely accumbent], longer [or equaling to shorter] than the radicle.
Base chromosome numbers 5-10. Lecrotype species: L. occidentalis (S. Wat-
1987] AL-SHEHBAZ, ALYSSEAE 223
son) S. Watson; see Payson, Ann. Missouri Bot. Gard. 8: 133. 1922. The
arbitrary designation of L. Lescurii (A. Gray) S. Watson as the lectotype species
of the genus by Britton & Brown should be rejected because it is in conflict
with Watson’s original description of Lesquerella. For further discussion on
the subject, see Payson and Rollins & Shaw (1973). (Name honoring Charles
Leo Lesquereux, Nov. 18, 1806-Oct. 25, 1889, a distinguished Swiss-born,
American paleobotanist and bryologist.)— BLADDERPOD.
A well-defined genus of some 90 species, the majority of which (83 species
and 27 infraspecific taxa) occur in North America, particularly in the south-
western United States and adjacent Mexico, the Rocky Mountains, and the
intermontane basin of the western United States. The remainder (probably up
to 12 species; Rollins & Shaw, 1973) are found in South America from Bolivia
southward. One species, Lesquerella arctica (Wormsk. ex Hornem.) S. Watson,
is widely distributed from the coasts of Greenland across the Canadian Arctic
and Alaska into Siberia. The genus is represented in the southeastern United
States by seven species, of which five are endemic
The sectional classification of Lesquerella, as proposed by Watson and Pay-
son, does not reflect the natural groupings of species. The former recognized
two sections: sect. ALysmMus S. Watson (five species; plants not canescent,
filaments dilated at base, cauline leaves usually auriculate) and sect. LESQUEREL-
LA (28 species; plants canescent, filaments slender at base, cauline leaves not
auriculate). Payson, on the other hand, redefined sect. ALysmus to include only
one species, L. Lescurii, with latiseptate fruits (flattened parallel to the septum).
e placed in sect. ENANTIOCARPA Payson three species said to have angusti-
septate fruits (flattened at right angles to the septum) and retained in sect.
ESQUERELLA (as sect. Eulesquerella) the remaining 48 species, with inflated,
globose or ovoid fruits. On the basis of chromosome numbers, fatty-acid con-
tent, cross-fertility, and several morphological features, L. Lescurii 1s very
closely related to several species with globose fruits. Therefore, sect. ALYSMUS
is clearly artificial. One of the three species assigned by Payson to sect. ENANTIO-
CARPA is a Draba, while the other two are definitely unrelated (Rollins & Shaw,
1973). Slightly angustiseptate fruits probably evolved independently a few times
within Lesquerella, and alone they can be unreliable indicators of relationships.
As indicated by Rollins & Shaw (1973), the sectional classification of Les-
querella was not based on well-founded facts. It is impractical to place a few
species in one or two sections and to retain the bulk of a genus in a highly
heterogeneous 0
Lesquerella ae (W. J. Hooker) S. Watson (Vesicaria gracilis W. J. Hook-
er, Alyssum gracile (W. J. Hooker) Kuntze, V. polyantha Schlecht., L. polyan-
tha (Schlecht.) Small), cloth-of-gold, 2” = 12, is represented in the Southeastern
States by subsp. gracilis. It grows on sandy loam or alkaline soil in prairies,
pastures, and old fields, as well as along roadsides and grassy banks, in Arkansas
(Howard and Little River counties), eastern Mississippi (Chickasaw, Lee, and
Lowndes counties), southern Oklahoma, and east-central Texas. It is weedy
and has been introduced in Tennessee in Shelby and Davidson counties (Rogers
& Bowers) and in Missouri and Illinois. The subspecies 1s distinguished by its
224 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
stipitate, glabrous, globose or ellipsoid fruits 3-6 mm long; cuneate, sessile or
short-petiolate cauline leaves; 4-10(-14) ovules per locule; stellate trichomes
with 4—7 bilaterally oriented rays; and straight, usually divaricate fruiting ped-
icels.
The records of Lesquerella gracilis subsp. Nuttallii (Torrey & Gray) Rollins
& E. Shaw from Arkansas by Small (1913) (as L. Nuttallii Torrey & Gray and
L. repanda (Nutt.) S. Watson) and by Payson (as L. gracilis var. repanda (Nutt.)
Payson) were shown by Rollins & Shaw (1973) to be based on plants from
Texas and Oklahoma, respectively. Subspecies Nuttallii differs from subsp.
gracilis in its obpyriform to narrowly obovoid fruits (4.5-)5.5-9 mm long with
a truncate base, instead of globose or ellipsoid fruits 3-6 mm long with a
rounded base. Small (1913, p. 471) also indicated that L. angustifolia (Nutt.)
S. Watson occurs “‘on prairies, near the Red River, Arkansas,” but the record
was from Red River County, Texas.
Lesquerella globosa (Desv.) S. Watson (Vesicaria globosa Desv., V. Shortii
Torrey & Gray, Alyssum globosum (Desv.) Kuntze, A. Shortii (Torrey & Gray)
Kuntze), 27 = 14, has no close relatives in the genus and is clearly unrelated
to any of the six species occurring in the Southeast. It is distributed in central
Tennessee (Maury, Davidson, Cheatham, and Montgomery counties), north-
central Kentucky, and Indiana (Posey County). The species was said to occur
in Benton and Franklin counties, Arkansas (Smith), but I have not seen any
material from this state, and neither Rollins & Shaw (1973) nor Kral has
indicated that it is found there. It is most common on open rocky areas,
limestone ledges, and cliffs along rivers but also grows in cedar glades and
pastures and on open talus slopes. Lesquerella globosa has numerous small,
globose, pubescent fruits (1-)2-2.8 mm long with a conspicuously wrinkled
septum (FIGURE 3b); usually one subhemispherical seed per locule; straight
fruiting pedicels; sessile or short-petiolate cauline leaves; and stellate trichomes
with three to six usually forked rays.
The five remaining species of Lesquere/la are endemic to the Southeastern
States. All are annuals with a mixture of simple and branched (but never stellate)
trichomes, auriculate cauline leaves, and staminal filaments with strongly di-
lated bases. They are diploids (2n = 16) that produce fully fertile offspring
when hybridized (see below) in any combination. Furthermore, they contain
high concentrations of densipolic acid, a unique seed fatty acid. The morpho-
logical, geographic, cytological, chemical, and interfertility data clearly support
the derivation of the five species from a common ancestor.
Lesquerella lyrata Rollins, 2n = 16, is a narrow endemic that grows in open
pastures, old fields, cedar glades, and bottom lands, on limestone hills, and
along roadsides in Franklin and Colbert counties, Alabama (Webb & Kral).
Although it is locally common in a few localities, it is an endangered species.
Lesquerella lyrata is readily distinguished from the other auriculate-leaved
species by its yellow flowers and its glabrous, depressed-globose fruits with an
opaque, complete septum and thick, leathery valves.
Lesquerella densipila Rollins, 2n = 16, occurs in open alluvial sites, fallow
fields, pastures, river bottoms, roadbanks, and cedar glades. It is abundant in
the Central Basin of Tennessee, particularly near the West Fork of Stones
1987] AL-SHEHBAZ, ALYSSEAE 225
Ficure 3. Fruits of Lesquerella. ac, L. globosa: a, fruit, x 3; b, septum and replum,
x |12—note eee) septum and position of seed; c, hemispherical seed, x 12. d, L.
gracilis, fruit, x 5. e, f, L. densipila: e, fruit, x 3%; f, replum and septum, x 5—note
midvein. g, h, L. eee g, fruit, x 214; h, septum and replum, x 5—note central
ston i, L. lyrata, fruit, x 3. j, L. stonensis, fruit, x 2%. k, L. “Lescurii, fruit, x 3.
River, the Duck River, and the upper Harpeth River (Rutherford, Bedford,
Williamson, Marshall, Maury, and Davidson counties). It also occurs, probably
as a recent introduction, in northern Alabama (Franklin, Lawrence, Morgan,
and Marshall counties). Lesquerella densipila is a very close relative of L. lyrata,
from which it can be distinguished by its dense indumentum of short, simple
or branched trichomes on the styles and the outer valve surfaces. According
to Rollins (1955), L. /yrata is morphologically and geographically intermediate
between L. densipila and L. auriculata (Engelm. & Gray) S. Watson (central
Texas and south-central Oklahoma) and may well be the evolutionary link
between the latter species and the auriculate-leaved members endemic to Ten-
nessee.
On the basis of fruit morphology, earlier authors considered Lesquerella
Lescurii (A. Gray) S. Watson (Vesicaria Lescurii A. Gray, Alyssum Lescurii
(A. Gray) A. Gray), 2” = 16, to be anomalous in the genus, and Payson placed
it in a monotypic section. As shown above, however, the species is very closely
226 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
related to and readily hybridizes with the auriculate-leaved lesquerellas of the
Southeast. It is most closely related to and was probably derived from L.
densipila. Lesquerella Lescurii is readily distinguished from all other species
of the genus by its fruits that are strongly flattened parallel to the septum and
its valves that are pubescent with a mixture of long, simple, bulbous-based
trichomes and short, branched ones. It grows in open areas of river-bottom
pastures, fields, and flood plains, as well as on thin soil over limestone in cedar
glades and on hill slopes, in north-central Tennessee (Summer, Wilson, Ruth-
erford, Davidson, Williamson, Cheatham, Montgomery, Dickson, and Stewart
counties), particularly along the Cumberland River and several tributaries of
the Harpeth River. Rollins (1981) listed L. Lescurii as having weedy tendencies.
It is adventive and has been recorded only recently from Alabama (Limestone
County) and Kentucky (Trigg County), by Kral and Chester, respectively.
A narrow endemic of central Tennessee, Lesquerella stonensis Rollins, 2n =
16, grows in pastures, flood plains, and fields and on knoll tops, as well as on
roadsides and stream banks along the East Fork of the Stones River (Rutherford
County). According to Kral, it is locally abundant in some years and almost
absent in others, with its present range restricted to a few fields along the Stones
River.
Lesquerella perforata Rollins, 2n = 16, is the nearest relative and perhaps a
direct descendant of L. stonensis. It is also a narrow endemic of Tennessee and
is presently known only within a radius of six miles around Lebanon (Wilson
County), where it grows in open fields, pastures, floodplains, and limestone
glades. Both species are easily distinguished from the other auriculate-leaved
lesquerellas by their perforated septa (FiGureE 3h) and their white petals with
yellow claws. Lesquerella stonensis has densely hirsute, depressed-globose to
subdidymous fruits, hirsute styles, and glabrous inner-valve surfaces, while L.
perforata has glabrous to sparsely hirsute, pyriform to obovoid fruits, glabrous
styles, and densely pubescent inner-valve surfaces.
In no other genus of Cruciferae has natural interspecific hybridization been
so well documented as in Lesquerella. In a series of papers, Rollins (1954,
1957) and Rollins & Solbrig (1973) demonstrated that species pairs involving
L. Lescurtt, L. densipila, and L. stonensis hybridize in all three combinations
in parts of Tennessee where their ranges come together. Hybrid populations
of L. densipila x L. stonensis (L. x maxima Rollins, L. densipila var. maxima
Rollins) were found in Rutherford and Davidson counties along the Stones
River downstream from the junction of its East and West forks, where L.
stonensis and L. densipila, respectively, grow. Those hybrids were more similar
to the former than to the latter species. The hybrid L. Lescurii x L. densipila
occupied a stretch of more than 40 miles downstream along the Harpeth River
between its junctions with Arrington Creek and the Cumberland River in
Williamson, Davidson, and Cheatham counties. The third hybrid combination,
L. stonensis x L. Lescurii, was found only once (in a vacant lot in the town
of La Vergne, Rutherford County) and was not directly associated with any
river system, unlike the other hybrid combinations.
The establishment, persistence, and population size of hybrids or their pa-
rental species in a given area are influenced by spring flooding of rivers, agri-
1987] AL-SHEHBAZ, ALYSSEAE Pie |
cultural practices, and factors controlling seed germination. For example, the
hybrid Lesquerella Lescurii x L. densipila, which was estimated in 1955 to
occupy approximately 600 acres around the junction of Arrington Creek and
the Harpeth River (Rollins, 1957), was reduced to less than 10 plants in 1966
because of the conversion of that area into pasture land (Rollins & Solbrig,
1973). Man’s agricultural activities in the Central Basin of Tennessee have
played a major role in bringing the ranges of the auriculate-leaved species of
Lesquerella into contact and consequent hybridization. These species are largely
allopatric and presumably evolved and persisted in isolation from each other
until a few decades ago. Very high degrees of interspecific fertility exist among
L. Lescurii, L. densipila, L. stonensis, L. perforata, and L. lyrata. Artificial
hybrids between any pair of these show very low levels of meiotic irregularities
that are not significantly different from those observed within each parental
species (Rollins, 1957; Rollins & Solbrig, 1973). The artificial first- and second-
generation hybrids have very high pollen quality, and their seeds germinate at
levels as high as 86 percent.
Many authors (e.g., Maguire & Holmgren; Mulligan; Payson; Rollins, 1939a,
1950, 1983; Rollins & Shaw, 1973) have emphasized the very close relationship
between Lesquerella and Physaria (Nutt.) A. Gray (22 species; Alberta, the
Pacific and Mountain states, Arizona, and New Mexico). It is generally agreed
that Physaria is derived from Lesquerella. The line separating them is artifi-
cially drawn, and there is a continuous morphological gradation from one to
the other. Both genera, however, should be maintained. Similar situations exist
between pairs of related genera throughout the Cruciferae, and it is not practical
to merge the larger Lesquere/la with the smaller and earlier-published Physaria
(Rollins, 1950; Rollins & Shaw, 1973). Physaria differs from Lesquerella in its
highly inflated, always didymous fruits either markedly constricted at the re-
plum or strongly flattened contrary to the septum (angustiseptate). In general,
the fruits of Lesquerella are not inflated, not didymous, and not constricted at
the replum. There are, however, some exceptions. In L. inflata Rollins & Shaw
and L. perforata the fruits are inflated, while in L. hemiphysaria Maguire and
L. stonensis they are subdidymous. Angustiseptate fruits are found in unrelated
species of Lesquerella and are well developed in L. carinata Rollins, L. Paysonii
Rollins, and L. /asiocarpa (W. J. Hooker) A. Gray var. Berlandieri (A. Gray)
Payson. Physaria oregona S. Watson, P. Geyeri (W. J. Hooker) A. Gray, and
P. alpestris Suksd. have slightly inflated fruits. They were transferred to Les-
querella by Mulligan but, as shown by Rollins & Shaw (1973), should be
retained in Physaria.
Lesquereila is also related to Synthlipsis A. Gray (three species; Texas and
northern Mexico), and L. Jasiocarpa var. Berlandieri was suggested as the
possible link between the two genera (Rollins, 1955; Rollins & Shaw, 1973).
The Old World genera Alyssoides (= Vesicaria Adanson) (four species; southern
France, Balkan peninsula, Turkey) and A/yssum were also said to be closely
related to Lesquerella (Rollins, 1950; Rollins & Shaw, 1973). Both Alyssum
and Alyssoides have stellate trichomes indistinguishable from those of Les-
guerella, but they differ in their winged or appendiculate staminal filaments,
their pollen morphology, their winged seeds, and their lack of a nerve in the
228 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
septum. In my opinion, Lesqguerella should be associated only loosely with
Alyssum and Alyssoides
The tribal disposition of Lesquerella 1s problematic. Schulz placed the genus
in the tribe Drabeae and assigned its nearest relative, Physaria, to the tribe
Lepidieae. He defined the latter tribe mainly on the basis of its angustiseptate
fruits. As delimited by Rollins & Shaw (1973), however, Lesquerella contains
several species (see above) with such fruits. According to Schulz’s key to the
tribes, various species of Lesquerella will be identified in the Alysseae, the
Drabeae, and the Lepidieae. Angustiseptate fruits evolved independently in at
least four tribes of the Cruciferae (Al-Shehbaz, 1986). Angustiseptate and la-
tiseptate fruits are found in Lesquerella, Graellsia Boiss., Smelowskia C. A.
Meyer, and Nerisyrenia Greene. Therefore, the type of flattening of fruits is
not always useful for assigning genera to tribes. Rather, problematic genera
such as Lesquerella should be placed in the tribe containing what seem to be
their nearest relatives.
On the basis of pollen morphology, Lesquerella should be associated with
Physaria, Synthlipsis, Nerisyrenia, and Dimorphocarpa Rollins. All these New
World genera have 5- to 10-colpate pollen grains not found elsewhere in the
Cruciferae (Rollins, 1979; Rollins & Banerjee, 1979; Rollins & Shaw, 1973).
The last genus is closely related to Dithyrea Harvey, which has 4-colpate pollen.
Because all of these genera have angustiseptate fruits and are traditionally
assigned to the Lepidieae, there is no major obstacle to placing Lesquerella
with them. Von Hayek’s grouping of these genera in one tribe was more natural
than Schulz’s, but he assigned them, along with several unrelated genera, to
the tribe Schizopetaleae Prantl, which was considered to have a polyphyletic
origin from tribe Thelypodieae Prantl. In this paper I have placed Lesquerella
in the Alysseae, following the modified tribal classification adopted earlier (Al-
Shehbaz, 1984). It is obvious, however, that the genus is more appropriately
placed with its nearest relatives in the Lepidieae.
Although individual flowers are not showy in Lesquerella, they are densely
grouped in compact inflorescences that can be quite attractive. These are visited
by various species of flies, butterflies, and solitary bees, but the most common
pollinator in the Southeast is the introduced honey bee (Apis mellifera). Self-
incompatibility is widespread in Lesquerella and occurs in all of the auriculate-
leaved species growing in our area (Rollins, 1957; Rollins & Solbrig, 1973,
Sampson).
Chromosome numbers have been reported for at least 52 species, the majority
of which are diploid with n = 5 to 10. Polyploidy did not play a major role in
the evolution of Lesquerella, and only ten species have both diploid and poly-
ploid populations. Three species, L. mendocina (Phil.) Kurtz (South America),
L. arctica, and L. peninsularis Wiggins (Baja California) are polyploid, with
2n = 50, 60, and ca. 40 or 48, respectively. Diploid, tetraploid, and hexaploid
populations are found in L. Engelmannii (A. Gray) S. Watson (x = 6) and L.
ludoviciana (Nutt.) S. Watson (x = 5). Complex aneuploid series occur in both
L. argyraea (A. Gray) S. Watson and L. ovalifolia Rydb. subsp. ovalifolia
(Clark; Rollins & Shaw, 1973). Except for L. grandiflora (W. J. Hooker)
A. Gray (2n = 18) and L. /asiocarpa (2n = 14), the remaining auriculate-leaved
1987] AL-SHEHBAZ, ALYSSEAE 229
species have 2m = 16. The uniformity in chromosome numbers and in the
presence of densipolic acid (see below) support the placement of these auric-
ulate-leaved species in a position somewhat remote from L. grandiflora and
L. lasiocarpa.
Seeds of the auriculate-leaved species of Lesquerella endemic to the Southeast
(L. densipila, L. Lescurii, L. lyrata, L. perforata, and L. stonensis) contain high
concentrations of densipolic acid (C,,) and lack lesquerolic acid (C,.). Those
of sixteen other species (including L. grandiflora and L. lasiocarpa) are rich in
lesquerolic acid (45-72 percent of the total fatty-acid content). In L. auriculata,
which is believed to be the link between the auriculate-leaved species of the
Southeastern States and the rest of the genus, small amounts of densipolic (two
percent) and lesquerolic (ten percent) acids were found, in addition to high
concentrations (32 percent) of auricolic acid. The last is lacking in our auric-
ulate-leaved species and is only a minor constituent in many other species of
Lesquerella (Appelqvist). It is a higher homologue of densipolic acid, while
lesquerolic acid is a higher homologue of ricinoleic acid, a trace acid present
throughout the genus.
Five glucosinolates were found in 13 species (Daxenbichler et a/., 1961,
1962). 6-Methylthiohexylglucosinolate occurs in all of the auriculate-leaved
species (Lesquerella auriculata was not analyzed) and in L. Engelmannii. Other
compounds were 4-methylthiobutyl, 3-methylthiopropyl, isopropyl, and 2-bu-
tyl glucosinolates.
Trichome diversity in Lesquere/la is probably greater than that in any other
genus of Cruciferae. Rollins & Banerjee (1975, 1976) studied the trichomes of
69 species of Lesquerella and observed well-marked trends of specialization.
From the dendritic type (stalked, with unequal branches forming an irregular
pattern), which is presumably primitive, stellate trichomes evolved by the
reduction of irregularities in branching and by the disposition of the rays in
one plane. Further specialization from stellate trichomes with few, simple rays
proceeded in two directions. The first trend, found in many species, 1s a pro-
gressive increase in the branching of rays. As a result, two- or four-forked rays
either without thickened bases (L. macrocarpa A. Nelson) or with massively
thickened and fused bases (L. Hitchcockii Munz, L. rubicundula Rollins, and
L. thamnophila Rollins & Shaw) probably evolved. In the second trend, found
in at least ten species, the increase in the number of simple rays 1s correlated
with a centrifugal increase of webbing between the rays. The two representative
extremes of this trend are L. Douglasii S. Watson (with about 13 rays webbed
only between their bases) and the highly specialized L. mexicana Rollins (with
ca. 50 rays webbed to their tips and forming peltate scales).
The trichomes of the Cruciferae are unicellular, and those of Lesquerella
have calcium carbonate deposited as calcite on the interior of the cell wall
(Lanning, 1961). Rollins & Shaw (1973) indicated that there is a broad cor-
relation between the density of trichomes and the availability of moisture.
Species growing in arid areas and at high elevations have the densest trichome
covering, while those of mesic areas have a sparse indumentum. Ancibor showed
that the fully developed trichome remains alive and has a very conspicuous
nucleus and a dense cytoplasm. She suggested that trichomes may have a water-
230 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
absorbing function, but Rollins & Shaw (1973) indicated that they probably
reduce water loss from plants of arid areas by reflecting light rays, by forming
a layer that slows down air movement, and by establishing a moisture gradient
between the epidermis and the open air.
Lesquerella has little if any economic value. Several species analyzed for
fatty-acid content show very high concentrations of hydroxy acids, which are
valuable in industry. Tough plastics and reinforced elastomers have been pro-
duced from the oils of L. Palmeri S. Watson. Hinman (1984, 1986) suggested
that L. Fendleri (A. Gray) S. Watson has superior qualities and can compete
with castor bean (Ricinus communis L.) in its industrial oils. It has no allergenic
or toxic properties, 1s capable of growing on sandy or calcareous soils of semiarid
areas, tolerates cold and drought, and can be harvested by combine. No species
of the genus, however, is a crop, and the agronomic values of most have not
been evaluated.
REFERENCES:
Under family references in AL-SHEHBAZ (Jour. Arnold Arb. 65: 343-373. 1984), see
BRITTON & BROWN; VON HAYEK; JONES; MANTON; ROLLINS (1966, 1981); ROLLINS &
BANERJEE (1979); ROLLINS & ie enone (1971, 1977, 1979); ScHULz; SMALL; E. B.
SmitH; and WELSH & REVEAL.
Under tribal references see BOLKHOVSKIKH et al.; GATTINGER; GOLDBLATT (1981, 1984,
1985); KUMAR & TsUNODA; Moore; PRINCEN & ROTHFUS; and SMALL.
AL-SHEHBAZ, I. A. The genera of Lepidieae (Cruciferae; Brassicaceae) in the southeastern
United States. Jour. Arnold Arb. 67: 265-311. 1986.
Ancisor, E. Ontogenia y morfologia de los pelos de Lesquerella mendocina (Phil.)
Kurtz var. microcap O. E. Schulz (Cruciferae). (English summary.) Physis C. 38:
63-67. 1978
APPELOVIST, L. bh oon in the Cruciferae. Pp. 221-227 in J. G. VAUGHAN, A.
MacLeop, & B. M. G. Jones, eds., The biology and chemistry of the Cruciferae.
London, New York, and San Francisco. 1976. [Lesquerella, 224-227.
AYENSU, E. S., & R. A. DeFiuipps. Endangered and threatened plants of the United
States. xv + 403 pp. Washington, D. C. 1978. [L. densipila, L. perforata, L. lyrata,
and L. stonensis endangered, 76; L. globosa and L. Lescurti threatened, 103; L.
macrocarpa assumed extinct, 76; pone species listed.
Barc ay, A. S., H. S. Gentry, & Q. J The search for new industrial crops II:
Lesquerella (Cruciferae) as a source ae new oilseeds. Econ. Bot. 16: 95-100. 1962.
[Seed weight, protein and oil content, percentages of C,, and C,, hydroxy acids; 17
species including those indigenous to the Southeastern States.]
Bass, L. N., & D. C. CLARK. Persistence of the dormancy-breaking effect : gibberellic
acid on — seeds. Proc. Assoc. Off. Seed Anal. 63: 102-105. 1973.*
. SAYERS. Germination experiments with seed . Lesquerella
species. Proc. ee Off. Seed Anal. 56: 148-153. 1966.*
Brnper, R. G., & A. Lee. Py ae acids of Lesquerella densipila seed oil.
Jour. Org. Chem. 31: 1477-1479.
Brooks, R. E. On the name and ss of white flowered Lesquerella Engelmannii in
northcentral Kansas. (Abstr.) Am. Jour. Bot. 73: 753. 1986. [Chromosome numbers,
CHESTER, E. W. Some new distributional records for Lesquerella Lescurti (Gray) Watson
(Brassicaceae), including the first report for Kentucky. Sida 9: 235-237. 1982. [New
to Dickson County, Tennessee, and Trigg County, Kentucky; map.]
1987] AL-SHEHBAZ, ALYSSEAE 231
CLarK, C. Ecogeographic races of Lesquerella Engelmannii (Cruciferae): distribution,
chromosome numbers, and taxonomy. Brittonia 27: 263-278. 1975. [L. ovalifolia
reduced to two subspecies of L. Engelmannii.]
CRANFILL, R., J. M. Baskin, & M. E. MEDLEY. Taxonomy, distribution and rarity status
of Leavenworthia and Lesquerella rasicacen) in Kentucky. Sida 11: 189-199.
1985. [Lesquerella globosa, L. Lescur
DAXENBICHLER, M. E., C. H. VANETTEN, aL “A.W OLFF. Identification ofa new, naturally
occurring, eee: tile isothiocyanate from Lesquerella ee seed. Jour. Org.
Chem. 26: 4168, cee 1961. [6-Methylthiohexyl isothiocyan
Lt, & I. A. Woxrr. Isothiocyanates from Seay hydrolysis
of Lesquerella seed Pec Jour. Am. Oil Chem. Soc. 39: 244, 245. 1962. [Analysis
of 17 ee: estimates of total volatile isothiocyanates, identification of five com-
poun
FREEMAN, 1D ., A. S. Causey, J. W. SHort, & R. R. Haynes. Endangered, threatened,
and special concern plants of Alabama. 25 pp. Auburn, Alabama. 1979. [L. densipila
and L. /yrata endangered. ]
Gentry, H. S., & A. S. BARCLAY. The search for new industrial crops III: prospectus
of Lesquerella Fendleri. Econ. Bot. 16: 206-211. 1962. [Variation, environmenta
requirements, yield, agronomic values.]
HINMAN, C. W. New crops for arid lands. Science 225: 1445-1448. 1984. [L. Fendleri,
1447]
——. Potential new crops. Sci. Am. 255(1): 32-37. 1986. [Lesquerella, 37.]
JAKOWSKA, S. The trichomes of Physaria Geyeri, P. australis and Lesquerella Sher-
woodii: development and morphology. Bull. Torrey Bot. Club 76: 177-195. 1949.
[L. Kingii var. diversifolia as L. Sherwoo
. The resting nucleus in ee and Lesquerella. Ibid. 78: 221-226. 1951. [Same
taxa as in the preceding pap
baste B. Lesquerella arctica (Wormskj,) Wats. in Siberia. Notul. Syst. 21: 148-157.
961. [Circumpolar distribution, ae aeons map.
ae G. F. SPENCER, F. R. E, H. J. NiESCHLEG, & A. S. BARCLAY. Tetra
acid triglycerides containing a new ede ao ato moiety in eee
auriculata seed oil. Lipids 7: 660-665. 1972.
Know .es, R. E., K. W. TAyLor, G. O. KoHLer, & L. A. GOLDBLATT. Hydroxy-unsat-
urated oils and meals from PL le and Lesquerella seed. Jour. Agr. Foo
Chem. 12: 390-392. 1964. [L. Fer
KRAL, R. A report on some rare, ser en or endangered forest-related vascular eae
of the South. U. S. Dep. Agr. Forest Serv. South Reg. Tech. Publ. R8-TP2. V
x + 718 pp. 1983. [L. Gaia L. globosa, L. Lescurii, L. lyrata, L. ee ZL,
stonensis; descriptions, habitats, maps, 508-527
LANNING, F. Calcite in Lesquerella ovalifolia trichomes. Science 133: 380. 1961.
[Trichomes contain calcium carbonate deposited as calcite on cell wall.
. Ash, silica, and calcium in Lesquerella ovalifolia. Trans. Kansas Acad. Sci. 67:
481-485. 1964.
LUNDELL, C. L. Studies of American plants— XIV. Wrightia 5: 331-351. 1977. [L.
gracilis var. pilosa, var. nov., 331; photo of type in ibid. 6: pl. 86. 1979; = L.
Lindheimeri.]
MAcurIrE, B., & A. H. Ho-mcren. Botany ofthe Intermountain Region—II. Lesquerella.
Madrofio 11: 172-184. 1951, [Infraspecific classifications of L. Hitchcockii, L.
Mikotasczak, K. L., F. R. EARLE, & I. A. Wotrr. Search for new industrial oils. VI.
Seed oils of the genus aoe Jour. Am. Oil Chem. Soc. 39: 78-80. 1962.
[Fatty-acid content of 14s
MILLER, oe ae H. REE. RL A. Wo.trr. Amino acid composition of Les a
seed m s. Jour. Am. Oil Chem. Soc. 39: 115-117. 1962. [Distribution ma amin
acids in on species. ]
Zag JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
MuLuicaAn, G. A. Transfers from Physaria to Lesquerella (Cruciferae). Canad. Jour.
Bot. 46: 527-530. 1968. [Four species transferred, chromosome numbers; see ROLLINS
& SHaw, 1973.,]
A. E. Porsitp. A new species of Lesquere/la (Cruciferae) in northwestern
Canada. Canad. Jour. Bot. 47: 215, 216. p/. J. soe [L. Calderi.]
Nixon, E. S., J. R. WARD, & B. L. Lipscoma, Rediscovery of Lesquerella pallida (Cru-
ciferae). Sida 10: 167-175. 1983. [Description, peers SEMs of pollen and
trichomes, comparison with L. gracilis. ]
Payson, E. B. A monograph of the genus Lesquerella. Ann. Missouri Bot. Gard. 8: 103-
236. 1922. del trends, distributions, sectional classification, taxonomic
treatment of 52 specie
Princen, L. H. New oilseed crops on the horizon. Econ. Bot. 37: 478-492. 1983.
[Lesquerella, 486.]
QUARTERMAN, E. Studies on the distribution and life history of two species of Lesquerella
(Cruciferae). (Abstr.) ASB Bull. 7: 37. 1960. [L. densipila, L. Lescurii.]
REVEAL, J. L. Comments on mae Hitchcockii. nike Basin Nat. 30: 94-98. 1970.
[Recognized ae species; see ROLLINS & SHAW,
Rocers, K. E., & F. D. Bowers. Notes o on csi plants III. Castanea 38: 335-339.
1973. [L. aracilis, infrequent in railroad yards in Memphis and Nashville, 338.]
Ro tins, R. C. The cruciferous genus Physaria. Rhodora 41: 392-415. pl. 556. 1939a.
pagers between Physaria and hates 393, 394]
n the genus Lesquerella. Am. Jour. Bot. 26: 419-421. 1939b. [Chro-
mosome ane of six species; three new mn
udies on some North American Cruciferae. Contr. Gray Herb. 171: 42-53.
1950. [L. carinata, L. Paysonii, L. Mcvaughiana, spp. nov.; relationships between
Lesquerella and both Physaria and ea 42-47. ]
Some Cruciferae of the Nashville Basin, Tennessee. Rhodora 54: aaa 1952.
[L. densipila and L. perforata (spp. nov.), L. Lescurii, L. globosa, k
. Interspecific hybridization and its role in plant evolution. Eighth Internat. Bot.
Congr. Paris Rapp. Comm. Sed. 9 & 10: 172- a 1954. [Hybridization between
L. densipila and L. Lescurii along the Harpeth River, Tennessee. ]
The auriculate-leaved species of Lesquerella (Craciferne) Rhodora 57: 241-
264. pls. 1207-1212. 1955. [Key, descriptions, distributions; L. /yrata and L. ston-
ensis (spp. nov.), L. lasiocarpa, L. Lescurti, L. densipila, L. perforata, L. auriculata,
L. grandiflora.
On the identity of Lesquerella epg le Ibid. a 199-202. 1956. [Distri-
orn
Interspecific hybridization in Lesquerella. oe nat Herb. 181: 3-40. 1957.
[Detailed study of natural hybridization between L. densipila and both L. Lescurti
and L. stonensis in Central Basin, Tennessee; see ROLLINS & SOLBRIG, 1973.]
otes on Lesquerella (Cruciferae) in hn Bol. Soc. Bot. Méx. 23: 43-47.
1958. [L. mexicana, L. Mirandiana, spp. nov.]
ithyrea and a related genus ae ae Publ. Bussey Inst. Harvard Univ.
1979: 3-32. 1979. [Taxonomic treatment of a aa gen. nov. (four spp.)
and Dithyrea (two spp.), SEMs of pollen; 24 figs., 4 pls.,
pecies of Draba, Lesquerella, and Sibara (Cruise oe Gray Herb. 211:
107- 113. 1982. [L. kaibabensis, sp. nov., 1.]
—. Studies in the Cruciferae of western North oe Jour. Arnold Arb. 64: 491-
510. 1983. [Seed dispersal, relationship between Lesquerella and Physaria, 491-
493; L. Goodrichii and L. parviflora, spp. nov., 503-507.]
. Studies in the Cruciferae of western North America. II. Contr. Gray Herb. 214:
1-18. 1984a. [Lesquerella, 7-11.]
. Studies on Mexican Cruciferae II. /bid. 19-27. 1984b. [Lesquerella, 22-24.]
— & U.S. Banersee. Atlas of the trichomes of Lesquerella (Cruciferae). Publ.
1987] AL-SHEHBAZ, ALYSSEAE 233
Bussey Inst. Harvard Univ. 48 pp. 1975. [SEMs of trichomes of 69 species and 11
suse trends of specialization, /20 SEM photos, 20 pls.]
. Trichomes in studies of the Cruciferae. Pp. 145-166 in J. G. VAUGHAN,
AJ. MacLeop, & B. M. G. Jones, eds., The biology and chemistry of the Cruciferae.
London, New York, and San a 1976. [Trichomes of Lesquerella, trends of
differentiation, 36 SEM figs., 6
& . SHAW. Neca aes in ee Rhodora 74: 76-79. 1972.
[Infraspecific taxa ae L. lasiocarpa, tauton
& The genus Lesquerella eS in North America. x + 288 pp.
Cambridge, Nee iccits 1973. [Taxonomic treatment of 69 species and 29 infra-
specific taxa, chromosome numbers, SEMs of pollen and trichomes, hybridization;
32 pls., 28 maps; the best and most comprehensive account.]
. T. SoLBRic. Spatial and east variation in hybrid aaa of Les-
querella densipila x L. Lescurii. (Abstr.) Am. Jour. Bot. 58: 466.
Interspecific hybridization in Lesquerella. Contr. ae roe 203: 3-
48. 1973. [Detailed analyses of 23 characters in L. densipila, L. Lescurit, L. lyrata,
L. perforata, and L. stonensis and 1 in their second-generation artificial hybrids, cross-
stonensis in eight localities in the Central Basin, Tennessee; see Rotuins, 1954,
ie
1957.]
SAMPSON, D. R. The genetics of self-incompatibility in Lesquerella ene and in the
F, hybrid L. densipila x L. Lescurii. Canad. Jour. Bot. 36: 39-56. 19
SHARIR, A., & H. GELMOND. Germination studies of een Fendleri - L. Gor
donii, with reference to their caer ne Econ. Bot. 25: 55-59. 1971. [Seed ee
mancy, gibberellic acid, light requiremen
SMALL, J. K. Flora of the southeastern ae States. ed. 2. xii + 1394 pp. New York.
1913. [Lesquerella, 468—471.]
Situ, C. R., Jr., T. L. Witson, R. B. BATEs, & C. R. SCHOLFIELD. Densipolic acid: a
unique hydroxydienoid acid from eee densipila seed oil. Jour. Org. Chem.
27: 3112-3117. save
_K. . H. Zoper, R. L. Lonmar, & I. A. WoLFr. Lesquerolic
acid. : new Nee boa from een seed oil. Jour. Org. Chem. 26: 2903-
2905.
SUPAVARN, rs a W. Knapp, & R. SIiGAFUS. Investigations of mucilaginous seeds as
potential biological control agents against mosquito larvae. Mosq. News 36: ae
182. 1976. [Mucilage of 16 species of Lesquerella, including all seven growing i
the southeastern United States, caused a to larvae of Aédes aegyptt, aes
mortality, 85 percent, was caused by L. argyra
VanatTra, E. G. Notes on the leaf hairs of inter Proc. Acad. Nat. Sci. Phila. 59:
247, 248. pl. 27. 1907. [Twenty-three species in five groups.]
VIELLION, M. K. A taxonomic study of Lesquerella S. Wats. (Cruciferae) i in the Great
Plains. xi + 137 pp. Unpubl. Ph.D. dissertation, Univ. Nebraska. 1973. [Numerical
study of 12 species, key, aes distributions, phylogenetic relationships.]
Warp, D. E. Chromosome counts fro pats xico and southern Colorado. Phytologia
54: 302- 308. 1983a. [L. aurea, n = 7; L. valida, n = 5.]
. In: A. Léve, ed., IOPB a ena number reports LX XX. Taxon 32: 504—
S11. 1983b. [L. purpurea, 510,n = 9.
Watson, S. Contributions to American botany. 1. Some new species of plants of the
United States, with revisions of Lesquerella (Vesicaria) and of the North American
species of Draba. Proc. Am. Acad. Sci. 23: 249-267. 1888. [Lesquerella, 249-255;
33 species in two sections. ]
Wess, D. H., & R. KRAL. Recent collections and status of Lesquerella lyrata Rollins
(Cruciferae). Sida 11: 347-351. 1986. [Distribution in Franklin and Colbert counties,
labama, recommended management practices, map.
234 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
26. Camelina Crantz, Stirp. Austriac. 1: 17. 1762.
Spring or winter annual or biennial herbs, with furcate-stellate and[/or] sim-
ple trichomes, sometimes glabrescent; stems simple or branched at base, often
branched above. Basal leaves petiolate or subsessile, usually not in a rosette,
entire to sinuate. Cauline leaves sessile, sagittate or amplexicaul, oblong to
linear or lanceolate, entire or dentate, gradually decreasing in size upward.
Inflorescences ebracteate, corymbose racemes, greatly elongated in fruit; fruit-
ing pedicels horizontal to ascending [or appressed to rachis]; rachis of infruc-
tescence straight [or flexuous], glabrous [or pubescent]. Sepals oblong, erect,
equal, not saccate at base, usually membranaceous at margin, villous in bud,
often glabrescent. Petals yellow to white, clawed, spatulate, attenuate at base,
longer than sepals. Nectar glands 4, | on each side of lateral stamens, median
glands absent. Stamens 6, usually in 3 different lengths; filaments linear, free,
unappendaged, slightly dilated at base; anthers oblong to ovate. Fruits usually
dehiscent, obovate or narrowly to broadly pyriform [or linear], somewhat
flattened parallel to the septum, short stipitate, strongly keeled and narrowly
winged at the replum, rounded [truncate or notched] at apex, ending abruptly
ina stylelike beak; valves thick, slightly to strongly convex, obscurely to prom-
inently reticulate, the midvein evident in the lower half or along the entire
length of the valve, glabrous [or pubescent], glossy on inside, the acuminate
apex extending 0.5-1.5 mm into the beak area; beaks longer to shorter than
styles; styles filiform, persistent; stigmas capitate; replum covered by the con-
nate margins of valves, becoming visible after fruit dehiscence. Seeds 4-12 per
locule, reticulate, oblong, copiously mucilaginous when wet, biseriately [or
uniseriately] arranged in each locule; cotyledons incumbent or rarely accum-
bent. Base chromosome numbers 6, 7, 10, 13. (Including Dorella Bubani, non
Weber-van Bosse, Linostrophum Schrank.) Type species: Myagrum sativum
L. = Camelina sativa (L.) Crantz. (Name of obscure origin, possibly derived
from Greek chamai, dwarf or on the ground, and /inon, flax, perhaps referring
to the stunting or suppressing influence of Camelina on the growth of flax.)
— FALSE FLAX, GOLD-OF-PLEASURE, FLAXWEED.
A well-marked genus of six or seven species centered in Turkey and adjacent
parts of southwestern Asia and southeastern Europe. Camelina is represented
in North America by four naturalized species, of which two occur in the south-
eastern United States.
Of the two sections recognized in Camelina by De Candolle (1821, 1824),
sect. CAMELINA (as sect. Chamaelinum DC.) is now retained in the genus, while
sect. PSEUDOLINUM DC. has been transferred to Rorippa Scop. Boissier’s sec-
tional classification, which was accepted by Schulz and neglected by many
subsequent authors, is more practical than any other infrageneric classification
of Camelina. The monotypic sect. ErystmAsTruM Boiss. (fruits linear-cylin-
drical, seeds uniseriate) includes C. anomala Boiss. & Hausskn. of southern
Turkey and the Bekaa valley, Lebanon. Section CAMELINA (fruits obovate to
pyriform, seeds biseriate) contains the remaining species of the genus. On the
basis of seed size and other characters of continuous nature, Mirek (1981)
1987] AL-SHEHBAZ, ALYSSEAE Pah,
recognized two series in Camelina. It is doubtful, however, that these improve
the taxonomy of the genus.
Camelina microcarpa Andrz. ex DC. (C. sativa subsp. microcarpa (Andrz.)
E. Schmid), false flax, 2” = 40, which is naturalized throughout North America,
grows in grainfields, meadows, waste places, and disturbed habitats, as well as
along roadsides, in North and South Carolina, Georgia, Tennessee, Arkansas,
and Louisiana. It is likely to be found in the remaining states of the Southeast.
It is distinguished by its mixed simple and furcate-stellate trichomes on the
lower part of the stem, fruits 2.5-5 mm long, petals to 4.2 mm long, and seeds
0.9-1.5 mm long.
Most of the earlier reports of Camelina sativa (L.) Crantz (Myagrum sativum
L., Alyssum sativum (L.) Scop., C. sativa var. glabrata DC., C. glabrata (DC.)
K. Fritsch), false flax, flaxweed, gold-of-pleasure, Dutch-flax (Small), 2n = 40,
from North and South Carolina, Tennessee, Arkansas, and Louisiana are doubt-
ful and may well represent misidentifications of plants of C. microcarpa. Ca-
melina sativa is less common in North America than C. microcarpa and may
be only a waif in the southeastern United States. It is distinguished from C.
microcarpa by its glabrous or sparsely stellate-furcate stems rarely with simple
trichomes, its longer fruits 7-9 mm long, and its shorter infructescences with
fewer fruits.
Camelina Rumelica Velen., which has only recently been recorded in the
United States (McGregor, 1984), is easily confused with C. microcarpa because
of similarities in fruit shape and size, seed length, and lack of trichomes on
the infructescence. It differs in its petals 5-9 mm long, in having only simple
trichomes with some usually 2-3.5 mm long, and in its fruits more widely
spaced in the lower than the upper part of the infructescence. The species is
naturalized in Oklahoma, Texas, Kansas, Colorado, Nevada, and Oregon.
Camelina has obovate-pyriform (very rarely linear) fruits, thick valves
abruptly acuminate into a stylelike beak (FiGuRE le), and connate valve margins
that make the replum invisible until dehiscence. The genus is related to Chry-
sochamela Boiss. (three species; Turkey and Syria) but certainly not to Nes/ia
Desv., Capsella Medicus, or Cochlearia L., as has been suggested by several
earlier workers.
The tribal disposition of Camelina is controversial, and there is little or no
agreement among the several classifications consulted. The genus was placed
in the Camelineae DC. (De Candolle, 1821, 1824; Bentham & Hooker), the
Lepidieae DC. (Von Hayek, Janchen), and the Sisymbrieae DC. (Schulz). Al-
though the placement of Camelina in the Alysseae is only a minor improve-
ment, it is obvious that the boundaries of the Alysseae defined above are not
natural and that this tribal disposition of the genus is not final.
Camelina is taxonomically troublesome, and only three species, C. anomala,
C. laxa C. A. Meyer (Turkey, Iran, the Caucasus), and C. hispida Boiss. (Syria,
Turkey, Iran) are distinct. The last is variable and contains three varieties
(Hedge). The remaining species are weeds, and the boundaries between some
of them are artificially drawn. Forms intermediate between C. Rumelica and
C. microcarpa and between the latter and C. sativa have been found. Camelina
236 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Alyssum (Miller) Thell., a weed of flax (Linum usitatissimum L.) fields in
Europe said to be naturalized in the Dakotas and southern Canada, is com-
pletely interfertile with C. sativa. Their hybrids produce a very large array of
intermediates found in nature (Tedin, 1925; Sinskaja & Beztuzheva). There is
continuous variation in every character said to distinguish the two species. The
nomenclature of the weedy camelinas can be quite misleading, and the number
of species recognized has varied from seven (Vasil’chenko) to five (Meikle),
four with several subspecies and varieties (Smejkal; Mirek, 1981; see both for
extensive synonymy), and two (one of which has four subspecies) (Markgraf).
In my opinion, C. Rumelica, C. microcarpa, and C. sativa are sufficiently
distinct to merit specific status, but C. A/yssum should be treated as a subspecies
of the last (as C. sativa subsp. A/yssum (Miller) E. Schmid). Interspecific hy-
bridization has probably been responsible for blurring species boundaries, which
otherwise are sharply defined in areas of allopatry. The pattern of continuous
variation between species has been interpreted as a series of evolutionary
differentiations from C. microcarpa to C. sativa and from the latter to C.
Alyssum (Zinger). Both man’s selection of flax (see below) and natural hybrid-
ization probably played major roles in creating the taxonomic complexity of
the weedy camelinas.
Although most chromosome counts for Camelina sativa (including subsp.
Alyssum) and C. microcarpa agree on 2n = 40, counts of 2” = 26 and 2n =
16 and 32 have been reported for these species, respectively. Manton suggested
that the base chromosome number for Camelina is eight and that all species
are pentaploid, while others have believed that they are tetraploids based on
ten. Stebbins, on the other hand, suggested that they may well be ancient
allotetraploids, the ancestral species of which are unknown. Diploid counts of
2n = 12 and 2 = 14 are reported for C. Rumelica and C. hispida, respectively
(Brooks; Goldblatt, 1984), but other counts (2n = 24, 26, and 40) are also
recorded for the former.
The chemistry of Camelina is poorly understood. The scant data indicate
that both C. sativa and C. microcarpa contain 10-methylsulfinyldecylglucosi-
nolate (Kjaer ef al.). These species and C. Rumelica have uniform fatty-acid
composition characterized by high concentrations (33-38 percent) of linolenic
acid, by lower and nearly equal amounts (9-19 percent) of oleic, linoleic, and
eicosenoic acids, and by negligible amounts (1-3 percent) of erucic acid (Kumar
& Tsunoda).
The mode of origin of Camelina species as weeds of flax fields was studied
by Zinger, Sinskaja & Beztuzheva, and Tedin (1925) and was reviewed by
Hjelmqvist, Stebbins, and Barrett. According to these authors, certain forms
of C. sativa (variously recognized as varieties, subspecies, or species) originated
under selection pressures (climatic, phytosociological, agricultural —e.g., thresh-
ing and winnowing) operating in the cultivation of flax. Whether flax is grown
for fiber or for seed oils, it is “mimicked” by plants of C. sativa in growth
habit, branching pattern, internode length, leaf width, stem diameter and pu-
bescence, flowering time, fruit dehiscence, and winnowing properties of the
seeds. Camelina sativa subsp. A/yssum (listed in the literature as a distinct
species or as a subordinate of C. sativa under the epithets Alyssum, macrocarpa,
1987] AL-SHEHBAZ, ALYSSEAE Zot
foetida, dentata, and linicola) scarcely grows outside flax fields, and it has
evolved winnowing characteristics so similar to those of flax that their seeds
remain mixed and are therefore resown the following season. Other aspects of
Camelina-Linum relationships have been discussed by Stebbins and Barrett.
It has been shown that competition between Camelina and flax reduces the
yield of the latter and produces in it smaller leaves, thinner stems, reduced
branching, and smaller infructescences (Balschun & Jacob, 1961, 1972; Kranz
& Jacob). Griimmer & Beyer demonstrated that the decline in productivity of
flax is caused by allelopathic effects of leaf phenolic compounds (e.g., vanillic,
p-hydroxybenzoic, and ferulic acids) washed from Camelina by rain. Others
(e.g., Lovett and various co-workers) indicated that in the presence of certain
free-living, nitrogen-fixing bacteria, aqueous washings of foliage of C. sativa
(presumably containing toxic degradation products of isothiocyanates) stim-
ulate the early growth of flax but inhibit its later growth and may cause marked
ultrastructural changes in its root tips.
Species of Camelina accompanied the spread of agriculture in prehistoric
times. Subfossil remains (as carbonized seeds) date back to the Neolithic and
the Iron Age. Camelina sativa was cultivated for its stem fibers and edible oils
by the Romans as early as 600 B.c. Despite the drastic decline in its cultivation,
it is still grown in parts of Europe and the Soviet Union for the same purposes.
The seeds contain 34—42 percent oil and about 33 percent protein. The seed
oil has been used as an illuminant and for making soap, while the seed cake
is fed to cattle.
REFERENCES:
Under family references in AL-SHEHBAZ (Jour. Arnold Arb. 65: 343-373. 1985), see
ApPELQVIST (1976); BAILLON; BENTHAM & HOOKER; BERGGREN; De CANDOLLE (1821,
1824); EASTERLY oe FERNALD; Von Hayek; HEDGE & RECHINGER; JANCHEN; LA
RTE; MANTON; MARKGRAF; Sap pia abel RADFORD et al.; RICKETT;
Rotuins (1981); cis and E. B. Sm
Under tribal references see BOLKHOVSKIKH ef al.; DUNCAN & KARTESZ; GATTINGER;
GOLDBLATT (1984, 1985); KuMAR & TsuNoDA; LITCHFIELD; MACRoBERTS, Moore, and
Ancev, M.E. In: A. Love, ed., Chromosome number reports LX XIII. Taxon 30: 829-
861. 1981. [C. sativa, 855, 2n = = 26.]
Baksay, L. Th d ical relations of some European
plant species. Ann. Hist.-Nat. ae Hungar. 8: 169-174. 1957. [C. microcarpa (2n =
40), C. Rumelica (n = 6), p. 172.]
BALSCHUN, H., & F. JAcos. on ns problem of the effect of Camelina species on flax
yield. (In German: English summary.) Flora 151: 572-606. 1961. [Yield of flax is
reduced by competition with Camelina.]
_ On the interspecific competition among Linum usitatissimum L. and
species of Camelina. (In German; English summary.) /bid. 161: 129-172. 1972.
[Evidence supporting the effects of competition and negating the role of allelopathy
on the reduction of growth in flax.]
Barrett, S. C. H. Crop mimicry in weeds. Econ. Bot. 37: 255-282. 1983. [Camelina,
263, 264.]
Bossier, E. Camelina. Fl. Orientalis 1: 311-313. 1867. [Recognized seven species in
two sections. ]
238 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Brooks, R. E. Jn: A. Léve, ed., Chromosome number reports LXXXVII. Taxon 34:
346- 351. 1985. [C. Rumelica, 347,n = 6.]
FritscH, K. Zur Kenntnis der eee oe Velenovsky. Sitz-ber. Akad, Wiss.
Math-Naturw. Wien 138: 347-370. 1929.*
GALLAND, N. In: A. LOveE, ed., Ouse. number reports LX XXYV. Taxon 33: 756-
Gus 1984. [C. sativa subsp. pilosa, 756, 2n = 26.
Grummer, G. Die Beeinflussung des Leinertrages durch Camelina-Arten. Flora 146:
158 177. 1958. [C. sativa, C. Alyssum, C. dentata
& H. Beyer. The influence exerted by species of Camelina on flax by means of
toxic substances. Pp. 153-157 in J. L. HARPER, ed., The biology of weeds. Oxford.
1960. [Allelopathy, influence of leaf phenolics of Camelina on growth and reduction
of yield in flax.]
Gunstong, F. D., & L. J. Morris. Fatty acids VI. The oxygenated acid present in
Camelina sativa (Crantz) seed oil. Jour. Chem. Soc. 1959: 2127-2132. 1959. [Epoxy-
fey acid.]
E, I. C. Camelina. In: P. me Davis, ed., Fl. Turkey 1: 490-493. 1965. [Excellent
accoun t; SIX species reco ]
Hutonen, I. Blick auf die nine. Arten Finnlands. Arch. Soc. Zool. Bot. Fenn. 1:
129-131. 1948.
Hyetmavist, H. The flax weeds and the origin of cultivated flax. Bot. Not. 1950: 257—
8. 1950. [Camelina, 258-261.]
IpARRA, F. E., & J. LA Porte. Las cruciferas del género Camelina adventicias en la
Argentina. (English summary.) Revista Argent. Agron. 14: 94-1 oe ra [Key,
eae chromosome numbers; C. sativa, C. microcarpa, C. Par
Kyaer, A., R. GMELIN, & R. B. JENSEN. Isothiocyanates XXI. (-)-10- Methloulphinyls
decyl isothiocyanate, a new mustard oil present as a glucoside (glucocamelinin) in
Camelina species. Acta Chem. Scand. 10: 1614-1619. 1956. [C. dentata, C. micro-
carpa, C. sativa. ]
Knorzer, K.-H. Evolution and spreading of gold of pleasure (Camelina sativa s.1.). (In
German; English summary.) Ber. Deutsch. Bot. Ges. 91: 187-195. 1978. [C. sativa,
C. microcarpa, C. Alyssum, C. pilosa; archeological findings, economic importance. ]
KRANZ, E., & F. JAcos. The competition of Linum with Camelina for minerals. I. The
uptake of **S-sulphate. (In German; English summary.) Flora 166: 491-503. 1977.
[Uptake of minerals by Camelina was higher than that of Linum; reduction of the
dry weight of Linum is caused by competition and not by allelopathy; for a related
paper on the uptake of phosphate and rubidium, see ibid. 505-516.
Lovett, J. V.,& A.M. DurrFietp. Allelochemicals of Camelina sativa. Jour. Appl. Ecol.
18: 283-290. 1981. [Aqueous washings of leaves of Camelina contain allelochem-
icals; benzylamine influences the association of Camelina and flax: role of bacteria
in the production of allelochemicals.]
H. F. Jackson. Allelopathic activity of Camelina sativa (L.) Crantz in relation
to its phyllosphere bacteria. New Phytol. 86: 273-277. 1980.
& B. E. Juniper. Electron microscopy of structures on the adaxial leaf surfaces
of Camelina Sativa and Spinacia oleracea. New Phytol. 81: 627, 628. pil. J. 1978.
[Crystalline structures probably associated with activity of the bacterium Entero-
bacter cloacae
. R. Sacar. Influence of bacteria in the phyllosphere of Camelina sativa
(L.) Crantz on germination of Linum usitatissimum. New Phytol. 81: 617-625.
1978. [Growth of radicle of flax is stimulated by leaf washings of Camelina in the
presence of bacteria.]
MAJsovskyY, J., ef al. Index of chromosome numbers of plea flora (part 3). Acta
Fac. Nat. Comen. Bot. 22: I- 20. 1974. [C. microcarpa, 4, 2n = 40.
ae K. Einiges tiber Camelina. Allg. Bot. Zeitschr. 15: 132, 133. 1909. [Nomenclature
C. Rumelica, C. microcarpa, C. sativa, and C. Alyssum.]
1987] AL-SHEHBAZ, ALYSSEAE 229
McGrecor, R. L. Camelina Rumelica, another weedy mustard established in North
America. Phytologia 55: 227, 228. 1984. [Distribution in Colorado, Kansas, Okla-
homa, Oregon, and Texas.]
Current status of the genus Camelina eames in the prairies and plains
of central North America. Contr. Univ. Kansas Herb. 15: 1-13. 1985. [Key, de-
scriptions, ae distributions of four species. ]
MEIKLE, R. D. melina. In: T. G. TutTtn et al, eds., Fl. Europaea 1: 315. 1964
MIREK, VA ae mae and nomenclature of Camelina pilosa auct. Acta Soc. Bot. Polon.
49: 553-561. 1980. [Numerical analysis of 14 characters in C. sativa and C. micro-
carpa, C. sativa var. Zingeri, var. nov., proposed to replace the ambiguous C. pilosa. ]
Genus Camelina in Poland—taxonomy, distribution and habitats. Frag. FI.
Geobot. 27: 445-507. 1981. [Numerical analysis of 37 characters in four species;
descriptions of two new series; habitats, key, descriptions, distributions, 32 figs., 10
tables. |
Monographic studies in genus Camelina Cr. 1. Camelina anomala Boiss. e
Hausskn. Acta Soc. Bot. Polon. 53: 429-432. 1984. [Morphology, distribution,
sectional ae of Camelina
MOouTERDE, P. uvelle flore du Liban et de la Syrie. Vol. 2. xii + 727 pp. Beirut.
1970. Seen 129, 130.]
PLessers, A. G. Beds. trials with oilseed plants. Il. Camelina. Canad. Jour. Pl. Sci.
2: 452-459. 1962.
Popiecu, D., & A. DieTERLE. Chromosomenstudien an afghanischen Pflanzen. Can-
dollea 24: 185-243. 1969. [C. Rumelica, 204, 2n = 40.
ScHwanitz, F. Die Evolution der Kulturpflanzen. xii + 463 pp. Munich, Basel, and
Vienna. 1967. [Camelina, 337-339.
SinskAJA, E.N. The oleiferous plants and root crops of the family Cruciferae. (In Russian;
English summary, 555-619.) Bull. Appl. Bot. 19(3): 1-648. 2 colored plates. 1928.
[Camelina, 535-554, figs. 102-108.
_BezTuzHEVA. The forms of Camelina sativa in connection with climate,
flax and man. (In Russian; English summary, 179-197.) Bull. one Bot. 25(2): 98-
200. 1931. [Six species; ecotypic variation, distributions, evolut n of Camelina
through selection of flax, effects of climatic and ghtosocological eae and of
threshing and winnowing, 7 figs., 8 tables, 30 map
SKALINSKA, M., E. PoGAN, R. Czapik, ef a/. Further aes in chromosome numbers
of Polish angiosperms, twelfth contribution. Acta Biol. Cracov. Bot. 21: 31-63.
1978. [Camelina, 35, 36.]
SMEJKAL, M. Revision der tschechoslowakischen Arten der Gattung Camelina Crantz
(Cruciferae). Preslia 43: 318-337. 1971. [Five species; key, descriptions, distribu-
tions, extensive synonymy.]
Srespins, G. L., Jk. Wariation and evolution in plants. xx + 643 pp. New York. 1950.
[Camelina, 123-134; excellent summary of the works of SINSKAJA & BEZTUZHEVA,
TepIN (1925), and ZiNGER.]
Strip, A., & R. FRANZEN. Jn: A. Love, ed., eee: number reports LX XII.
Taxon 30: 829-861. 1981. [C. Rumelica, 834, = 26.
Tepin, O. Zur Bliiten- a ee der Tendeie: (Camelina sativa). Bot.
Not. 1922: 177-189.
. The inheritance - oad leaves in Camelina. Hereditas 4: 59-64. 1923.
[Two genes with four alleles control leaf shape; double recessive genes control the
formation of entire ee
. Vererbung, Variation und Systematik in der Gattung Camelina. (English sum-
mary, 380-385.) Ibid. 6: 375 386. 1925. [Crosses between four “pure” lines yaa
in height, leaf shape, hairiness, fruit shape, and seed color and weight; 25 figs., 4
tables
—_. Zur Vererbung in der Gattung Camelina, eine Antwort. Ibid. 8: 359-362. 1927.
240 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
VASIL’CHENKO, I. T. Camelina. In: V. L. KomMARov & N. A. Buscu, eds., Fl. USSR 8:
596-602. 1939. [English translation by R. Lavoorrt, 8: 445-450. Jerusalem. 1970.]
VAUGHAN, J. G. The ee and utilization of oil seeds. xv + 279 pp. London. 1970.
[C. sativa, 62, fig.
ZINGER, H. B. On the ae of Camelina and S| |
of flax and their origin. (In Russian.) Trav. Mus. Bot. Acad. Sci. St.-Pétersb. 6: I-
303. pls. 1-9. 1909, a 1-234, pls.
1987] CHANNELL & WOOD, BUXACEAE 241
THE BUXACEAE IN THE SOUTHEASTERN
UNITED STATES!
R. B. CHANNELL? AND C. E. Woon, JR.?
BUXACEAE Dumortier, Comment. Bot. 54. 1822, nom. cons.
(BoxwooD FAMILY)
Monoecious [or dioecious, rarely perfect-flowered] evergreen shrubs, sub-
shrubs, or rhizomatous herbs [rarely trees], with entire or dentate, alternate or
opposite, exstipulate leaves, the hypogynous flowers actinomorphic, borne in
usually dense racemes, spikes, or heads, the staminate above the carpellate,
the latter rarely solitary, both usually subtended by | to several bracts and
bracteoles. Perianth of 4 [or 6] imbricate tepals in 2 pairs [or in whorls].
Androecium of 4 [or 6] stamens opposite the tepals, sometimes abate a
central nectary [or a rudimentary gynoecium], the distinct filaments n thick-
ened and bearing large anthers. Gynoecium [2- or] 3- canpenaie rn fa many
or] twice as many locules, each with [2 or] | (respecti
ovules, the carpels connate below, distinct above: srading at length into linear-
subulate style branches stigmatic along the inner surface, often becoming di-
vergent, divaricate or recurved in fruit. Fruit [a loculicidal capsule forcibly
ejecting the seeds (Buxus) or] apparently indehiscent but capsular, baccate {or
drupaceous], regularly failing to dehisce, disarticulating below, falling entire
and freeing the enclosed seeds from the base or by degeneration of fruit pulp.
Seeds dark brown or black, shining, sometimes conspicuously carunculate;
endosperm fleshy. (Buxacées Loiseleur, Man. PI. Us. Indig. 2: 495. 1819, nom.
inval.) TYPE GENUS: Buxus
‘Prepared for the Generic Flora of the Southeastern United States, a long-term project currently
made possible through the support of National Science Foundation grants BSR-841 $769 (Carroll E.
Wood, Jr., principal investigator) and BSR-8415637 (Norton G. Miller, principal investigator). The
116th in the series, this paper follows the format established in the first one (Jour. Arnold Arb. 39:
296-346. 1958) and continued to the present. The area covered by the Generic Flora includes North
was prepared by Dorothy H. Marsh, with Channell’s panies and eee from plants grown
We are indebted to Barbara Nimblett for her ve with the ene and the m t.
>Department of General Biology, Box 1501, Vanderbilt University, Nashville, ae 37203.
3Arnold Arboretum of Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 241-257. pees 1987.
242 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
An apparently ancient group, to judge from the diversity of the constituents
(including about 70 species of Buxus, seven or eight of Notobuxus Oliver, 11
of Sarcococca Lindley, and four of Pachysandra Michaux) and the widespread
intercontinental distribution, especially in the Old World tropics. The family
has been treated as a tribe of the Euphorbiaceae (Bentham; Bentham & Hooker),
or a “series” (Buxeae) of the Celastraceae (Baillon, 1875), or assigned variously
to the Celastrales (Scholz), the Euphorbiales (Takhtajan, 1959, 1969; Cronquist,
1968, 1981), the Hamamelidales (Takhtajan, 1954, 1980; Hutchinson, 1969,
1973), the Pittosporales (Thorne, 1976, 1983), and a separate order Buxales
in Dahlgren’s (1983) Rosiflorae. Airy Shaw (in Willis) thought the Buxaceae
to be related to the Euphorbiaceae and perhaps the Celastraceae.
The pollen of Pachysandra and Sarcococca is similar. The spheroidal grains
are radially symmetrical. In Pachysandra they are polyforate, with more than
12 apertures or foramina. They have a pattern characteristic of Croton L. and
other Euphorbiaceae, with well-defined, regularly arranged triangular excres-
cences of the sexine that form reticulate polygons. The pollen of Sarcococca
Conzattii (Standley) I. M. Johnston (S. guatemalensis 1. M. Johnston) is quite
different from that of the Asiatic species and that of Pachysandra (see Gray &
Sohma), strengthening Sealy’s (1986) exclusion of this species from Sarcococca.
The pollen of Buxus evidently in no way resembles that of either Pachysandra
or Sarcococca, being similar, however, to that of Styloceras Juss. (Stylocera-
laceae), a genus comprising four species of glabrous trees of western tropical
South America and related to Buxaceae but differing, according to Airy Shaw
(in Willis), in the naked staminate flowers with many more or less sessile anthers
borne on a solitary bract, and in the locules of the ovary that are completely
divided by secondary longitudinal septa (but cf. Pachysandra procumbens).
The staminate flowers of Sty/oceras appear to be simple androphylls, as in
Didymeles, a genus of two arborescent Madagascan endemics and the basis for
the Didymeleaceae, the fruits of which are large, one-seeded drupes (cf. the
berries of P. terminalis), the flowers evidently primitively simple, and the genus
possibly having some relationship to the Buxaceae through the Stylocerataceae.
It is of interest in passing that the distribution of the Buxaceae, the Styl
cerataceae, and the Didymeleaceae together includes the major continental
land masses of the world, suggesting that comparative knowledge of the alli-
ances may contribute to a refined appreciation of phytogeography and a better
understanding of the efficacy of isolation in phylogeny.
The family is notable for the alkaloids that occur in its members. Gibbs
noted, “steroid alkaloids in bewildering numbers occur in: Buxus, Pachysandra,
Sarcococca” (p. 1217) and “the steroid alkaloids of the Buxaceae certainly
define that family” (p. 1221).
The Buxaceae are economically important for various ornamentals used in
horticulture, principally Buxus (various species, especially B. sempervirens L.
and B. microphylla Sieb. & Zucc., selections, and cultivars, including those
variously known as English, Japanese, Korean, Chinese, and ‘California’ box
or boxwood), and to a lesser extent representatives of four or five (see Bailey
et al. and Sealy, 1949) species of Sarcococca (including S. ruscifolia Stapf, sweet
box, and S. saligna (D. Don) (Miill.-Arg., willow-leaf box) and Pachysandra
1987] CHANNELL & WOOD, BUXACEAE 243
terminalis Sieb. & Zucc., Japanese spurge or Japanese pachysandra, widely
used as a ground cover. The firm, close-grained wood of certain members of
Buxus (especially B. Serpe TVitens) is used for turning and engraving. Other
products also find use in commerce.
A useful liquid wax, difficult to synthesize commercially, is obtained from
the seeds of Simmondsia chinensis (Link) Schneider (S. californica Nutt.), the
jojoba or goatnut. Contrary to the connotation of the original epithet, this long-
lived, low shrub (under cultivation attaining a height of three feet when staked)
is indigenous to large areas of the Sonoran Desert of California, Arizona, and
northern Mexico. Although long assigned to the Buxaceae (Miller; Pax, 1890;
Hutchinson, 1967; Scholz), it constitutes a separate family taxonomically, the
immondsiaceae.* The seeds contain up to 25 percent liquid, unsaturated wax,
which can be solidified by hydrogenation, used in the manufacture of extreme-
high-pressure lubricants, especially for transmissions in heavy-duty vehicles.
It is a suitable substitute in uses calling for spermaceti (sperm-whale ‘‘oil,”
itself technically a wax), carnauba wax, and beeswax, and it is currently also
used in a number of cosmetics. Resistant to pests and diseases, jojoba thrives
without irrigation in areas where rainfall is less than 24 cm annually, with
mature plants producing up to 12 pounds dry weight of seeds during the period.
It has elicited considerable interest as a potential agricultural crop, not only in
desert regions of the North American Southwest, but in arid regions of Argen-
tina, Chile, Israel, Africa, and Australia.
The monogeneric Simmondsiaceae have flowers with (4) 5 (6) tepals, nu-
merous (8-12) stamens, unique pollen, and a solitary ovule in each of the three
locules (cf. Pachysandra), usually a single large seed with a large embryo and
little or no endosperm; and anomalous wood siEUctiite with several concentric
rings ue eae strands, besides other distincti 1 features. In pollen
and an Simmondsia has much in common with some members of the
acres although an actual relationship with them is difficult to en-
vision, according to Airy Shaw (in Willis), who suggested that the most probable
affinity of Simmondsia is with the Monimiaceae (sensu stricto), from which it
differs in the syncarpous gynoecium and fruit and the scanty or absent endo-
sperm. However, Wettstein (1924, 1935), Takhtajan (1969), Cronquist (1981),
Thorne (1983), and Dahlgren all have associated Simmondsia with the Eu-
phorbiales. Scogin (see references to Simmondsiaceae) found that in taxa from
a wide array of families examined for cross-reactivity with Simmondsia anti-
serum, a reaction was detected only with three species of Euphorbiaceae. It
‘The name Simmondsiaceae is usually attributed to Van Tieghem (Ann. Sci. Nat. VIII. 5: 289-
338. 1897, the 008 on mateo repeated almost verbatim in Jour. Bot. Morot 12: 103-112.
1898), | did he use the Latin form. He always referred to the family as ‘““Simmondsiacées.”
(And in none of his papers ee we find anything but the French vernacular form for family names,
and frequently for generic ones, e.g., “le genre Simmondsie.’”’) Not being in Latin, these names are
invalidly published. Insofar as we have been able to determine, the name Simmondsiaceae should
be cited as Takhtajan ex Dostal, Botanickéa Nomenklatura, 217. 1957. Airy Shaw’s citation (in Willis)
attributing the name to (Pax) Van Tieghem is certainly incorrect, for nowhere did Van Tieghem either
use the correct form of the family name or mentio
244 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
seems clear that the relationships of Simmondsia, whatever they may be, are
not with the Buxaceae as formerly thought.
Because of the economic interest of Simmondsia, a selected series of refer-
ences to this genus is included here following those for the Buxaceae.
REFERENCES TO BUXACEAE:
ALPHIN, T. H. A descriptive study of varietal forms in Buxus. Am. Jour. Bot. 27: 349—
357. pls. 1, 2. 1940.
BAILEY, L. H., E. Z. BAILEY, & STAFF OF L. H. BAILEY Hortorium. Hortus third. xxi +
1290 pp. New York and London. 1976. [Buxaceae, 192; Pachysandra, 809; Sar-
cococca, 1006; Simmondsia, 1046.]
Baton, H. Monographie des Buxacées et des Stylocerées. 89 pp. 3 pls. Paris, 1859.
[Buxacées, including Sarcococca, Pachysandra, Buxus sects. Eubuxus and Tricera,
71, 81-86; Stylocéracées, 72-81.
: Célastracées, Hist. Pl. 6: 1-50. 1875. [Série des Buis, 16-19; Buxeae, 47-49.
Includes Buxus, Pachysandra, Sarcococca, Simmondsia, Styloceras.] English transl.
by M. M. Hartoa, Celastraceae. Natural History of Plants 6: 1-51. 1880. [Box
series, “ 19: Heads 48-51.
BALpwin, J. T. Boxwood. Boxwood Bull. 14(1): 10-13. 1974. [Popular account of
uXUS.
Barase, D., Y. BERGERON, & G. A. VINCENT. The position of Daphniphyllaceae, Bux-
aceae, Simmondsiaceae and Cecropiaceae in the subclass Hamamelididae. A nu-
merical ae or t. Rend. Séances Acad. Sci. Sér. II. Sci. Vie 294: 891-893,
895, 896.
BENTHAM, G. me on Euphorbiaceae. Jour. Linn. Soc. Bot. 17: 183-267. 1878. [Dis-
agrees with BAILLON; Buxeae maintained as a tribe of Euphorbiaceae, 205, 206.]
J.D. Hooker. Euphorbiaceae. Tribus III. Buxeae. Gen. Pl. 3: 265-267. 1880.
[Simmondsia, Styloceras, Sarcococca, Buxus, Pachysandra.]
BOLKHOVSKIKH, Z., V. GRIF, T. MATVEJEVA, & AKHARYEVA. Chromosome numbers
of flowering plants. A. A. FepERov, ed. 928 pp. V. L. Komarov Bot. Inst., Acad.
Sci. USSR, Leningrad. 1969. [Buxaceae, 182.]
Cerny, V., & F. Sorm. Steroid alkaloids: alkaloids of Apocynaceae and Buxaceae. /n:
R. H. F. MAnskKe, ed., The alkaloids 9: 305-426. 1967.
CHENG, M., & T. L. Mina, eds. Angiospermae: Dicotyledoneae: Daphniphyllaceae,
Callitrichaceae, Buxaceae, Empetraceae, Coriariaceae, Anacardiaceae, Pentaphyla
caceae. (In Chinese.) Fl. Reipubl. Pop. Sinicae 45(1). vi + 152 pp. Beijing. 1980.
[Buxaceae, 16-60.
CRONQUIST, A. The evolution and classification of flowering plants. xii + 396 pp. Boston.
1968. Su phorbiales 257-260; Buxaceae, Euphorbiaceae, Daphniphyllaceae, Aex-
toxicaceae, Pandaceae. ]
ntegrated system of classification of flowering plants. xviii + 1262 pp. New
York. 1981, [Euphorbiales, 729-740; Buxaceae, Simmondsiaceae, Pandaceae, Eu-
phorbiaceae; Pachysandra terminalis and nine em illustrated. ]
DAHLGREN, R. General aspects of angiosperm evolution and macr osystematics. Nordic
Jour. Bot. 3: 119-149. 1983. [Simmondsiaceae placed in Euphorbiales in Malvi-
florae, Buxaceae in Buxales in Rosiflorae.]
DANG-VAN-LiEM. Embryogénie des Buxacées; développement de I’ seas chez le Bux-
us SEMIDETVIECHS L. Compt. Rend. Acad. Sci. Paris ek 1844-1847.
Davis, G. L. of glosp + 528 pp. New . London,
and Sydney. 1966. [Buxaceae, 65, 66.]
GAMBLE, M. A. The Edgar Anderson Balkan boxwoods. Boxwood Bull. 14(4): 57-63.
197
5.
Gentry, A. H. Family 99. Buxaceae. /n: R. E. Woopson, Jr., R. W. SCHERY, & COLLAB-
1987] CHANNELL & WOOD, BUXACEAE 245
oRATORS. Fl. Panama 6. Ann. Missouri Bot. Gard. 65: 5-8. 1978. [Buxus citrifolia
(Willd.) Sprengel, Panama and Venezuela; includes discussion of family.]
& R. Foster. A new Peruvian Sty/oceras (Buxaceae): discovery of a phytogeo-
graphical missing link. Ann. Missouri Bot. Gard. 68: 122-124. 1981. [S. Brokawii,
from lowland Amazonian Peru (Madre de Dios); illustrated.]
Gipss, R. D. Chemotaxonomy of flowering plants. 4 vols. (paged continuously). 2372
pp. Montreal. 1974. [Buxaceae, 2: 916, 1217-1219, 1221; Simmondsiaceae, 2: 1074,
1075.
GOLDBLATT, P. Taxonomy ofthe cultivated boxwoods, Buxus, Buxaceae. Boxwood Bull.
16(1): 12, 13. 1976. [Chromosome numbers.]
Gray, J., & K. Souma. Fossil Pachysandra from western America with a comparative
eae of pollen in Pachysandra and Sarcococca. Am. Jour. Sci. 262: 1159-1197.
map. 1964.
Hatusima, 8. A revision of the Asiatic Buxus. Jour. Dept. Agr. Kyushu Univ. 6: 261-
342. pls. 16-27. 1942. [26 species.]
HEGNAUER, R. Chemotaxonomie der Pflanzen. Band 3. Dicotyledoneae: Acanthaceae-
Cyrillaceae. 473 pp. Basel and Stuttgart. 1964. [Buxaceae (including Simmondsia),
BH my
Howarp, R. A. Notes on Buxus in the Lesser Antilles and on Mathou’s ee
publication. Jour. Arnold Arb. 44: 96-100. 1963. [Crantzia Sw., Tricera Sw., and
Buxus L.; Buxus species in ee Lesser Antilles; MATHOU’s monograph. 1
HUuTCHINSON, J. Buxaceac. Gen. Pl. 2: 105-109. 1967. [Stvloceras, Simmondsia,
Sarcococca, Buxus ( — Pachysandra, Austrobuxus Miq. (1861) =
Longetia nitida (Mia. ) Van Steenis (Euphorbiaceae), Reg. Veg. 34: 59. 1964.]
. Evolution and phylogeny of flowering plants. Dicotyledons: facts and theory.
xxvi + 717 pp. London and New York. 1969. [Buxaceae (“including Pachysandra-
ceae (1858). Stylocerataceae Baill. Simmondsiaceae van Tieghem (1898)’), 138-
141, in Hamamelidales, 132-142; Pachysandra axillaris, illustrated, 140, and genus
mapped, 141.]
. The families of flowering plants. ed. 3. xx + 968 pp. Oxford. 1973. [Buxaceae,
228, 229, in Hamamelidales.]
JouHNsTON, I. M. Some undescribed species from Mexico and Guatemala. Jour. Arnold
Arb. 19: 117-128. 1938. [Sarcococca guatemalensis I. M. Johnston, 121; thought
by Johnston to be the only New World species of the genus; but see SEALY (1986),
who excluded it, and Gray & SoHma, who noted its very different pollen.]
New phanerogams from Mexico. /bid. 20: 234-240. 1939. [Sarcococca guate-
malensis antedated by Buxus Conzattii Standley, described from Oaxaca, Mexico,
on the basis of incomplete material (fruit lacking); S. guwatemalensis = S. Conzattii
(Standley) I. M. Johnston, 240.]
KOu.eEr, E. Pollen types in the genus Buxus L. s.1., their geographical distribution and
Piatt for taxonomy (Buxaceae). Proc. Fourth Internat]. Palyn. Conf. 1: 264—
7. Lucknow. 1978.*
. Pollen nine of the West Indian—Central American species of the genus
Buxus L. (Buxaceae) with reference to taxonomy, (French summary.) Pollen Spores
23: 37-91. 1981. [Pollen of 37 species of Buxus examined by light and scanning-
electron microscopy. Eight pollen types and five major systematic groups recog-
d.
nize
Kupcuan, S. M., R. M. KeNNepy, W. R. SCHLEIGH, & G. OHTA. Buxus alkaloids. XII.
Benzamide eee from Buxus sempervirens L. Tetrahedron 23(12): 4563-4586.
1967
Martin, A. C. The comparative internal morphology of seeds. Am. Mid]. Nat. 36: 513-
660. 1946. [Buxaceae, 574, 575; Buxus microphylla and Pachysandra terminalis
illustrated; Simmondsia, 646.]
MarTIN-SANS, E. Généralité de la présence d’alcaloides chez les Buxacées. Compt. Rend.
246 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Acad. Sci. Paris 191: 625, 626. 1930. [Alkaloids in Buxus, Simmondsia, Pachysan-
dra, Sarcococca, and Styloceras.]
. PoNcHET. Sur l’appareil sécréteur des Buxus. Bull. Soc. Hist. Nat. Toulouse
60: 231, 232. 1930.*
MatTuHou, T. Recherches sur la famille des Buxacées; étude anatomique, microchimique
et systématique. Thése Fac. Sci. Toulouse Doc. Sci. Nat. 449 pp. pls. 28-33. (pls.
MaAuvriTzon, J. Kritik von J. Wigers Abhandlung ‘““Embryological studies on the families
Buxaceae, Meliaceae, Simaroubaceae and Burseraceae.” Bot. Not. 1935: 490-502.
1935
MELIKIAN, A. P. On the position of the families Buxaceae and Simmondsiaceae in the
system. (In Russian; English summary.) Bot. Zhur. 53: 1043-1047. 1968. [On the
basis of anatomical study of 12 species in four genera, concludes that Simmondsi-
aceae should be separated from Buxaceae.
MULLER, J. Buxaceae. DC. Prodromus 16(1): 7-23. 1869. [Tribes Buxeae (Styloceras,
Sarcococca, Buxus, Pachysandra) and Simmondsieae (Simmondsia).]
Naumova, T. N. Nucellar polyembryony in Sarcococca (Buxaceae). (In Russian.) Bot.
Zhur. a 230-240. 1980. [S. humilis Hort. and S. Hookerana Baillon.]
he embryology of the representatives of the family Buxaceae. (In Russian;
English summary.) /bid. 66: 1135-1145. 1981. [Buxus sempervirens, B. colchica, B.
balearica, Notobuxus acuminata, Sarcococca.]
— zky, F. Anatomie der Angiospermen-Samen. Handb. Pflanzenanat. II. Arche-
n. 10. vi + 365 pp. 1926. ee 189, 190.]
oa. M. Y. Polyembryony in Sarcococca ruscifolia, Stapf. Notes Bot. Gard. Edinburgh
14: 21-23. 1923. [Up to seven embryos ina single seed; position of embryos suggested
a nucellar origin
Pax, F. Buxaceae. Nat. Pflanzenfam. III. 5: 130-135. 1890. [Tribes Buxeae (Sarcococca,
Pachysandra, Buxus), Stylocereae (Notobuxus, Styloceras), Simmondsieae (Sim-
mondsia). Nachtr. I-IV: 213. 1897.
. Buxaceae KI. et Gcke. Pflanzenareale 1: 82. map 70. 1927. [Map showing the
worldwide distribution of the family; range of Pachysandra in the eastern United
States inaccurate. ]
RADCLIFFE-SMITH, A. A remarkable new species of Notobuxus (Buxaceae) from Tan-
zania. Kew Bull. 36: 39-41. 1981.
Recorp, S. J. Boxwoods of commerce. Bull. Torrey Bot. Club 47: 297-306. 1922.
& G. A. GARRATT. Boxwoods. Yale School Forestry Bull. 14: 1-81. pls. J-4.
Rupert, E. A., & G. L. Wesster. A procedure for staining pollen nuclei when obscured
by cytoplasmic a Stain Technol. 47(4): 185-187. 1972. [Euphorbiaceae,
Malvaceae, Buxaceae.
Scuoiz, H. Reihe ine ie Pp. 289-300 in H. Metcnior, A. Engler’s Syllabus der
Pflanzenfamilien. ed. 12. Vol. 2. [viii +] 666 pp. Berlin. 1964. [Buxaceae in Un-
terreihe Buxineae, 297, 298.
SEALY, J. R. Species of Sarcococca in cultivation. Jour. Roy. Hort. Soc. 74: 301-306.
fig. 108. 1949. [S. Hookerana var. digyna, S. saligna, S. humilis, S. confusa, S.
rusclfolia vars. ruscifolia and chinensis. ]
. A revision of the genus Sarcococca (Buxaceae). Bot. Jour. Linn. Soc. 92: 117-
159. 1986. [Eleven aa including one with three varieties, two with two varicties
each, another with tw :
SimoneT, M., & C. i eauaaieers Etud i é I t
ou sarmenteuses d’ormement. Compt. Rend. ‘Soc. Biol. Paris 111: 969. 1932. [Chro-
mosomes of Buxus and Sarcococca; x =
TAKHTAJAN, A. Proiskhozhdenie pokrytosemennykh rastenil. Soviet Sciences Press,
1987] CHANNELL & WOOD, BUXACEAE 247
Moscow. 1954. English translation by O. H. GANKIN. Origins of angiospermous
plants. G. L. Sressins, ed. 68 pp. AIBS, Washington, D. C. 1958. [Buxaceae and
Simmondsiaceae in order SappuNenin 59.
. Die Evolution der Angiospermen. vili + 344 pp. Jena. 1959. [Simmondsiaceae
questionably assigned to eras 197, 198; Buxaceae in Euphorbiales, 215,
216.
. Flowering plants: origin and dispersal. Authorized translation [of The origin of
angiospermous plants. ed. 2. Moscow. 1961] from Russian by C. JEFFREY. x + 310
pp. Edinburgh and Washington, D. C. 1969. ‘Euphorbiales, including Buxaceae
(including Stylocerataceae), Simmondsiaceae (Simmondsia), Daphniphyllaceae, Eu-
phorbiaceae, Dichapetalaceae, Pandaceae, Picrodendraceae, 221.]
. Outline of the classification of flowering plants (Magnoliophyta). Bot. Rev. 46:
225-359. 1980. [Order Hamamelidales, suborder Buxineae, Buxaceae and Sim-
mondsiaceae, 265, 266, 350.]
THorn_e, R. F. A phylogenetic classification of the Angiospermae. Evol. Biol. 9: 35-
106. 1976. [Buxaceae in order Pittosporales, suborder Buxineae; Simmondsia in
Incertae sedis.]
d new realignments in the angiosperms. Nordic Jour. Bot. 3: 85-117.
1983. [Simmondsiaceae in Euphorbiales between Euphorbiaceae and Thymelae-
aceae; Buxaceae in Pittosporales, — Buxineae, including Buxaceae, Daphni-
phyllaceae, Didymelaceae, Balanopac
TIEGHEM, P. vAN. Sur les Buxacées. Ann. Sci Nat. VIII. 5: 289-338. 1897. [Morpho-
logical and anatomical; Simmondsiacées recognized as a distinct family (but name
never given in Latin form, hence invalidly published). Buxacées divided into tribes
uxées and Pachysandrées. ]
UNDERHILL, T. L. The genus Sarcococca. Pl. Propag. 21(2): 4, 5. 1975.
VASILEVSKAYA, V. A., & G. M. BorisovskaAya. Life forms and their evolutional trans-
formations in the Buxaceae Dum. (In Russian English summary.) Trudy Mosk.
Obshch. Ispyt. Prir. Biol. 56: 9-104. 1981.
WETTSTEIN, R. Handbuch der systematischen Botanik. ed. 3. viii + 1081 pp. Leipzig
and Vienna. 1924. [Reihe Tricoccae, including on ee Dichapetalaceae,
Buxaceae, Callitrichaceae, 591-599.] ed. 4. x + 1152 pp. Leipzig and Vienna. 1935.
[Reihe Tricoccae, including the same families plus Daphnighyil eee 672-684. ]
Wicer, J. Ein neuer Fall von autonomer Nuzellarembryonie. Bot. Not. 1930: 368-370.
1930. [Nucellar polyembryony in Sarcococca pruniformis Lindley.]
ryological studies on the families Buxaceae, Meliaceae, Simarubaceae and
nee Thesis. Printed by H. Ohlsson, Lund. 1935.* [Buxaceae, 5-38.]
. Reply to remarks on my paper on Buxaceae, Meliaceae, etc. Bot. Not. 1936:
585-589. 1936. [See MAURITZON. ]
WILLAMAN, J. J.. & H. L. Lit. Alkaloid-bearing plants and their contained alkaloids,
1957-1968. Lloydia 33(suppl. 3A). 286 pp. 1970. [Buxaceae, 64-66; Buxus, Pachy-
ees Sarcococca, Simmondsia.
B. G. ScHuBERT. Alkaloid-bearing plants and their contained alkaloids. U. S.
Dep. Agr. Agr. Res. Serv. Tech. Bull. 1234. 287 pp. 1961. [Buxaceae, 56, 57,
including species of Buxus, ae Sarcococca, Simmondsia, and Styloceras. ]
Witus, J.C. A dictionary of the flowering plants and ferns. ed. 7. Revised by H. K.
Airy SHAW. xxii + 1212 + lili pp. Cambridge, England. 1966. ees ak 169;
Simmondsiaceae, 1040. Buxaceae i
showing relationships with Euphorbiaceae and perhaps Celastraceae.” "Re Sim-
mondsiaceae, ‘““The most p prooarle | affinity would seem to be with Monimiaceae
(s. str.), from which Sim principally in its syncarpous gynoecium and
fruit, and in the scanty scent |
248 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
REFERENCES TO SIMMONDSIA AND SIMMONDSIACEAE:
Under references to Buxaceae above, see BAILEY et al.; BAILLON, 1875; BARABE, BER-
GERON, & VINCENT; BENTHAM & HOOKER; CRONQUIST, 1981; DAHLGREN; GIBBS;
lea gnet Hutcuinson, 1967, 1969; MARTIN; MARTIN-SANS; Mera MULLER; PAX,
1890, 1927; TAKHTAJAN, 1959, 1969, 1980: . 1976, 1983: VAN TIEGHEM; WIL-
LAMAN & Lr, WILLAMAN & SCHUBERT; and WILLIS. Also see Jojoba Happenings, a pe-
ee published by the Office of Arid Lands Studies, University of Arizona, Tucson,
since 1972.
AL-ANI, H. A., B. R. Strain, & H. A. Mooney. The physiological ecology of diverse
populations i. desert shrub Simmondsia chinensis. Jour. Ecol. 60: 41-57. 1972.
[Dot map.
ALCARAZ, M. L. Air layering method for vegetative propagation of jojoba (Simmondsia
chinensis). Environ. Sci. Res. 23: 435-437. 1982.]
BaiLey, D. C. Anomalous ie and vegetative anatomy of Simmondsia chinensis.
Am. Jour. Bot. 67: 147-161. 0.
Brown, L. High hopes for the a. Can this shrub save the sperm whale and bring
sane agriculture to the Southwest? Horticulture 57(1): 34-36, 38, 39. 1979.
CONSEJO NACIONAL DE CIENCIA Y TECHNOLOGIA. Memorias de la II Conferencia Inter-
México, 10 al 12 de febrero de 1976. (In Spanish and English.) 338 pp. México,
F *
DauGHERTY, P. M., H. H. Sincatu, & T. A. WASTLER. Industrial raw materials of plant
origin. IV. A survey of Simmondsia chinensis (jojoba). Econ. Bot. 12: 296-304.
1958.
Gentry, H. S. Apomixis in black pepper and jojoba? Jour. Hered. 46: 8. 1955.
. The natural history of jojoba (Simmondsia chinensis) and its cultural aspects.
Econ. Bot. 12: 261-295. 1958.
HopceE, W. H. Jojoba—an overlooked ornamental shrub of the arid Southwest. Am.
Hort. Mag. 40: 346, 347. 1961. [S. chinensis.]
MauGu, T. H. Guayule and jojoba’s agriculture in semiarid regions. Science 196: 1189,
1190. 1977
NATIONAL ACADEMY OF ScIENCE. Underexploited tropical plants with promising eco-
nomic value. x + 188 pp. Washington, D. C. 1975. [Jojoba (Simmondsia), including
description, oil, culture, limitations and special requirements, research needs, and
selected readings, 105-110.
Rost, T. L., A. D. Stmper, P. SCHELL, & S. ALLEN. Anatomy of jojoba Selle
chinensis) seed and the ce of iquid wax during germination. Econ. Bot
140-147. 1977
ScHMID, R. Floral a fruit anatomy of jojoba (Simmondsia chinensis). Boxwood Bull.
21(2): 25-28. 19
Scoam, R. ere ee of Simmondsia chinensis (Simmondsiaceae). Aliso 9: 555-
559. 1980. [A wide spectrum of families tested; cross reaction detected only with
three species of Euphorbiaceae. ]
. Brown. Leaf flavonoids of Simmondsia chinensis ee Aliso
9: 475-477. 1979. [One well-characterized and three previously unreported iso-
rhamnetin glycosides * ‘of little utility as a systematic discriminant see of their
a
n
SHERBROOKE, W. C., & E. F. HAASE. Jojoba: a wax-producing shrub of the Sonoran
Desert; literature review and annotated bibliography. iv + 141 pp. map. Arid Lands
Resource Information Paper No. 5. Univ. Arizona, Office of Arid Lands Studies.
Tucson. 1974.*
TIEGHEM, P. VAN. Sur le genre Simmondsie considéré comme type d’une famille dis-
tincte, les Simmondsiacées. Jour. Bot. Morot 12: 103-112. 1898. [A repetition of
1987] CHANNELL & WOOD, BUXACEAE 249
the section on Simmondsia in his paper “Sur les Buxacées” published in 1897 (see
references under Buxaceae and footnote 4); Simmondsiaceae not validly published.]
VASUDEVA Rao, P. H. V., & E. R. R. IYENGAR. Studies in seed morphology and ger-
mination in jojoba (Simmondsia chinensis Link). Curr. Sci. Bangalore 51: 516-519.
1982.*
WALLACE, C. S., & P. W. RUNDELL. Sexual dimorphism and resource allocation in male
and female shrubs of Simmondsia chinensis. Oecologia 44: 34-39. 1979.
YERMANOS, D. M. Agronomic survey of jojoba in California. Econ. Bot. 28: 160-174.
1974.
1. Pachysandra Michaux, Fl. Bor.-Am. 2: 177, 178. pi. 45. 1803.
Evergreen or semi-evergreen, erect, decumbent, or prostrate, sympodial
[shrubs, subshrubs, or] perennial herbs, usually with [woody or] fleshy rhi-
zomes, fibrous roots, and simple, alternate, exstipulate, petiolate leaves with
glabrous, glabrescent, or pubescent, variously toothed, subdentate to nearly
entire blades with prominently 3-nerved pinnate venation. Inflorescences spi-
cate, basal [or axillary or terminal], the distal portions occupied by 5—40 ped-
icellate, subsessile, or sessile carpellate flowers. Staminate flowers subtended
by a single ciliate-pubescent bract, and with a perianth of 4 decussate, imbricate,
ciliate tepals sometimes with accompanying bracteoles, and (2 or) 4 (or 6)
distinct stamens with long-exserted, thickened or compressed clavate filaments,
each surmounted by a linear-oblong, rotund to sagittate, dorsifixed, longitu-
dinally dehiscent, introrse anther, the connective sometimes prolonged as an
appendage; pollen spheroidal, polyforate, with polygonal ornamentation. Car-
pellate flowers inserted on the inflorescence axis below the staminate flowers,
subtended by 7-13 distinct imbricate herbaceous bracts, with 4 or more acute
tepals; ovary [2- or] 3-carpellaie, the carpels connate below, each with 2 locules
separated by a false partition, each locule then containing a single pendent
ovule; the styles [2 or] 3, subulate to linear, erect or spreading at anthesis,
becoming recurved in fruit; stigma linear or linear-lanceolate, papillose, usually
sulcate, covering the inner surface of the style branches. Fruit capsular [or
baccate], indehiscent but becoming detached basally and falling entire. Seeds
trigonal, with [or sometimes without] a micropylar caruncle, the smooth, glossy
testa finally hard and dry, dark brown or black, the endosperm whitish and
oily, the embryo straight, the cotyledons considerably broader than the radicle.
TYPE SPECIES: P. procumbens Michaux. (Name from Greek pachys, thick, and
andros, of a man, alluding to the thick filaments of the stamens.)
The genus includes four species: Pachysandra procumbens, Allegheny spurge
or Allegheny-Mountain spurge, indigenous to the southeastern United States,
and three indigenous to eastern Asia. Pachysandra terminalis Sieb. & Zucc.
(China and Japan) and P. axillaris Franchet (Yunnan, China), as their epithets
connote, are well marked and, together with P. procumbens, easily distinguished
by the disposition of the inflorescences; P. stylosa Dunn (China) is characterized
by its long, prominent styles, recurved in fruit. Treated by Robbins (1962) on
the basis of herbarium material as comprising six taxonomic varieties, the last
species deserves reexamination on the basis of more and better material. Cheng
has recently (1980) treated P. stylosa as a variety of P. axillaris.
250 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
The geographic distribution of the genus, with a single species endemic to
the eastern United States disjunct from the four of eastern Asia, exemplifies
the well-known affinities of the floras of the two regions pointed out by Asa
Gray in 1840 and elaborated by him in 1846. Fossil evidence of the occurrence
of Pachysandra in the western United States might be thought to lend credence
to the belief that the genus was a member of the northern temperate ‘“‘Arcto-
Tertiary” flora, but Leopold & Macgintie pointed out that Pachysandra “‘ap-
pears to have had ancestral types at middle latitudes in America well before
the ‘Arcto-Tertiary’ flora came into being.” They further suggested that the
‘“‘Pachysandra-Sarcococca group may well have had a New World origin.” The
center of morphological and taxonomic diversity, however, is clearly in eastern
Asia.
Gray & Sohma have shown that Sarcococca and Pachysandra have distinc-
tive and related pollen structure. Although the pollen morphology of the two
genera merges when they are viewed as a whole, some types are distinctive—
for example, the pollen of P. procumbens has a sculpture pattern and a pore
frequency that set it off from that of other species of Pachysandra and Sar-
cococca. Muller noted that Pachysandra-type pollen is known from the Cam-
panian of Canada and from the Campanian-Danian interval of Germany. Later
Cretaceous records of the group come from deposits of Maestrichtian age in
Wyoming (Leopold & Macgintie), California, Montana, and Canada (see Mul-
ler). In the eastern United States pollen of the P. procumbens type 1s known
from the middle Eocene and from the Miocene (Leopold & Macgintie). ““Most
of the ooo western records of the group appear to be related to Old World
specie
Believed to have survived the geologic changes of the past few million years
in the limestone plateau country of central Kentucky, Tennessee, and adjoining
states, Pachysandra procumbens 1s now of local occurrence, for the most part
in rich woods of moist ravines near streams.
Braun considered Pachysandra procumbens to be a characteristic herbaceous
plant of the Western Mesophytic Forest Region, an area having as its eastern
boundary the western escarpment of the Cumberland and Allegheny plateaus
and as its western boundary the loess bluffs of the Mississippi River. She
remarked that, although commonly thought to be rare, it is an abundant plant
of mesophytic woods in the region.
Pachysandra procumbens is of some floristic, phytogeographical, and wild-
multaneously replaced by new growth from scaly basal buds in spring. (As
might be expected, anomalous flowers with two or four styles instead of three,
and five stamens instead of four, have been described.) Old reports of the
occurrence of the plant in West Virginia and New Jersey, as well as at Memphis,
Tennessee, are probably erroneous but are of considerable interest if verified.
The species has been reported from central Kentucky, central and eastern
Tennessee, western North Carolina, western Georgia, Alabama, Mississippi,
the Marianna Caverns in Jackson County, Florida, and the Tunica Hills of
1987] CHANNELL & WOOD, BUXACEAE 251
Figure |. Pachysandra. a-h, P. a a, portion of plant with fruit, x '4; b,
inflorescence, carpellate flowers below the staminate, x 1/2; c, staminate flower
stamen, x 4; e, carpellate flower, x 2; f, carpel, re ee portion in longitudinal section
to show the single ovule suspended in each of the 2 locules, x 4; g, semidiagrammatic
cross section ety ovary to show 6 locules, each with a single ovule, x 6; h, mature
ruit, x 1; 1, seed, x
Louisiana. Its distribution outlines the dissected portion of the Highland Rim
Province surrounding the Central Basin of Tennessee, but it is not known to
occur on outlying portions of the Rim isolated within the Basin.
Robbins (1962) has reported results of investigations of embryology and life
history in Pachysandra procumbens. Microspore mother cells undergo the first
meiotic division, giving rise to two dyad nuclei without cytokinesis. The second
division, followed by simultaneous furrowing, results in a tetrahedral arrange-
ment of the pollen tetrad. Excrescences of the sexine, arranged as reticulate
polygons, develop after the microspores are released from the mother-cell wall.
(See J. Gray and J. Gray & Sohma for illustrations of pollen.)
The ovary is divided into six locules by three true and three false partitions.
Early in development the ovary is three-locular, each locule enclosing two
ovules. Later it becomes six-locular by formation between the ovules of sec-
ondary partitions that are thinner than the original septa (see FiGure 1). Dif-
ferentiation of the ovules and the appearance of the archesporial cells occur in
late June of the year prior to anthesis, which happens as early as February of
the following year.
The anatropous ovule has two integuments. The archesporial cell, differ-
entiating from a hypodermal cell of the nucellus, divides, forming a primary
parietal cell and a primary sporogenous cell, the latter, pushed into the nucellus,
functioning as the megaspore mother cell. The primary sporogenous cell is
separated from the nucellar epidermis by several layers of cells, a characteristic
of crassinucellate ovules
Megagametophyte (embryo sac) development follows the Polygonum or nor-
mal course, being of the monosporic, eight-nucleate type. Pollen tubes make
a2 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
their way between the stigmatic papillae down through the tissues lining the
stylar canal, passing between the cells rather than penetrating them. Before
fertilization a protuberance (obturator) develops by proliferation of placental
cells immediately dorsal to the ovule. Growing downward, it meets the outer
integument, which has grown over and enclosed the ovule, and continues to
develop until it forms a hood over the nucellus, possibly functioning in the
penetration of the pollen tube into the ovule. Proliferation of the outer integ-
ument in the micropylar region results in the formation ofa prominent caruncle,
a conspicuous feature of the mature seed. All of the nucellar tissue, except the
epidermis and groups of cells at the chalazal and micropylar ends of the seed,
disappears with development of the cellular-type endosperm, which constitutes
the bulk of the seed.
The trigonal, black, shining, carunculate seeds are shed in July and August,
remain dormant through the following fall and winter, and germinate in late
March and early April in close proximity to the parent plants, beneath leaf
litter, usually in direct contact with moist mineral soil. Rupture of the seed
coat occurs as the rapidly growing primary root emerges through the caruncle.
Eventually the seed coat is shed, the cotyledons and epicotyl still enclosed
within the surrounding fleshy endosperm. The cotyledons and epicotyl even-
tually emerge, with only remnants of the endosperm then being evident.
The cotyledons, which persist for as long as a year, have thick, glossy, green
blades and short petioles, the latter undergoing elongation as development
proceeds. Within six weeks of germination, the minute epicotyl develops into
a short aerial stem bearing three to five small foliage leaves. Growth is slow,
the shoot attaining a height of only five inches by the end of the first year.
Secondary aerial shoots develop from the base of the initial shoot at the point
of attachment of the cotyledons. The rapid growth of these shoots results in
the establishment of lateral branches that soon overtop the primary axis. Pro-
tuberances that develop along with the lateral aerial branches are evidently the
source of the rhizomes and adventitious roots. Eventually the seedling develops
a sympodially branched rhizome system. The flowering of seedlings was not
observed, although their development was followed for three years.
Established clones of Pachysandra procumbens have a well-developed sym-
podial system of rhizomes terminated by decumbent aerial shoots surmounted
by a cluster of approximate leaves, diminishing in size distally and mottled
pale green (if not silver) on dark green in late fall and winter. Upon excavation
clones with as many as 38 aerial shoots were found to be interconnected by
underground rhizomes. Each aerial shoot dies as the result of abscission, which
occurs in spring as a new vegetative bud gives rise to a replacement shoot. A
lateral vegetative bud, already established at the base of the old shoot stub,
now rapidly develops into a new aerial shoot. This process, repeated succes-
sively year after year, results in the prominent and characteristic sympodium,
one actual analysis of which estimated the age at 34 years! “‘Dichotomous”’
branching of the axis occurs when two lateral buds of a single segment develop
into leafy shoots. Indeed, the typical circular growth habit of well-established
clones of considerable age is attributable to the repetition of such branching.
1987] CHANNELL & WOOD, BUXACEAE 20
It seems plausible, therefore, that an entire “population” may in fact represent
a single clone, having developed from one plant by repeated sympodial growth
and dichotomous branching, followed by subsequent fragmentation or degen-
eration of older portions of the rhizome system
By early May or June of the year prior to the spring in which a given aerial
shoot abscises, not only is a lateral shoot bud developed but so also are one
to three flower buds. These occupy a lateral position, well below the point of
shoot abscission. Anthesis occurs as early as mid-February in the vicinity of
Nashville, Tennessee, but usually during the last week of March and early
April. Either the staminate or the carpellate flowers may open first. In some
instances the staminate flowers will have fallen before the carpellate ones open.
There appears to be no single, regular, progressive order of events with respect
to the details of flowering—probably the differential effect of short-term en-
vironmental influences upon the preformed flower parts. The conspicuous white
staminal filaments elongate rapidly, well overtopping the sepals. The originally
erect style branches diverge, curve outward, and expose the inner stigmatic
surface, which is covered with minute papillae
The whitish pollen grains are exposed by longitudinal splitting and slight
recurving of the anther walls. A heavy “rain” of pollen onto the carpellate
flowers below commonly occurs, with a glistening appearance of the stigmatic
surface presumably indicating receptivity. Soon after anthesis the anthers fall,
carrying with them adherent pollen and thus possibly providing a second op-
portunity for pollination.
During anthesis the staminate flowers emit a rather penetrating odor faintly
resembling that of carnations or, to some people, the essence of ammonia or
of an amine. It has been described as being pleasantly fragrant at the outset,
later becoming sharp and penetrating. Insects, including beetles and bees, have
been reported to visit the staminate flowers but never the carpellate ones. Red
spiders (mites) covered with pollen have been observed on and in both types
of flowers. There is no question but that the staminate flowers are structurally
equipped to attract insects and are effective in doing so. In addition to the
attractant features of the conspicuous white filaments and the abundant whitish
pollen, the existence of a central nectary in the staminate flowers would appear
to be especially significant. Evidence that self-pollination occurs is unques-
tionable. That insects provide a medium for cross-pollination seems not only
possible but probable.
The plant is hardy well to the north of its native range and is sometimes
grown as an ornamental novelty in partial shade, where it spreads slowly.
Isolated clones in cultivation—indeed, those in nature— behave as though they
may be genetically self-sterile, apparently never setting seeds. That fruits are
seldom seen in nature 1s De to reflect actual absence, as opposed to faulty,
casual, or cursory observation.
In the self-pollination oe meats of Robbins (1962), 50 plants, collected
from five different populations, were used. These bore a total of 288 carpellate
flowers, representing a potential of 1368 seeds (on the basis of six seeds per
fruit). Of the 228 flowers self pollinated, only 104 set fruit, with a potential of
254 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
624 seeds. However, only 17 seeds were actually produced, and these by only
three plants. Ten fruits contained only one seed each, two contained two seeds
each, and only one contained three.
The crossing experiments made use of 43 plants with a total of 191 carpellate
flowers, representing a potential of 191 fruits and 1146 seeds. Only 57 (30
percent) of the flowers set fruit, and only six of these produced seeds, the total
number of seeds being 12.
Of the 423 carpellate flowers on 40 plants used in the initial apomixis test
in which pollen was withheld, 87 (21 percent) set fruit, but none produced
seeds. In a “replication” of the experiment the following spring, 98 carpellate
flowers on 28 plants were observed. None of these produced fruits. In a third
replication 95 carpellate flowers on 13 plants were observed, but again no fruits
were set. In all experiments 248 fruits out of a potential of 1991 were produced.
The total number of seeds produced was only 29, these being recovered from
only nine fruits.
The experiments indicate that the species is both self- and cross-compatible.
The possibility of apomixis being involved, although unlikely, cannot be ruled
out entirely on the basis of the negative results obtained. The general paucity
of fruits in nature tends to mitigate against apomixis, however. Pollination
could be shown to be a stimulus to apomictic development, for in most pseu-
dogamous species the embryo begins development autonomously, but the en-
dosperm will not develop unless it is fertilized.
The overall results of these experiments further emphasize previous obser-
vations that development of fruits in Pachysandra procumbens is sporadic and
that seed production 1s generally rare. This implies that the species may indeed
have a very low degree of sexual fertility. Vegetative propagation has assumed
a major role, with reproduction involving seeds occurring only rarely. As pre-
viously pointed out, the possibility exists that an entire population occupying
the slopes of a ravine could represent a single clone, having originated vege-
tatively from a single plant.
The combination of rhizomatous habit, morphological constancy, restricted
habitat, and low ee fertility indicates that Pachysandra procumbens is a
nonaggressive 1f not “senile” species with a very low evolutionary potential.
Like other persistent Sent of low sexual capacity inhabiting ecologically
closed communities, P. procumbens no doubt benefits from either a sustained
low or a sporadic incidence of sexual output. Whether or not genetic self-
incompatibility operates between and among clones is not known, although
the crossing results suggest the existence of such a possibility.
Of the species of Pachysandra, P. terminalis is clearly the most important
economically, being widely used in horticulture as an ornamental ground cover
since its introduction into the United States in the 1800's. It is known in the
trade by the somewhat contrived name Japanese spurge. Its glossy evergreen
leaves, low, creeping growth habit, and tolerance of shade make it an attractive
subject for ground-cover use. It is propagated vegetatively. It is unique in having
terminal inflorescences and two-carpellate, white, baccate fruits (cf. Didymeles),
described as about the size of a ‘Delaware’ grape, the pulp decidedly sweet.
Other distinctive characters include the elevated veins of the adaxial surface
1987] CHANNELL & WOOD, BUXACEAE 250
of the leaf blades, the comparatively small stigmatic area occupying only the
distal one-third of the style branches, the presence of a coriaceous bract and
two bracteoles subtending each staminate flower, and the somewhat elongated
pedicel of the carpellate flowers.
Although Pach) terminalis in cultivation is subject to attack by various
insect pests and fungus diseases, it is in general resistant to them. Dodge (1944a)
reported that canker blight or leaf-spot disease of the plant is due to the fungus
Volutella pachysandricola. He also noted susceptibility to fungi of the genera
Phyelosticta and Glocosporum and to attack by the scale insect Chionaspis
evonyml.
While the leaves of Chinese plants of Pachysandra terminalis are reportedly
somewhat smaller than those of the Japanese ones, no taxonomic significance
has yet been attached to the difference. Variegated selections with ivory-white
areas confined to the leaf margins have been described and are extant in the
horticultural trade.
Pachysandra stylosa Dunn var. glaberrima Hand.-Mazz. also finds limited
use in ornamental oe mainly as a ground-cover subject, being similar
in gross aspect to the preced
Horticultural use of eee axillaris and the varieties of P. stylosa is
rarely, if ever, encountered, although individuals of these species are occa-
sionally grown for exhibition in botanical gardens. These plants for the most
part present a more nearly woody, even shrubby habit of growth and have
thicker, more coriaceous leaves than do the other two. It would appear that
they deserve greater attention horticulturally.
REFERENCES:
Under references to Buxaceae, see BAILEY et BaILLon, 1859, 1875; BENTHAM &
Hooker; J. Gray & SOHMA; HUTCHINSON, 1967, 69: MARTIN; MARTIN-SANS; MATHOU
MULLER; PAx, 1890, 1927; VAN TIEGHEM; = eros) ene and WILLAMAN & SCHUBE a
BrAuNn, E. L. Deciduous forests of eastern North America. xiv + 596 pp. map. New
York. 1950. [P. procumbens, 124, 139, 157, 301, 488.]
aces E. S. Pachysandra Sioeunabeny. Am. Garden 11: 346. 1890. [A popular ac-
unt.]
Grateeey L. Morphologische und biologische Mittheilungen. 4. Ueber den Frucht-
knoten von Pachysandra procumbens Michx. Ost. Bot. Zeitschr. 43: 317. pl. 14, fig.
iy nes [Gynoecium of three carpels but six locules by development of false
partitions. ]
CHANG, T. T. Pollen eta of Hamamelidaceae and Altingiaceae. (In Russian;
English summary.) Acta Inst. Bot. Acad. Sci. URSS. 1. Fl. Syst. Pl. Vasc. 13: 173-
232. pls. I-17. 1964. saree 220.]
CHENG, M. New taxa of Buxaceae from China with discussions on some species. (In
Chinese; Latin diagnoses.) Acta Phytotax. Sinica 17(3): 97-103. 1979. [Buxus, Sar-
cococca, Pachysandra; includes P. stylosa Dunn, P. axillaris var. tricar,
P. axillaris Franch. var. stylosa (Dunn) M. Cheng. Fl. Reipubl. Popul. Sinicae
45(1): 59. 1980.
CLEWELL, A. F. Guide to the vascular plants of the Florida Panhandle. viii + 605 pp.
Tallahassee, Florida. 1985. [P. procumbens on calcareous bluffs, Jackson Co.; con-
sidered endangered in Florida 7
256 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Co.uierR, C. W. Cultivation of ais pachysandra (Pachysandra terminalis). W. Va.
Univ. Ext. Misc. Publ. 4 . 1972.
Dire, M. A., & J. H. ALEXANDER. The Allegheny Pachysandra. Arnoldia 39(1): 16-21.
1979, fletudes photographs of leaves and inflorescences (in bud) of P. procumbens
and P. terminalis. ]
Dopce, B. O. Canker blight of Pachysandra. New York Bot. Gard. Bull. 45: 159-163.
44a.
—— ew Pseudonectria on Pachysandra. sey ae 36: 532- see 1944b.
DUNCAN, W. H. Preliminary reports on the flora of Georgia—4. Notes on the distri-
bution of flowering plants, er species new to the state. Catan 15: 145-159.
1950. a procumbens, 154,
FREEMAN, J. D., A. S. Causey, J. . SHORT, & R. W. Haynes. Endangered, threatened,
and special concern plants of Alabama. Auburn Univ. Dept. Bot. Microbiol. Agr.
Exper. Sta. Dept. Ser. 3. 25 pp. Auburn, Alabama. 1979. [P. procumbens, 20, color
photo, treated as a species of special concern; known from “rich woods, usually over
limestone.” Cleburne, DeKalb, Lauderdale, Lawrence, Limestone, Marion, and
FREEMAN, O. M. Notes on the flora of Polk County, North Carolina. Castanea 20: 37-
57. 1955. [P. procumbens.
Gray, A. Dr. Siebold, Flora Japonica: sectio prima, plantas ornatui vel usul inservientes;
digessit Dr. J. G. Zuccarini. Am. Jour. Sci. Arts 39: 175, 176. 1840.
Analogy between the flora of Japan and that of the United States. Ibid. 52: 135,
136. 1846.
Gray, J. Northwest American Tertiary palynology: the emerging picture. Pp. 21 -30 in
L
Honolulu. 1960. [Includes drawing of Pachysandra pollen from the Moose Creek
sediments (Oligocene?), Salmon River Mountains, Idaho. See also J. Gray & SOHMA
under references to Buxaceae. ]
HuTT.esTon, D. G. Allegheny aie ail as a groundcover. Am. Hort. Mag. 39: 236,
Kikucul, T., & T. Toy is olation and structure determination of pachysandiol-A
and a note on the areas of cerin. Tetrahedron Lett. 1967: 3181-3185.
1967. [From P. terminalis.]
, 8S. Uyeo, T. NisHinaGa, T. IpukA, & A. Kato. Pachysandra alkaloids. en
Mass spectra of Hp ee alkaloid. (In Japanese; English summary.) Phar
Soc. Jap. Jour. 87: 631-639. 1967.* [From P. terminalis. ]
LeEopo_p, E. B., & H. D. oe Development and affinities of Tertiary floras in
the Rocky Mountains. Pp. 147-200 in A. GRAHAM, ed., Floristics and paleofloristics
of Asia and eastern North America. Amsterdam. 1972. [Pachysandra(-Sarcococca),
174, 175, 183-185; includes map showing fossil and modern occurrences.
Mutter, J. Fossil pollen records of extant angiosperms. Bot. Rev. 47: 1-146. 1981.
[Buxa aceae, 48. Pachysandra-type pollen known from Campanian of Canada, Cam-
panian-Danian interval of Germany, and ee of California, Montana, and
Canada. Buxus type from the lower Mio
Rickett, H. W. Wildflowers of the United Sites Vol. 2. The Southeastern States. Part
1. 323 pp. (including 112 pls.). New York. 1966. [P. procumbens, 152, pl. 53.)
Rossins, H.C. The nature of the species, Pachysandra | (Buxaceae). (Abstr.)
ASB Bull. 7: 38. 1960.
. Amonographic study of the genus Pachysandra (Buxaceae). v + 124 pp. (type-
script). Ph.D. thesis, Vanderbilt University. 1962. (See Diss. Abstr. 23(4): 1179.
1962.) [History, morphology, geographic distribution, and systematic treatment.]
. The genus Pachysandra (Buxaceae). Sida 3: 211-248. 1968. [Systematic part of
thesis above.]
TIEGHEM, P. vAN. Recherches sur la structure du pistil. Ann. Sci. Nat. V. 9: 127-226.
1987] CHANNELL & WOOD, BUXACEAE 257
pls. 9-12. 1868. [Buxacées, 171, 172; describes the vasculation and false partitions
, T. NisHINAGA, M. YASUNISHI, & A. YAMAMOTO.
Pachysandra alkaloids. 1. oe isolation a alkaloids. (In Japanese; English
summary.) Pharm. Soc. Jap. Jour. - Lene 227.
WARD, ae B. a Plants. Jn: P. C. H. P HARD, Ser ee Rare and endangered biota
of Florida. Vol. 5. xxix + 175 pp. Gainesville 1979. [P. procumbens, 47, illustration
and map; oe ee ieaied known in Florida only from one eee ae in J ackson Coun-
ty; treatment prepared by R. K. Goprrey & D. B. W.
Wuerry, E. T. Ne glected natives: mountain ern ‘Natl. Hort. Mag. 8: 130-
132. 1929. [P. procumben
. Neglected eee ee Ibid. 34: 211, 212. 1955. [Includes P. procumbens.]
ne
OS
HAYNES & HOLM-NIELSEN, ZANNICHELLIACEAE 259
THE ZANNICHELLIACEAE IN THE SOUTHEASTERN
UNITED STATES!
RoBERT R. HAYNES? AND LAuRITZ B. HOLM-NIELSEN?
ZANNICHELLIACEAE a Anal. Fam. Pl. 59, 61. 1829, “‘Zanichel-
liaceae,’> nom. cons.
A small family of annual [or perennial], glabrous, monoecious, aquatic herbs,
growing entirely submersed in fresh or brackish waters, rooting at the lower
nodes. Roots unbranched, 1-7 at a node, nonseptate. Stems slender, dimorphic,
the lower often stoloniferous, the upper erect and leafy, without teeth along
internodes; turions and tubers absent. Leaves alternate, opposite, or pseudo-
whorled, scalelike, without vascular tissue or foliaceous, linear, 1- [or rarely 3-]
veined, subterete, sessile, with basal sheaths, the sheath adnate to or free from
the blade, the infravaginal scales membranaceous. Inflorescences axillary, with
2 [to several] imperfect flowers. Staminate flowers short- se perianth
absent [rarely minute and 3-lobed], androecium consisting of 1 stamen, the
connective extended into a blunt appendage, the anthers (O)4(-8)L- iO ocw:
late, dehiscing by longitudinal slits; pollen inaperturate, globose, often in a
gelatinous matrix. Carpellate flowers short-pedicellate, often enclosed in a
membranaceous, spathelike envelope; perianth absent [or a small cuplike sheath;
or segments 3, separate]; carpels (1-)4 or 5(-8), separate, short-stipitate,
1-loculate; ovule solitary, bitegmic, pendulous, anatropous, placentation apical;
style short [long], stigma enlarged, + funnel shaped [feathery or peltate]. Fruit
drupaceous, with a membranaceous exocarp, fleshy mesocarp, and stony en-
docarp. Seed solitary; embryo curved; endosperm helobial in development,
absent in mature seed. (Zannichelliaceae sensu stricto, excluding genera that
‘Prepared for the Generic Flora of the Southeastern United States, a long-term project currently
made possible through the support of National Science Foundation grants BSR-8415769 (Carroll E
Wood, Jr., principal investigator) and BSR-8415637 (Norton G. Miller, eae ay Eee The
117th in the series, this paper follows the format established in the first one (Jo
296-346. 1958) and continued to the present. The area covered b Lg the Generic ee santa Nort
members of a family or genus in brackets [ ]. References that we have not verified are marked wit
an asterisk.
We are indebted to Drs. Wood and Miller for their advice, suggestions, and help with the literature
during the preparation of the manuscript.
The illustration was drawn by Karen Stoutsenberger at the Arnold Arboretum under Haynes’s
direction from material he collected in Alabama.
2Department of Biology, University of Alabama, P. O. Box 1927, Tuscaloosa, Alabama 35487-
7
3Botanical Institute, University of Aarhus, DK-8240, Risskov, Denmark.
© President and Fellows of Harvard College, |
Journal of the Arnold Arboretum 68: 259-268. pee 1987.
260 JOURNAL OF THE ARNOLD ARBORETUM [vVOL. 68
are better placed in the Potamogetonaceae, Cymodoceaceae, and Posidoni-
aceae.) TYPE GENUS: Zannichellia L.
Four genera and ten to twelve species; represented in the southeastern United
States by one species of Zannichellia, a nearly cosmopolitan genus consisting
of four or five species. Zannichellia differs from Pseudalthenia Nakai, A/thenia
Thouars, and Lepilaena Drumm. ex Harvey in lacking a creeping rhizome, in
having mostly four- or five-carpellate flowers (rarely fewer than four-carpellate),
and in having warty fruits. Tomlinson & Posluszny indicated that no clear
lines of evolution are recognizable in the family.
Pseudalthenia Aschersoniana (Graebner) Den Hartog (V/eisia Aschersoniana
(Graebner) Tomlinson & Posluszny) is an endemic of the Cape Southwest region
of South Africa, where it grows in vleis (depressions in which water collects
during the wet season), The species 1s unique in the family in having leaves
with a submarginal vascular strand and transverse strands continuous with the
midvein. The staminate flower lacks a perianth, is eight-sporangiate, and has
a pair of vestigial appendages on the connective. The carpellate flower is always
unicarpellate and produces a papillate fruit, with the papillae not arranged in
Althenia, with two species in northern Africa, the west-central Mediterranean
region, and the Atlantic coasts of Morocco, Spain, Portugal, and France, is
characterized by peltate stigmas and styles about 3 mm long.
Lepilaena consists of three species endemic to Australia and a fourth oc-
curring in New Zealand and Australia. Diagnostic features of the genus include
two- or twelve-sporangiate staminate flowers and carpellate flowers with short
styles and funnel-shaped or feathery stigmas.
Cronquist placed the Zannichelliaceae in the Najadales, whereas Dahlgren,
Dahlgren & Clifford, and Thorne (1976, 1983) put the family in the Zosterales.
The Zannichelliaceae as here interpreted have been combined variously with
members of the Potamogetonaceae, Najadaceae, Zosteraceae, and Cymodo-
ceaceae under the names Zannichelliaceae (Taylor), Zosteraceae (Fernald), Na-
jadaceae (Gleason & Cronquist), and Potamogetonaceae (Ascherson & Graeb-
ner). Miki considered Najas L. to be closely related to the Zannichelliaceae,
especially A/thenia, less so to Zannichellia.
Pollen is mostly dispersed as single grains but 1s occasionally contained in
a gelatinous matrix (as in Zannichellia palustris). The grains are spherical,
nonaperturate or rarely monosulcoidate, binucleate, and sparsely and unevenly
verrucate. Adjacent verrucae are often in contact. The endexine, according to
Pettitt & Jermy (see generic references), is very indistinct, and the intine is
The family is known to have secondary compounds, including flavonoid
bisulphates, flavones (Gornall et a/.), and apiose (Van Beusekom).
Cytological data are incomplete for the family, but the reported chromosome
numbers include 2n = 12, 24, 28, 32, and 36 (x = 6 or 8)
The Zannichelliaceae are all aquatic herbs and grow clonally in shallow,
generally brackish coastal waters or in inland freshwater lakes.
he roots are all adventitious and unbranched; they arise from nodes of the
1987] HAYNES & HOLM-NIELSEN, ZANNICHELLIACEAE 261
creeping and sympodially branched rhizomes or from those of the erect and
richly branched leafy stems. The leaves are linear, sheathing at the base, and
with rounded, pointed, truncate, or toothed apices. Pairs of inconspicuous,
filiform squamules (nonvasculated scales) occur laterally at the nodes.
The unbranched roots have a thin-walled epidermis of large cells and con-
spicuous root-hairs that arise from short trichoblasts. The outer cortex is com-
pacted into an exodermis of one or two layers of narrow, slightly lignified,
thick-walled cells, while the inner part is lacunose, the endodermis aie
and thin walled, and the stele narrow, surrounding a metaxylem lacun
The stems are nearly without mechanical tissue, and the pian c co orex:
endodermis, and stele resemble those of the roots. Vascular bundles supporting
ee organs diverge directly from the stele, and there is no cortical vascular
system.
The leaf blade is glabrous, with the epidermis uniform, thin walled, and
chlorophyllous. The epidermis mostly lacks stomata, although they do occur
in the apices of leaf blades of certain species of Zannichellia. The mesophyll
is lacunose either throughout or only on each side of the midvein. The vascular
system is reduced to a single median vascular bundle surrounded by a uniseriate
endodermis. The leaf blades have submarginal fibers.
The plants are monoecious, with complex, terminal, sympodial inflorescences
of reduced, specialized flowers subtended by reduced bractlike leaves. Each
inflorescence usually has one staminate flower terminating the first-order mer-
istem and one to several carpellate flowers terminal on branches of higher
orders.
The staminate flowers are short-pedicellate, reduced to one stamen, and with
or without a short, three-lobed, scalelike perianth. The anther consists of one
or more bisporangiate units, sometimes with a short connective appendage;
dehiscence is longitudinal. The tapetum is of the periplasmodial type, micro-
sporogenesis is of the successive type, and the pollen grains are three-celled at
dispersal.
The carpellate flowers are short-pedicellate and have one to eight separate,
short-stalked, slightly asymmetric carpels. The carpels are surrounded by a
biseriate perianth that consists of a closed tubelike structure in Zannichellia
and Pseudalthenia, and of three separate segments in A/thenia and Lepilaena,
with the segments opposite the carpels. The styles are more or less elongate
and are terminated by enlarged peltate or funnel-shaped stigmas that have
more or less lacerate margins or are occasionally feather shaped. Each of the
stipitate carpels contains a solitary, pendulous, anatropous, bitegmic ovule.
The embryo sac is of the Allium type, with embryo formation of the cary-
ophyllad type. Endosperm is of the helobial type but is absent in the mature
seed.
REFERENCES:
ER, A. Water plants. xvi + 436 pp. Cambridge, England. 1920. [Review of the
biology of aquatic vascular plants; Zannichelliaceae discussed throughout.]
ASCHERSON, P. Potamogetonaceae. Nat. Pflanzenfam. II. 1: 194-214. 1889.
262 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
& RAEBNER. Potamogetonaceae. Pflanzenr. IV. 11(Heft 31): 1-184. 1907.
[Tribe Zannichellieae, 153-160.
Aston, H. I. Aquatic plants of Australia. xv + 368 pp. Melbourne. 1973. [Zannichel-
liaceae, 301-325, including Amphibolis, Cymodocea, Halodule, Lepilaena, Syrin-
godium, Thalassodendron, Zannichellia palustris.
BaILton, H. Najadacées. Hist. Pl. 12: 99-126. 1894. [Série des Zannichellia, 105, 106;
Zannichellieae, 122, 123.
BEAL, E.O. A manual of marsh and aquatic vascular plants of North Carolina. N. Carolina
Agr. Exper. Sa Tech. Bull. 247. iv + 298 pp. 1977. [Zannichelliaceae, Z. palustris,
fag.).J
BENTHAM, G., & J. D. Hooker. Naiadaceae. Gen. Pl. 3: 1009-1019. 1883. [Tribe
Zannichellieae, 1016, 1017, apes one Althenia, Lepilaena.]
BEUSEKOM, C. F. vAN. Ueber einige Apiose-Vorkommnisse bei den Helobiae. Phyto-
chem mistry 6: 573-576. 1967. iZannichelliaceae, including Zannichellieae (Zanni-
chellia palustris), Cymodoceae.]
BOLKHOVSKIKH, Z., V. GRIF, T. MATVEJEVA, & O. ZAKHARYEVA. Chromosome number
of flowering plants. A. A. Feporov, ed. 926 pp. Leningrad. 1969. [Althenia, Zan-
nichellia palustris.
CAMPBELL, D. H. A morphological study of Naias and Zannichellia. Proc. Calif. Acad.
Sci. III. Bot. 1: 1-70. pls. 1-5. 1897. [Zannichellia (Z. palustris), 35-60, 67-70, pls.
4, 5.
Casper, S. J., & H.-D. KrAuscH. Pteridophyta und Anthophyta. |. Teil: Lycopodiaceae
bis Orchidaceae. Band 23 in H. Err i, J. GeRLorr, & H. HEYNIG, Siisswasserflora
von Mitteleuropa. 403 pp. Stuttgart and New York. 1980. [Zannichelliaceae, 150,
152-155, 402; Althenia ae Zannichellia
CLAPHAM, A. R., T. G. TutTin, & E. F. WARBURG. Flora of the British Isles. ed. 2.
xviii + 1269 pp. Cambridge, England. 1962. [Zannichelliaceae, 960.
CLEWELL, A. F. Guide to the vascular plants of the Florida Panhandle. 605 pp. Talla-
hassee. eg [Zannichelliaceae, 199.]
Cook, C. D. K. Zannichellia ee Pp. 275, 276 in V. H. HEywoop, Flowering plants
of the world. New York.
CRANWELL, L. M. New ne pollen studies. The monocotyledons. A comparative
account. Bull. Auckland Inst. Mus. 3: 1-91. 1952. [Zannichelliaceae, 25, 26.
Cronaquist, A. The evolution and classification of flowering plants. x + 396 pp. Boston.
1968. [Najadales, 327-330, including Aponogetonaceae, Scheuchzeriaceae, Juncag-
inaceae, Najadaceae, Potamogetonaceae, Ruppiaceae, Zannichelliaceae, Zostera-
ceae.
. An integrated system eae are of flowering plants. xviii + 1262 pp. New
York. 1981. [Zannichelliaceae, 1068, 1069.
DaGHLIAN, C. P. A review of the fossil ae of monocotyledons. Bot. Rev. 47: 517-
55. 1981.
DAHLGREN, R. M. T. A revised system of classification of the angiosperms. Bot. Jour.
Linn. Soc. 80: 91-124. 1980. [Zosterales, 98, including Scheuchzeriaceae, Juncagi-
naceae, Najadaceae, Potamogetonaceae, Teter. Posidoniaceae, Cymodocea-
ceae, Zannichelliaceae. ]
& H. T. CuirForp. The monocotyledons. A comparative study. Bot. Syst. 2.
XIV + 378 pp. London. 1982. [Zannichelliaceae discussed throughout.]
, . F. Yeo. The families of the monocotyledons. Structure, evolution
and taxonomy. xii + 520 pp. Berlin, Heidelberg, New York, and London. 1985.
[Najadales, 307-322, including Scheuchzeriaceae, Juncaginaceae, Potamogetona-
ceae, — Posidoniaceae, Cymodoceaceae, Najadaceae, Zannichelliaceae,
318-320,
Danpy, J. E. Zannichelliaceae. a: T.G. Tutin, V. H. HEywoop, et al., eds., Fl. Europaea
5: 12, 13. 1980. pana palustris, Whee filiformis, Cymodocea nodosa. ]
1987] HAYNES & HOLM-NIELSEN, ZANNICHELLIACEAE 263
ERDTMAN, G. Pollen morphology and plant taxonomy. Angiosperms. Frontisp. + xii +
539 pp. Uppsala. 1952. [Zannichelliaceae, 454.]
Fassett, N. C. A manual of aquatic plants (with revision appendix by E. C. OGDEN).
iv + 405 pp. Madison, Wisconsin. 1957. [Najadaceae, 55-77, including Potamo-
geton, Ruppia, Najas, Zannichellia, 74, 75, fig.)
FERNALD, M. L. Gray’s manual of botany. ed. 8. Ixiv + 1632 pp. New York. 1950.
[Zannichellia, 80, 81.]
G.eason, H. A., & A. Cronquist. Manual of vascular plants of northeastern United
States and adjacent Canada. li + 810 pp. Princeton, New Jersey. 1963. [Najadaceae,
33-40; pores Ruppia, Najas, Zannichellia palustris.]
GorRNALL, R. J., . Boum, & R. DAHLGREN. The distribution of flavonoids in the
angiosperms. oe Not. 132: 1-30. 1979. [Zannichelliaceae with luteolin and/or
apigenin, methylated flavones, and flavone bisulphates
Hartoo, C. DEN. Pseudalthenia antedates Vieisia, a nomenclature note. Aquatic Bot.
9: 95. 1980. [Pseudalthenia Nakai; P. Aschersoniana (Graebner) Den Hartog, comb.
nov.]
Hutcuinson, G. E. A treatise on limnology. Vol. 3. Limnological botany. xi + 660 p
New York. 1975. [A discussion of the biology—especially chemical ecology— 1
aquatic vascular plants; Zannichelliaceae, 108, 129.]
Hutcuinson, J. The families of flowering plants. ed. 2. Vol. 3. Monocotyledons. viii +
792 pp. Oxford. 1959. Kangeccrar ie 16, 17, 78, 81, 90, 92.]
LE ey E., & J. DECAISNE. Traité général de botanique, descriptive et analytique.
+ 745 pp. Paris. 1868. IZannichelliaceae, 646, 647.]
eee D. Histogenese und Anatomie von Prim arwurzeln und sprossbiirtigen Wurzeln
einiger Potamogetonaceae L. (English summary.) Beitr. Biol. Pflanzen 46: 247-313.
1969. [Groenlandia, Halodule, Pomona Ruppia, Zannichellia, Zostera. |
MarkarafF, F. Bliitenbau und V' hsten Helobiae. Ber. Deutsch.
Bot. Ges. 54: 191-229. pls. 1-8. 1936. [Althenia, Zannichellia, 212-214.]
Mixt, S. The origin of Najas and Potamogeton. Bot. Mag. Toky 0 51: 472-480. 1937.
[Najas i is closely related to Zannichelliaceae, especially Althenia.]
Morona, T. L. The Naiadaceae of North sao a. Mem. Torrey Bot. Club 3(2): 1-65.
pls. 20-74. 1893. [Zannichellia, 56, 57, pl. 6
PosLuszny, U., & P. B. Tomiinson. Morphology in development of floral shoots and
organs in certain Zannichelliaceae. Bot. Jour. Linn. Soc. 75: 21-46. 1977. [Zanni-
chelliaceae, including A/thenia, Lepilaena, Vieisia (= Pseudalthenia).]
RENDLE, A. B. The ae of flowering plants. ed. 2. Vol. 1. Gymnosperms and
monocotyledons. + 412 pp. Cambridge, England. 1930. [Potamogetonaceae,
202-208; Zannichellia' in hae Zannichellicae ]
SAUVAGEAU, C. Sur les ues monocotylédones aquatiques. Ann. Sci. Nat
Bot. VII. 13: 103. 296. Ta 7 ener 959-264: Zannichellia, Althenia, Lep-
ilaena.]
SCHUMANN, K. Morphologische Studien. Heft 1. x + 206 pp. + 6 pls. Leipzig. 1892.
eee 154-174, pl. 6.]
ichelliaceae. Jn; K. F. P. von Martius, FI. Brasil. 3(3): 703-714. pi. 122.
1894. I Zannichellia nen ]
se eae C. D. The biology of aquatic vascular plants. xviii + 610 pp. London.
967. [A review of the biology of aquatic vascular plants; Zannichelliaceae discussed
. see especially 297-299.
TAKHTAJAN, A. Flowering plants. Origin and dispersal. (Authorized translation from
the Russian by C. Jerrrey.) x + 310 pp. Edinburgh. 1969. [“Najadales or Pota-
mogetonales,” 234, including Scheuchzeriaceae, Juncaginaceae, Aponogetonaceae,
Zosteraceae, Posidoniaceae, Potamogetonaceae, Ruppiaceae, Zannichelliaceae, Cy-
modoceaceae, and Najadaceae.
264 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
TAyLor, N. Zannichelliaceae. N. Am. Fl. 17: 13-27. 1909. [Zannichellia, Ruppia, and
THORNE, R. F. A phylogenetic classification of the angiosperms. Evol. Biol. 9: 35-106.
1976. [Zannichelliaceae in Zosterales suborder Potamogetonineae, along with Pota-
mogetonaceae, Juncaginaceae, and Posidoniaceae.]
Proposed new realignments in the angiosperms. Nordic Jour. Bot. 3: 85-117.
1983. [Placement of Zannichelliaceae as in preceding reference. ]
). In:
1
TOMLINSON, : B. Helobiae -C. R. METCALFE,
ed., Anatomy of the Soncebitons. Vol. 7. xvi + 559 pp. “Oxford. 1982. [Zan-
nichelliaceae, 336-369. ]
U. Pos_uszny. Generic limits in the Zannichelliaceae (sensu Dumortier).
Taxon 25: 273-279. 1976. [Vieisia, a new genus proposed to accommodate Zan
nichellia Aschersoniana, is antedated oh oe Nakai, with the single meee
P. Aschersoniana (Graebner) Den Hart
. Aspects of floral ses and development in the seagrass Sy-
Sel aA a a ea Bot. Gaz. 139: 333-345. 1978. [Includes table
ison of Syringodium with Lepilaena, Althenia, and Vleisia (= Pseuda
ena) .
1. Zannichellia Linnaeus, Sp. Pl. 2: 969. 1753; Gen. Pl. ed. 5. 416. 1754.
Annual or rarely perennial, monoecious plants of fresh or brackish waters.
Roots single or in pairs at the nodes. Leaves in pseudowhorls of 3 but usually
also alternate and opposite on same plant, entire, stipulate, mostly less than |
mm wide, | veined. Inflorescence usually consisting of 2 flowers, 1 staminate
and | carpellate. Flowers without a perianth. Staminate flowers with a single
usually 4-loculate [2-8-loculate] stamen, the connective prolonged into a blunt
appendage. Carpellate flowers with (1-)4 or 5(-8) carpels surrounded basally
membranaceous envelope, the style less than 1 mm long, the stigma
asymmetrically funnel shaped. Fruit endocarp often coarsely papillose. TyPE
SPECIES: Z. palustris L., the only species of the genus in Species Plantarum.
(Named after Gian Girolamo Zannichelli, 1662-1729, a Venetian apothecary
and botanist.)— HORNED PONDWEED.
A nearly cosmopolitan genus of perhaps five species, represented in the
southeastern United States only by Zannichellia palustris L. The genus has
been variously interpreted as consisting of one highly variable species (e.g.,
Dandy) or as many as five species (e.g., Holm-Nielsen & Haynes; Van Vierssen,
1982a). We recognize the genus to comprise at least one near-cosmopolitan
species (Z. palustris) and four others of restricted distribution, of which three
(Z. major Boenn., Z. pedunculata Reichb., and Z. peltata Bertol.) are in north-
ern Europe and one (Z. andina Holm-Nielsen & Haynes) is in the high Andes
of South America.
Zannichellia has an unusual pollination system in which the anther of the
staminate flower arches over the funnel-shaped stigmas of the carpellate flower.
Pollen transfer is entirely underwater: it is released from the anther in a ge-
latinous mass and falls directly into the stigma. Such a system limits outcrossing
but is valuable for a submersed annual aquatic since pollination is essentially
assured.
Reported chromosome numbers for Zannichellia are n = 12, 2n = 24, 28,
1987] HAYNES & HOLM-NIELSEN, ZANNICHELLIACEAE 265
Figure 1. Zannichellia. a—j, Z. palustris: a, branch of plant with fruit, x 1; b, node
with staminate and carpellate flowers, base of leaf (to right), portion of stem, and base
as 2 branches, x 12: c, staminate flower (a single stamen) and carpellate flower with 2
rpels, x 12—note expanded stigmas; d, anther shedding pollen, x 25; e, cross section
ee anther before dehiscence, showing 4 locules, a few pollen grains indicated diagram-
flower, 1 carpel undeveloped, x 6; g, h, endocarps of 2 fruits, with tip of style still covered
by outer part of pericarp, x 6; 1, fruit, the ovary in longitudinal section to show embryo,
x 12; Jj, embryo, x 12.
32, 36 for Z. palustris (Bolkhovskikh et al.), 2n = 36 for Z. pedunculata and
2n = 12, 36 for Z. peltata (Van Vierssen & Van Wijk).
Daghlian did not report the Zannichelliaceae in the fossil record, although
Katz and colleagues listed three species from the Quaternary in the Soviet
Union. Miller reported Zannichellia from lateglacial deposits in western New
266 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
York, and Pierce & Tiffney have reports from the postglacial Holocene in
Connecticut.
REFERENCES:
Under family references see ARBER; BAILLON; BEAL; BOLKHOVSKIKH ef al.; CAMPBELL;
AGHLIAN; DANDY; FASSETT; FERNALD: GLEASON & CRONQUIST; G. E. HUTCHINSON;
ee SAUVAGEAU; SCULTHORPE; TAYLOR; TOMLINSON; and TOMLINSON & POSLUSZNY.
BURGMEISTER, H. Entwicklungsphysiologische Untersuchungen zur Heterophyllie und
Stomatabildung bei Zannichellia palustris L. Beitr. Biol. Pflanzen 44: 67-121. 1968.
&
Press. 1975.) [Zannichelliaceae (Zannichellia palustris), 117, 119 (fig.), 120. M
. JOHNSTON. Manual of the vascular plants of Texas. Fronts,
1881 DD. + map. Renner, Texas. 1970. [Zannichelliaceae (Zannichellia Be eee
|
CRONQuIST, A., A. H. HOLMGREN, N. H. HOLMGREN, J. L. REVEAL, & P. K. HOLMGREN.
Intermountain flora. Vascular plants of the Intermountain West, U.S.A. Vol. 6. 584
pp. New York. 1977. [Zannichellia palustris, 44, 45, fig.]
DoroFEEV, P. I. On the Pleistocene flora of the locality of village Vyshgorod on Dniepr
(In Russian; English summary.) Bot. Zhur. 49: 1093-1100. pls. 7, 2. 1964. [Includes
Zannichellia palustris. |
e Pliocene flora of the Matanov Garden on the River Don. Bot. Inst. V. L.
Komarova, Izdat. Nauka, Leningrad. 87 pp. 1966.* [Includes Zannichellia, ae
Potamogeton.|
Evtes, D.E., & J. L. ROBERTSON, JR. A guideand key to the aquatic eee of southeastern
United States. U. S. Publ. Health Bull. 286. iv + 151 pp. + map. [Zannichellia
GLEASON, H. A. The new Britton and Brown illustrated flora of the northeastern U. S.
and adjacent Canada. Vol. |. Ixxv + 482 pp. New York. 1952. [Najadaceae, 74—
87, including Potamogeton, Ruppia, Zostera, Najas, and Zannichellia, Z. palustris,
86, 87, fig.]
Goprrey, R. K., & J. W. Wooren. Aquatic and wetland plants of the southeastern
United States. Monocotyledons. x + 712 pp. Athens, Georgia. 1979. [Zannichelli-
aceae (Zannichellia palustris), 29, 30.]
— NER, Le & M. FLAHAULT. 6 Familie. Potamogetonaceae. Pp. 394-543 in O. von
R et al., Lebensgeschichte der Bliitenpflanzen Mitteleuropas. Band 1, Abt.
1. areas 1908. [Six genera; Zannichellia (by P. GRAEBNER), 509-516.
HARRINGTON, H. D. Manual of the plants of Colorado. For the identification of ferns
and oe plants of the state. x + 666 pp. Chicago. 1954. [Zannichellia (Z.
naliie. 3
Haynes, R. R. re atic plants of Alabama. I. Alismatidae. Castanea 45: 31-50. 1980,
[Zannichellia palustris. ]
& . HoLtM-NIELSEN. A generic treatment of Alismatidae in the Neotropics
with special reference to Brazil. Acta Amazonica Suppl. (In press.) [Zannichellia
engine, Z. palustris
Zan
]
nichelliaceae. Jn: G. HARLING & B. Sparre, eds., Flora of Ecuador.
Re a [Zannichellia andina. |
ci, G. Potamogetonaceae. | Mitteleuropa 1: 120-144. pls. 16-18. 1907. [Zanni-
re ohellia, 140, 141, pi. 17,
HELLQuisT, C. B., & G. E. Con Aquatic vascular plants of New England: part 1.
1987] HAYNES & HOLM-NIELSEN, ZANNICHELLIACEAE 267
Zosteraceae, Potamogetonaceae, Zannichelliaceae, Najadaceae. New Hampshire Agr.
xper. Sta. Bull. 515. iii + 68 pp. 1980. [Zannichelliaceae (Zannichellia palustris),
9, 60.]
Histncer, E. Recherches sur les tubercules du Ruppia rostellata et du Zannichellia
polycarpa, provoqués par le Tetramyxa parasitica. Medd. Soc. Faun. Fl. Fenn. 14:
53-62. pls. I-10. 1887.
Hitcucock, C. L., & A. CRONQUIST. Flora of the Pacific Northwest. An illustrated
manual. xix + 730 pp. Seattle. 1973. [ ), 566.]
HocuHReEvuTINER, C. Etudes sur les phanérogames aquatiques du Rhone et du port de
Genéve. Rev. Gén. Bot. 8: 90-110, 158-167, 188-200, 249-265. pl. 7, text figs. 5-
65. 1896. [Premiére partie. Morphologie et anatomie du Zannichellia palustris L.,
91-110; Zannichellia also in the six sections the second part.]
How, L., J. V. PANCHO, J. P. HERBERGER, & D. L. PLucknett. A geographical atlas
of world weeds. xlix + 391 pp. New York. 1979, [Zannichellia, 389.]
Hoim-NIeE.sen, L. B., & R. R. HAYNES. Two new Alismatidae from Ecuador and Peru
(Alismataceae and Zannichelliaceae). Brittonia 37: 17-21. 1985. [Zannichellia an-
dina, sp. nov.
Hotcukiss, N. Underwater and floating-leaved plants of the United States and Canada.
. Dept. Int. Fish Wildlife Serv. Bur. Sport Fish. Wildlife Res. Publ. 44. vii +
124 pp. ae 63.
Wear Ne Jecoe ¥ z, & M. G. Kipranr. Atlas and keys of fruits and seeds occurring
in the Seen deposits of the USSR. (In Russian; English and Russian title
pages.) 365 pp. Moscow. 1965. [Zannichellia, 128, 129,
LAKSHMANAN, K. K. Note on the endosperm formation in Zannichellia palustris L.
ee Buenos Aires 22: 13, 14. 1965.
Lona, R. W., & O. Laketa. A flora of tropical Florida. xvi1 + 962 pp. Coral Gables.
1971. [Zannichelliaceae (Z. palustris), 118
MacRoserts, D. qT The vascular plants of Louisiana. An annotated epee as
bibliography of the vascular ted to grow without cultivation in Lou
Bull. Mus. Life Sci. Louisiana State Univ. 6. 165 pp. 1984. a elite. (Z.
palustris), 54.]
Martin, A. C. The comparative internal morphology of seeds. Am. Midl. Nat. 36: 513-
660. 1946. Fede ae ie including et Scat Ruppia, Zannichellia,
Zostera, all in the category “linear embryos,” none illustra
Mason, H. L. A flora a the marshes ae California. xi + 878 ro Berkeley, California.
1969. ata as (Zannichellia hey, 89, 90, fig. 37.]
McATEE, W. L. Wildfowl food plants. Thei ue, propagation, and management. x +
141 pp. Ames, Iowa. 1939. ea ieheIGe: 16. 17, 78, 81, 90, 92.]
McCvure, J. W. Secondary constituents of aquatic angiosperms. Pp. 233-268 in J. B.
Harsorne_, ed., Phytochemical phylogeny. London. 1970. [Zannichelliaceae, 7. pa-
lustris.
Mutter, N. G. Lateglacial plants and plant communities in northwestern New York.
se Arnold Arb. 54: 123-159. 1973. [Zannichellia palustris var. major (Boenn.)
W. D. J. Koch, 145, 146 (fig.).]
Morone, T. L. How to collect ae plants. Aquatic plants (Naiadaceae, etc.). Bot.
Gaz. 11: 139, 140. 1886. [Includes Zannichellia palustris.]
MUuENSCHER, W. C. Aquatic plants of the United States. x + 374 pp. Ithaca, New York
1944, [Potamogetonaceae, 27-65, including ees ty Ruppia, Zannichellia (Z.
palustris), Phyllospadix, Zostera, Halodule, Cymodoce
OcpeEN, E. C. Anatomical patterns of some aquatic ae ants of New York. New
bee State Mus. Bull. jae v + 133 pp. 1974. Saas palustris, 11, map 52,
Dies
= K. Dean, C. W. Boy en, & R. B. SHELDON. Field guide to aquatic plants of
Lake George, New York. Ibid. 426. iv + 65 pp. 1976. [Zannichelliaceae, 16.]
268 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Pettitt, J. M., & A. C. Jermy. Pollen in hydrophilous angiosperms. Micron 5: 377—
405. 1975. [Includes Zannichellia palustris.
—— e S., & B. H. Tirrney. Holocene fruit, seed, and leaf flora from riverine
ments near New Haven, Connecticut. Rhodora 88: 229-252. 1986. [Zannichellia
eanete 238, 249, fig. 12.
PosLuszny, U., & R. SATTLER. Floral development of Zannichellia palustris. Canad.
Jour. Bot. 54: 651-662. 1976. [The fertile node complex of Zannichellia palustris
appears at first to be a perfect flower. ]
RaApForD, A. E., H. E. AHLEs, & C. R. BELL. Manual of the vascular flora of the Carolinas.
Ixi + pe pp. Chapel Hill, North Carolina. 1968. [Zannichelliaceae (Z. palustris),
48, fig., map.
REESE, G. Ober die deutschen Ruppia- und Zannichellia-Kategorien und ihre Verbrei-
tung in ae Holstein. Schr. Naturw. Ver. Schlesw.-Holst. 34: 44-70. 1963.*
tologische und taxonomische Untersuchungen an Zannichellia palustris L.
Biol. Fae. 86(Suppl.): 277-306. 1967.*
REINECKE, P. A contribution to the morphology of Zannichellia Aschersoniana Graebn.
Jour. S. Afr. Bot. 30: 93-101. 1964. [Carpellate flowers “persistently” one-carpellate. ]
Rose, E. Le mode de fécondation du Zannichellia palustris L. Jour. Bot. Morot 1: 296-
299. 1887.
SCHENCK, H. Vergleichende Anatomie der submersen Gewachse. Bibliot. Bot. 1(Heft
1). 67 pp. + 10 pls. 1886. [Zannichellia, 16, 44, pl. 3, fig. 12.]
SMALL, J. K. Manual of the southeastern flora. xxii + 1554 pp. New York. 1933.
(Reprinted by Univ. N. Carolina Press, Chapel Hill.) [Zannichellia palustris, 15.]
STEYERMARK, J. S. Flora of Missouri. Ixxxiti + 1725 pp. Ames, Iowa. 1962. [Zanni-
chellia palustris, 56.]
SUBRAMANYAM, K. Aquatic angiosperms. vill + 190 pp. Calcutta. 1962. [Zannichellia,
7.
Soieen, W. Miocene flora from Stare Gliwice in Upper Sein Prace Inst. Geolog. 33:
1-205. ae [Includes Zannichellia, Potamogeton, Ruppia.]
UoTILA, P., W. VAN VIERSSEN, & R. J. VAN WuK. Notes on the = catieeranl taxonomy
of Zannichellia in Turkey. Ann. Bot. Fenn. 20: 351-356. 1983. [Chromosome num-
bers of 2n = +32 for Zannichellia major and 2n = 24 for Z. palustris.]
VENKATESH, C. 8. Anther and pollen grains of Zannichellia palustris L. Curr. Sci. Ban-
galore 21: 225, 226. 1952.*
VIERSSEN, W. vAN. The ecology of communities dominated by Zannichellia taxa in
western Europe. I. Characterization and autecology of the Zannichellia taxa. Aquatic
Bot. 12: 103-155. 1982a. [Morphological as cytological characteristics of Zanni-
chellia from Europe, including Z. major, Z. palustris, Z. pedunculata, Z. peltata.]
II. Distribution, synecology and productivity aspects in ere to environmental
factors. Ibid. 13: 385-483. 1982b. [Environmental factors as in distribution
of European Zannichellia, including Z. major, Z. palustris, Z. pedunculata.] UI.
Chemical ecology. Ibid. 14: 259-294. 1982c. [Chemical constituents of European
Zannichellia, including Z. palustris and Z. pedunculata. |
. Reproductive strategies Zannichellia taxa in western Europe. Pp. 144-149
in J. J. Symoens, S. S. Hooper, & P. Compére, eds., Studies on aquatic vascular
plants. eee 1982. ‘Gann abd Z. major, Z. pedunculata, Z. peltata.|
.J. VAN Wyk. On the identity and shad’ of Zannichellia peltata Bertol.
western Europe Aquatic se 13: 367-383.
VUAVARAGHAY ,M. R., & A. V. KUMARI. ie and a position of
Zan Beet palisine L. an ‘Indian Bot. aa 53: 292-302. 1974.
Voss, E. G. Michigan flora. Part |. Gymnosperms and monocots. c nbrook Inst. Sci.
Bull. 55. xv + 488 pp. Bloomfield Hills, Michigan. 1972. [Zannichelliaceae (Zan-
nichellia palustris), 93 (map), 94, 96 (fig.).]
Warp, D. B. ecklist of the vascular flora of Florida. Part 1. Univ. Florida Agr. Exper.
Sta. Tech. Bull. 726. 72 pp. 1968. [Zannichellia palustris, 19.]
GEORGE RALPH COOLEY
May 29, 1896-September 27, 1986
We record with regret the death of George R.
Cooley, a friend of botany and botanists and a gen-
erous supporter of the Generic Flora of the South-
eastern United States during its initial years.
Journal of the Arnold Arboretum April, 1987
CONTENTS OF VOLUME 68, NUMBER 2
The Genera of Cinchonoideae (Rubiaceae) in the Southeastern United
States.
SHORE 1 GEES ac 48 koi re a OE eR ee eR 137-183
The Genera of Alysseae (Cruciferae; Brassicaceae) in the South-
eastern United States.
IHSAN A, AL-SHEHBAZ «ie 6556055000800) ¥o~ ue) 44 bbe odes 185-240
The Buxaceae in the Southeastern United States.
R. B. CHANNELL AND C. E. Woop, JR. 00.2 ee 241—257
The Zannichelliaceae in the Southeastern United States.
Ropert R. HAYNES AND LAuRITZ B. HOLM-NIELSEN .......... 259-268
Volume 68, Number |, including pages 1-136, was issued January 6, 1987.
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JOURNAL
OF THE
ARNOLD ARBORETUM
VOLUME 68 Juty 1987 NUMBER 3
A CLADISTIC ANALYSIS OF CONIFERS:
PRELIMINARY RESULTS
JEFFREY A. HART!
A data matrix of 123 binary and multistate characters of 63 genera of conifers
was constructed based on an extensive literature review and study of herbar-
ium ue living specimens. Subsequent cece 8 of this matrix strongly
supports the monophyly of conifers; there is no reason to exclude the taxads.
Sciadopitys should be considered as constituting a separate family, the Scia-
and can be divided into two groups, one of northern and the other of southern
appear to be paraphyletic. The Taxaceae and Cephalotaxaceae also come out
as sister taxa. The Pinaceae appear to be the sister group of the other living
conifers. The placement of Araucariaceae and Podocarpaceae in relationship
to the other living conifers is problematic.
Conifers have long been of interest to morphologists, anatomists, paleobot-
anists, and foresters. A cosmopolitan group, conifers include 60 to 63 genera
and 500 to 600 species. Known from the fossil record from as far back as the
ng gymnosper ate, the
monophyly of the conifers and the phylogenetic relationships of the families
and genera have not been determined.
Most modern textbooks follow Pilger (1926) in dividing the group directly
into seven families (Taxaceae Sprengel, Podocarpaceae Endl., Araucariaceae
Strasburger, Cephalotaxaceae Neger, Pinaceae Lindley, Taxodiaceae Neger, and
Cupressaceae S. F. Gray), but other classifications have also been proposed.
Buchholz (1933) divided the Coniferae into two suborders: the Pinineae (in-
"Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138.
© President and Fellows of Harvard College, 1987
Journal of the Arnold Arboretum 68: 269-307. July, 1987.
270 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
cluding Pinaceae, Cupressaceae, Taxodiaceae, and Araucariaceae), with ob-
vious cones, and the Taxineae (including Podocarpaceae, Taxaceae, and Ceph-
alotaxaceae), without obvious cones. Sahni (1920) and Florin (1948b, 1951)
elevated the Taxaceae to the Taxales, equal to all other conifers in ordinal
rank. Keng (1973, 1975) has recently recognized eight families, elevating Phy/-
locladus Rich. (Podocarpaceae) to family rank. For a more complete review,
see the excellent summaries by Florin (1955) and Turrill (1959).
The phylogenetic and evolutionary relationships among these families and
genera have been widely debated. The lack of precise, explicit methodologies
for assessing phylogenetic relationships has resulted in a diversity of views
about conifer relationships. Historically, schemes of evolutionary relationships
have been based primarily on assertions as to the usefulness of individual plant
characters as phylogenetic markers
With the introduction of adie theory as developed by Hennig (1950,
1966) and his followers, there has been a renewed interest in the study of
higher-level taxonomic relationships in systematic biology. The purposes of
this paper are to review the kinds of evidence used historically in assessing
phylogenetic relationships among conifers; to construct a comprehensive char-
acter data matrix both to serve in the analysis and to provide the basis for
further studies; to utilize cladistic methodology in the study of phylogenetic
relationships of coniferous genera; to compare these results with previously
held notions of relationships; and to suggest new areas of research needed to
test my hypotheses of relationships among coniferous genera.
HISTORICAL CONCEPTS OF CONIFER
SYSTEMATICS AND PHYLOGENY
The history of conifer studies shows somewhat closer relationship to the
history of zoological systematics (at least in some groups) than to that of
angiosperm systematics. The reasons for the similarity are precisely those that
make conifers well suited for a cladistic analysis. First, gymnosperms, including
conifers, have a clear fossil record compared to angiosperms (Florin, 1951;
Stewart, 1983). Their remains are well preserved and have yielded a great deal
of information. The relative antiquity of gymnosperms was realized very early.
Brongniart (1849) recognized three principal plant groups—cryptogams, gym-
nosperms, and angiosperms—thought to follow one another in time and in a
progression from “lower” to “higher” forms. Second, early anatomical and
evelopmental studies of vegetative and reproductive structures have proved
useful in elucidating relationships among conifers. Anatomical studies have
also been employed in demonstrating relationships to other fossil and living
groups of gymnosperms (Strasburger, 1872, 1878, 1879; Bertrand, 1879; Coul-
ter, 1909; Buchholz, 1918, 1920, 1933, 1939, 1941: Jeffrey, 1926; Phillips,
1941; Greguss, 1955). Third, the small number of coniferous taxa, together
with their economic and horticultural importance, has permitted botanists (e.g.,
Chamberlain, 1935: Sporne, 1965) to stress comparative biology more than
species identification based on external morphology. Since the quantity is small,
however, it is surprising that so few systematic revisions (for example, Shaw,
1987] HART, CONIFERS 2d
1914; De Laubenfels, 1969; Liu, 1971; and Liu & Su, 1983) have been com-
pleted.
Evolutionary hypotheses concerning conifers have been characterized by
attempts to link extant groups in evolutionary time, very different relative
importances attributed to characters, preconceived notions of the nature of
evolution or evolutionary trends, and ideas regarding correlation of characters.
The result has been confusion in determining phylogenetic relationships and
classification.
LINKING ExTANT GROUPS IN EVOLUTIONARY TIME
A common problem, not unique to phylogenetic studies on conifers, has
been the tendency to link extant groups in evolutionary time, an apparent
holdover from the ancient scala naturae or “great chain of being’? theme
(Lovejoy, 1936). Living taxa, instead of characters, are viewed as either ad-
vanced or primitive. There are numerous examples in the systematics of both
gymnosperms and conifers. For example, Eichler (1889) considered the Tax-
aceae advanced, while Penhallow (1907) considered them primitive. Other
families and genera—Abietinae (= Pinaceae) (Jeffrey, 1917), Podocarpaceae
(Sporne, 1965), and Phyllocladus (Core, 1955; Keng, 1973, 1975)—have been
chosen as the most “primitive.” Similarly, some groups such as the Taxodiaceae
are considered relicts, while others such as the Cupressaceae are considered
progressive (De Laubenfels, 1965). A few early morphologists saw the fallacy
of lining up living taxa in this manner. Coulter (1909, p. 92) correctly remarked
that “living forms .. . do not represent a series, but the ends of many series.”
SPECIALIZATION OF RESEARCH
Gymnosperm biologists have often specialized in particular aspects of the
plant body or life cycle. While many interesting studies have resulted from this
approach, an unfortunate outcome has been systematic and phylogenetic spec-
ulation based on limited subsets of characters. Chamberlain (1935, p. 230)
aptly stated that, ‘““The grouping into families and the sequence of families and
genera will depend upon each investigator. If he is an anatomist, anatomy will
determine the grouping and sequence. . . . If the gametophytes are emphasized,
there will be still another arrangement.”
Examples of single-character analyses in conifer studies are common. The
most frequently emphasized set of characters has involved the ovulate cone.
For example, Celakovsky (fide Florin, 1955) assumed that the Pinaceae, Tax-
odiaceae, Cupressaceae, and Araucariaceae constitute a phylogenetic series
based on increasing fusion of the bract and scale. The principal classification
followed today is that of Pilger (1926); it is based primarily on the structure
of the ovulate cone (although vegetative characters were also used).
The excessive attention paid to the ovulate cone structure is evident in the
debate about the status of conifers without “evident” cones. Pilger’s (1903)
monograph on the Taxaceae included the conifers without (evident) cones; he
later (1926) divided this group into the Taxaceae sensu stricto, the Cephalo-
212 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
taxaceae, and the Podocarpaceae. Sahni (1920) proposed an independent order,
the Taxales, of equivalent rank with the Ginkgoales, the Cordianthales, and
the Coniferales. Florin (1948b) also concluded that the taxads should be seg-
regated from the rest of Pilger’s families; he therefore placed them in the
separate order Taxales. He maintained that the taxads are distinct from the
conifers and traced their more immediate ancestry not to the Cordaitales but
to the Devonian Psilophytales. His principal evidence was that both living and
fossil members of the Taxales and the Psilophytales have a solitary ovule that
is a direct continuation of the axis (uniaxial). Thus, the uniovulate strobilus of
the Taxaceae was considered primitive rather than derived. Florin (1951) main-
tained that in the Podocarpaceae, in contrast, the uniovulate strobili are in-
dependently derived from taxa with multiovulate strobili. Others are reluctant
to accept Florin’s separation of the taxads from the rest of the conifers, at least
at the ordinal rank. Chamberlain (1935) and Takhtajan (1953) have suggested
that the uniovulate, uniaxial strobilus of taxads is derived from the multi-
ovulate, biaxial cone. The argument becomes dangerously circular when the
very character wl evolution 1s being discussed has been used as the principal
line of evidence in forming the groups under discussion.
Other subsets of characters have been used to a lesser extent as the basis of
phylogenetic and systematic speculation. Saxton (1913) and Moseley (1943)
produced classifications based entirely on characters of the gametophyte and
the embryo. Thomson and Sifton (1926) thought the Pinaceae to be the most
highly evolved of conifers on the basis of the arrangement and structure of
resin canals. Flory (1936) proposed a phylogeny using chromosome numbers.
Praeger and colleagues (1976), relying on antigenic distances, suggested rela-
tionships among genera of Pinaceae.
Finally, as an extension of this approach, relationships of entire families of
conifers are occasionally suggested based on characters found only in a few
taxa. For example, the peltate, perisporangiate microsporophyll is often at-
tributed to all Taxaceae (Stewart, 1983), although it is found only in Taxus
L. and Pseudotaxus Cheng.
PRECONCEIVED NoTIONS OF How EvoLuTION WorKS
Interpretations of the evolution of conifers have been influenced by general
notions of evolution. Florin (1951) made use of Zimmerman’s (1930) telome
theory to explain various aspects of the evolution of the ovulate cone of conifers.
Jeffrey’s (1917) three canons of comparative anatomy include the doctrine of
conservative organs, which considered the leaf, reproductive axis, root, first
annual ring of the stem, seedlings, and sporangia as “‘conservative.” This idea
was apparently borrowed from zoological embryology, in which it was thought
that ancestral features, such as gill slits, are apt to persist in the earlier stages.
Ideas about complexity have also influenced perceptions of relationships. Pen-
hallow (1907) claimed that resin canals are more advanced than resin cells
since they are more complex. Other preconceived theories can lead to just the
opposite results. Jeffrey (1905) believed that resin canals disappear and are
replaced by resin cells.
1987] HART, CONIFERS 273
Another of Jeffrey’s (1917) canons of comparative anatomy was the doc-
trine of reversion, in which wounding induces ancestral traits. The presence of
resin canals after wounding was thus seen to be a reversion to a more primitive
condition. Celakovsky (1890) also argued that teratological structures and wound
tissues indicate evolutionary direction. Guédés and Dupuy (1974) observed
hypertrophied, leaflike segments of ovulate cone scales and interpreted the
ovules to be dorsal appendages (‘‘Ieaves’’) of scale components. Chamberlain
(1935) thought that the occasional abnormal occurrence of bisporangiate cones
represent the ancestral state.
Botanists have long ranked characters according to preconceived notions of
adaptive significance. Adaptive characters have generally been considered less
useful at higher (less inclusive) taxonomic categories than at lower (more in-
clusive) ones (Stevens, 1980). Saxton (1913) thought that the stability of plant
parts or organs is proportional to their distance from the surface of the plant
and their proximity to, or connection with, the reproductive structures. Thus
the external characters of the vegetative organs, such as shape and position of
leaves—characters most susceptible to adaptive change—are less important
than those of the reproductive structures (e.g., micro- and megagametophytes),
embryology, and the internal anatomy of vegetative structures (such as the
vascular system). Lawson (1907) similarly thought that various reproductive
structures of conifers that are buried deep within the tissues of the sporophyte
are less likely to be modified by external factors and more likely to preserve
ancestral characters. Coulter (1909, p. 86) believed that gymnosperm leaves
respond to “conditions of living” and so largely ignored them in his taxonomic
studies. Holgar Erdtman (1963) emphasized the taxonomic importance of con-
stituents excreted into dead conifer heartwood as metabolic end products since
he believed they were not subject to external influence.
CORRELATION OF CHARACTERS
The notion of correlation of characters has been common in conifer studies.
Gaussen (1944, 1950) believed that the most recent species of a group are
generally more evolved in all characters than were their ancestors. Stevens
(1980) aptly pointed out that character states may occur in any combination:
all primitive, all derived, or mixed.
A somewhat more reasonable class of correlations comprises functional ones.
Sporne (1965) noted that the loss of the pollination drop is correlated with the
loss of pollen wings. Coulter (1909) suggested that the position of the arche-
gonium is related to the position of the pollen tube that reaches the embryo
sac before the archegonial initials are evident.
Given such diverse views on how to classify organisms, the importance
attributed to certain characters by some botanists, and how evolution is thought
to proceed, it is littke wonder that attempts at reconstructing phylogenetic
relationships have been stuck in a morass of confusion, contradiction, uncer-
tainty, and appeal to authority.
274 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
CLADISTIC THEORY
Several excellent discussions of cladistic methods now exist (e.g., Hennig,
1966; Hecht & Edwards, 1977; Wiley, 1981; Bremer, 1983). In a cladistic
analysis, certain conditions are sought: the group being studied must be mono-
phyletic, characters selected must be homologous (inherited from a common
ancestor), there must be a known outgroup, and character states must be
designated as either primitive or derived (Arnold, 1981). Hull (1967) and others
have pointed out that there is not necessarily a precise order or progression in
cladistic analysis. A systematist may work at several levels of analysis simul-
taneously.
Initially, a group being studied may not be known to be monophyletic. In
this situation, a group may be selected based on previous taxonomic judgments
or phenetic similarity.
Characters are recognized by similarity of structure in different Organisms,
Recently there has been considerable discussi nabout ch t d homology
(Sattler, 1984; Stevens, 1984; Tomlinson, 1984). During the first stages of
phylogenetic reconstruction, it is not known if the characters are homologous
in the cladistic sense (i.e., equivalent to apomorphies—see Patterson, 1982:
Stevens, 1984). Homologies should, however, meet several opie Boer
ocation, similarity, and connection of intermediate forms (Remane, 1952).
Patterson (1982) recommended three tests of homology: similarity cer
ic, ontogenetic, compositional), congruence (with other hypothesized homol-
ogies), and conjunction (two homologues cannot coexist in the same organism).
Of these, the criterion of similarity is the first and thus the most important—
the tests of congruence and conjunction can be applied only after an initial
determination of the similarity of characters (Stevens, 1984).
Distinguishing between primitive and derived characters is one of the critical
problems in phylogenetic reconstruction. Recently, attention has been devoted
to the criteria by which this distinction 1s made (e.g., Crisci & Stuessy, 1980;
Stevens, 1980; Watrous & Wheeler, 1981; Maddison et a/., 1984). Outgroup
analysis based on parsimony is considered to be the most defensible criterion
(Stevens, 1980). Wiley (1981, p. 139) defined the outgroup rule as follows:
“Given two characters that are homologous and found within a single mono-
phyletic group ... the character found only within the monophyletic group is
the apomorphic character.” The underlying methodological principle of the
outgroup rule is parsimony. The simplest hypothesis—the one that minimizes
the number of parallelisms and convergences (homoplasy)—is preferred (Ste-
vens, 1980; Farris et a/., 1982). This means that the preferred tree is congruent
with the majority of apparent apomorphies. The use of parsimony does not
mean that homoplasy is rare or uninteresting; it only seeks to minimize it.
>
MATERIALS AND METHODS
This study was based on a literature survey, an examination of herbarium
specimens, and observations of living plants. The 63 genera of conifers used
in the analysis were selected from the treatments of Dallimore and colleagues
1987] HART, CONIFERS yg)
(1966), Quinn (1970), and Silba (1984). I chose a set of characters using three
criteria: a reasonable argument of similarity could be made supporting the
homology of the different states of the character; character-state transforma-
tions could be determined on the basis of outgroup analysis; and character
states could provide discrimination of families and genera (see APPENDIX, TA-
BLE) (Rodman et al., 1984). Characters or character states unique to individual
genera (autapomorphies) were not included in the analysis. Morphological and
anatomical information from all aspects of the life cycle, as well as chemical
and chromosomal data, was utilized to avoid favoring certain subsets of char-
acters.
A number of characters were not used for a variety of reasons, one of the
most common being insufficient sampling. Quantitative characters showing
apparently continuous variation or considerable overlap between possible states
were avoided as much as possible (Almeida & Bisby, 1984; Hart, 1985). Char-
acters showing considerable overlap between taxa were excluded. On some
occasions when derived character states were rare and when the character was
not recorded in many taxa, I assumed the primitive condition for missing
characters (e.g., characters 75 and 76).
Different classifications of characters are often found in the literature. Thus
Ueno’s (1960) classification of pollen (character 61) based on extensive sam-
pling using light microscopy differs somewhat from Reyre’s (1968; character
62) system based on a more limited sampling using scanning electron micros-
copy. In this situation I have used Reyre’s system but have included Ueno’s
in the TABLE for purposes of comparison.
Binary as well as multistate coding was used. The number 0 (primitive or
plesiomorphic) was assigned to the character state found in one or all of the
outgroups. With multistate coding, both unordered and ordered coding were
used (APPENDIX, TABLE), depending upon whether or not there was justification
for a transformation series. For example, leaves tetragonal in cross section
(character 28) are found in the fossil conifer outgroups, and a variety of shapes
are found among modern conifers (De Laubenfels, 1953); @ priori, it is not
possible to determine a transformation series of bifacially flattened, scalelike,
or needlelike leaves. In certain situations it was possible to justify a transfor-
mation series. Thus, the presence of specialized winter bud scales (character
37) can be interpreted as having had intermediate steps in evolution.
The PAUP program used in the analysis allows for the coding of missing
data (“9” in TABLE), treating them as equivalent to “all possible states.” The
missing states are filled in by the program according to what would be the most
parsimonious character states, had they not been missing, and the tree length
is then computed. Variable character states were also coded as “missing”
(9).
A data matrix including 63 genera and 123 characters was assembled. Since
current programs such as Swofford’s PAUP cannot guarantee parsimony with
such a large data matrix, the information was broken up into several smaller
units. The first was a family-level analysis using eight representative genera:
Taxus (Taxaceae), Cephalotaxus Sieb. & Zucc. ex Endl. (Cephalotaxaceae),
Araucaria Juss. (Araucariaceae), Podocarpus L’Hér. ex Pers. (Podocarpaceae),
276
JOURNAL OF THE ARNOLD ARBORETUM
[VOL. 68
Data matrix for character states of conifers and outgroup gymnosperms listed in
Appendix.*
5 0 15 20 6 59 3 40 45 SO 55 60
0100000090000000000000000008 1 000008009000200000 10980008 1000000
Ginkgo
Cordaitales 000000990990000900000900990800900000999992999990098 1 1190900000
Lebachiaceac 00099999099000090000090099 100090000000999099999010100000900900
10000000101 1100000000100001 1010000004 1000000000111110101120120
Austrotaxus 900000001001 1 10000000001 11190101920190
Pseudotaxu 000000000001 100000010100001 1010100004 10000000001 11011101920190
Taxus 000000000001 100000010100001 1090100004 10000000001 11011101120120
Torreya 100000001011 100000010100001 1010000004 100000000011 1010101120120
Cephalotaxus
Agathis 000019000000000 1 000000000101090000004 100020000091 1000101010133
aria 0000 19000000000 10000000091 1100000000 100002000009 10000101010133
Acmopyle 00000000100 100000000000000 1300000000290000000001 10000001 000000
Dacrycarpus 00000000100 100000000000000 1200000000 100000000001 10000001 000000
Dacry: 00000000 100100000000000000 1 900000000 100000000001 10000001 000000
000000001001 1100000000300009000001 11900001000000
Falcatifolium 00000000 1001. 00000000000000 1 300000000200000000001 10000001 000000
alocarpus 120000000000000000000 11 1
Lagarostrobos 00000000000 120000000000000000000 1 10000001 000000
epidothamnus 00000000000 10000000001 0000 120000000000000000000 1 1 1
Microcachrys 1202000000020000000001 10000001 000000
icrostr 00000000101 100000001 100000120000000002000000000010000001000000
Parasitaxus 1001 00000000000000 12 1
hyllocladus 01000000000 100000000000000 18000000002 100000000011 1100001000000
Podoc Ss 00000000 100 100000000000000 110000000041 11100001
rumnopitys 00000000100 1000000000000001 1000000003 1000000000111 100001000000
axegothaea 100000001 10100000001 9000101 1000000002 1000000000011 100001010000
Abies 00010101 111100000001 1100101 10002001049001 11000001 1000021000000
Cathaya 010101001111010019111010001 1000200104 111111000001 1000001000090
Cc S 010101 100000111111010101001000040101 1 10001011000001000000
Keteleeria 00010101111100001101 1100001 10002001049001 11000001 1100021000000
rix 010101101111900010111110101110010001401011 1000001 1000001 110020
Picea 10008 1190001011111110190001000049191110001011000001
Pinus 01010100081 100001019901900142 11191 112 1 000000
Pseudolarix 01010101111100000001 11000011 1001000141001 11000001 1100021000000
Re ee ee
Tsuga 000101001 111000000011 110001 000049001 11000001 1000021000010
throtaxis 000000001101 190000000012000000000010000001030122
“ryptomeria Dain rai -snveouacIG Docu boeocHes lauonapioOn meron
-unninghamia Ge 10000000100010200101031122
alyptostrobus 011000001 19 100000001010000190000000020000000101010000101031122
Metasequoia 1¥99x019%eo00co;0000 cpp 110101 1031122
Sciadopitys 000010000001 15000000002 100000000 1010300001020119
Sequoia 000000001 19100000 1010090001 910000000020000000100010000101031122
Sequoiadendron 000000001 191000000010090001 200000000: 10000101031122
Taiwania 00000000110 Cm
Taxodium 011000001 1110000000101000019000000002000000010001110010112
\ctinostrobus 100100000000000000120300000000¢ Patenennustraran naa ones
Aust rus baboons 2201 oa oxBREsNDIE 0101030121
allitris 000000001001 sesearTereit ea
spaalee {000000115 OBO D000 20 sa0 ecococosooLl ouibc imitans
000000001 99 10000000 1090000 120200100000000909 1 10010010101030121
- 191100000001 1900001 20200900000000901110010010101030121
Diselma 00000000 199 100000001 11000012021 10010001
Fitzroya 000000001 19100000009 1900001203 10000000000900900110010101030121
Fokienia 00000000 119100000001 0000001202001 00000000900900010010101030121
iperu: 000000001 1110000000011 1 0111091901010103012
00000000 1001000000091 1202001 10000000900900010010101030121
Aicrobiot 100000001090000120200000000000900990010010101030121
allitropsis 00000000 1001.000000000000001 102000000000009009000100101 1
ap 1s 100100000000000000 1202001 100000009009 10010010101030121
ilgerodendron 000000001091 00001202 10000002000900900010010101030121
latycl 00000000119 100000001010000120200¢ 1009 10010010101030121
etraclinis 1 20: 000000009009 10010010101030121
uja 000000001 19100100001010000120200100000000900910010010101030121
Thujopsis earns dic tied tctenen Coa 10010010101030121
Viddringtonia 00000000 100 1 00000000000000 120200000000000900100010010101030121
or ae
= T r
h Leahle hy &
*p]
states by 9 pp
1987]
Data matrix for character
HART, CONIFERS
f ‘ff, A t 1 ee
(continued)
Ginkgo
Cordaitales
Lebachiaceae
6 7 7 80 8 © 9 100 10 110 115 120
00 888888880081000810000
0000909900099909099000999989999090980000000008800081 190899989
0000999900099909099000999989999999900 1 00000009000080090809989
Amenotaxus
Austrotaxus
Pseudotaxus
Taxus
Torreya
901 101000000001 000000002010000000001 18888888888880 12001110008
901901000000001000000002010000000001 1888888888000002001010008
10190100000000 10000000020100000000011 88888000002001010078
101101000000001 01000000001 1 1888888888000012001110008
Cephalotaxus
Agathis
Araucaria
ssocarpus
Falcatifolium
Halocarpus
Lagarostrobos
eee
i
mnopitys
eee
nus
Pseudolarix
Pseudotsuga
Tsuga
101102001 10000901 100100101001 111000002 10000000000000090000008
1102001 10000901 100100101001 111000002 10000000000000090000908
0000020000000090001000020101 0000009 10200001010010001000010098
0008
002001000000 1000001100020101
009009000000009000 1000020101 00000090020000101000000000001
jen20ror oro ese ee0eeo1 co
000002000000000000 10000201 010000001 1020000101 000000000001 009
000002000000000000 10000201 010000009 1020000101000000000001 ne
00000201010000000010000201 0100000001 0200001 101 10000000001 0008
010000101001000100000003 1 1100000000 10201000000001 100110001110
0000001010010001 00000003 11 100000001 102000000000000001 10000110
1010010001 00000003 1 1100000001 10200000000001 100110001110
Peat ae
10100010101 1000100000003 1 11000000001 02000000000009001 1
0000000311 100000001 102010000000000001 10001110
\throtaxis
equoiadendron
Taiwani
axodium
peaer ors
ee
ane
Se
Thujopsis
Widdringtonia
00110101000000100001 000401 1 110000011
00119101000000100001 000301 000000001002 100000000000001 10000011
001 10101000000100001 000301 000000001002 10000000000000
00110101000000100001000301000000001002101000000000001 10000011
110100000000000000000201 000000001 002 100000000000001 10000002
0011010100000010000101 150200000000 1002 101000000000001 10000111
00110101000000000001 01 130100000000 1002 101000000000001 10000001
00110101000000100001 000301 000000001 002 100000000000001 10000011
00119101000000100001000301000000001 10210100000000002 110000111
0011110100000110000101040200000011110210010000000002110009091
00110101000000100001090301000000001 002 10010000000002 110009011
0111101000001 10000101040200000011 110210010000000002 1 10009091
0011010100000010000100030100000000 1002 10000000000002 1 10009091
00111101000000100001 000301 000000001002 10100000000002 110009011
00111101000000100001000301000000001 10210100000000002 110009191
0011010100000010000109030100000009 10010000000002 1100091091
00111101000000100001090301
00111101000000100001000301000000001 902 1010000000000201 0019091
00110101000000100001090301 00000000 1 002 10010000000002 110009091
001 10101000000 100001090301 0000000010021 ospecieeeac pd
001 10101000000100001090301 00000000 1002 10010000000002 11000909
SEGRE EESTI RTGTECVER RTE TA
001101010000001 psi ecco 10010000000002 110009011
0011010100000010000 301000000001 002 10000000000002010019011
cr oreo enone eneeeReC oz 00 eo
00110101000000100001 10000000000002 110009011
Bis 1G TOOL BIT ORSUT oR GOCSOT 0000000000002 110009011
001111010000001000010104020000001 1110210010000000002 110009001
278 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Pinus L. (Pinaceae), Taxodium Rich. (Taxodiaceae), Cupressus L. (Cupressa-
ceae), and Sciadopitys Sieb. & Zucc. Sciadopitys was added to the list since it
does not seem to share obvious synapomorphies with the Taxodiaceae, with
which it is normally associated. In this analysis the characters chosen for the
representative genera were consistent (with minor exceptions) within the family
but varied across the families. This analysis was conducted using the branch-
and-bound algorithm (Hendy & Penny, 1982). Next, a series of analyses of the
separate families, such as Pinaceae, Podocarpaceae, and Taxaceae, or pairs of
families, such as Taxodiaceae and Cupressaceae, was run. These analyses were
conducted using the local-branch-swapping algorithm.
The selection of outgroups requires some discussion. The Lebachiaceae,
Cordaites Unger, Ginkgo L., and other gymnosperms were chosen as outgroups
(see FiGure 1). For many characters, only the living gymnosperms—Ginkgo,
cycads, and the Gnetales—could be used as outgroups. Other characters were
represented in the fossil record. Paleobotanists generally accept the family
Lebachiaceae— which includes Lebachia Florin, Ernestiodendron Florin, and
Walchiostrobus Florin—as the “stem” conifer group (Florin, 1951). It is, how-
ever, not certain that the ““Lebachiaceae” represent a monophyletic group; C.
N. Miller (pers. comm.) indicated that the family is paraphyletic and thus
constitutes a series of outgroups. For some characters the various genera of
“Lebachiaceae” were individually used as outgroups. On the other hand, the
family Voltziaceae Florin—including Pseudovoltzia Florin, Ullmannia Gop-
pert, and G/yptolepis Schimper—seems to comprise taxa intermediate between
the Lebachiaceae and modern conifers (Stewart, 1983); these were not used as
outgroups since they may be ingroups to conifers. The next outgroup chosen,
Cordaites, is generally acknowledged to be represented earlier in the fossil record
than Lebachia and its relatives and is considered to share a common ancestor
with them (Florin, 1951; Taylor, 1981; Stewart, 1983; Clement-Westerhof,
1984; Mapes & Rothwell, 1984). The position of Ginkgo and then cycads as
the next most inclusive outgroups is supported by the work of Meyen (1984),
Doyle and Donoghue (1986), and Crane (1985). Occasionally it was possible
to use the initial cladogram of the families of conifers to determine polarity of
particular characters (Watrous & Wheeler, 1981). Thus, the presence of inverted
ovules in the Pinaceae, which seem to form a basal clade or functional outgroup
(FiGuRE 2), and in many members of the Lebachiaceae lent credibility to the
polarity of this character. In determining the polarity of the characters generally,
the algorithm developed by Maddison and colleagues (1984) was followed.
RESULTS
In this section I describe the results of attempts to analyze relationships 1)
of conifers to other gymnosperms, 2) among families of conifers, and 3) among
the genera of conifers within the different families. A complete resolution of
the cladistic relationships among the genera and families of conifers requires
more data. However, several hypotheses of phylogenetic relationships can be
proposed with the information available.
In the larger data sets, only the most parsimonious cladograms—those with
1987] HART, CONIFERS 219
Other Ext
ant
Gymnosperms Ginkgo Cordaitales Lebachiaceae Conifers
ar
urE |. Hypothesized relationships of modern conifers to outgroups, including
fossil and living gymnosperms, used as basis for polarization of character states. For
some characters other taxa related to Lebachia used as outgroups intermediate between
modern conifers and Cordaitales.
the fewest reversals, parallelisms, and convergences—are presented. The branch-
and-bound algorithm, which generates the most parsimonious cladograms, can
only work with smaller data sets. This algorithm was used solely in the family-
level analyses and for the Taxaceae. The other data sets were analyzed using
the local-branch-swapping algorithm, which unfortunately does not generate
most parsimonious cladograms. A basis for comparing parsimony among
cladograms is the consistency index, which is the minimum range of character-
state changes in the data divided by the actual length of the tree—or the sum
of character-state changes or patristic distances along all branches. Fractions
close to unity indicate a tree with little homoplasy (Kluge & Farris, 1969).
MONOPHYLY OF CONIFERS AND PHYLOGENETIC
RELATIONSHIPS WITH OTHER GYMNOSPERMS
A manually generated cladistic hypothesis for the monophyly of living co-
nifers and the relationships of these ae with fossil and living gymnosperm
outgroups is presented in FiGure 1. The distinguishing eeu that
separate extant conifers from all ae extant g
and hence suggest monophyly —are embryological. There are at least two char-
acters of importance. First, the number of free nuclear divisions in embryo-
280 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Taxa Ceph Arau Scia Taxo Cupr Podo Pina
68"
115"
Ln ae a oes 54
mmpee 58-3
mmm 68-2
13
30-1 am aca i eas
57 = as"
63 — 116
114°
mes 62-3
=e 101
mee 116
mee 119
ome 54
mes 66
meee 58-2
mes 60)
ge 65
me 68-1
68-2
74
119
86-1
URE 2. Hypothesized relationships between families of conifers, using represen-
tative ete (Arau = Araucaria (Araucariaceae); Ceph = Cephalotaxus (Cephalotaxa-
ceae); Cupr = Cupressus ps sae Pina = Pinus (Pinaceae); Podo = Podocarpus
aaesenieccnea Scia = S$ iadopitys; Taxa = Taxus (Taxaceae); Taxo = Taxodiaceae:
*= ersal; ' = one Sia = character evolved twice.)
genesis (character 86) 1s greatly reduced in living conifers (five or fewer) com-
pared to Ginkgo and cycads (eight and ten, respectively). Second, the structure
of the proembryo of conifers (character 88) is unique. In contrast to the proem-
bryo of cycads and Ginkgo, which is characterized by an unstratified cell ar-
rangement, that of conifers is stratified or tiered. The proembryo of Gnetum
L. differs from them in having no free nuclear stage and no definite arrangement
of cells, and in the elongation of each cell to form a suspensor (Johansen, 1950).
In conifers the primary proembryo is the first cellular structure formed after
the wall. It has two morphological units: an open tier and a lower primary
embryonal cell group (Chowdhurry, 1962; Dogra, 1978). This is characteristic
of nearly all conifers, including the Araucariaceae (Haines & Prakash, 1980)
and the Taxaceae (Chen & Wang, 1984). Since these characters are not known
for the Cordaitales or the Lebachiaceae, they may be placed at one of three
Taxa Ceph Scia Arau Taxod Cupre Pina Podo Taxa Ceph Arau Scia Taxod Cupr Podo Pina oO
oo
=
Taxa Ceph Scia Arau Taxod Cupr Podo Pina ae
>
ve)
7
Taxa Ceph Arau Taxod Cupre_ Scia Pina Podo a)
Z
—
|
™
A
"A
Ficure 3. Hypothesized relationships of families, using representative genera; 4 cladograms involving | more step than in FiGure 2.
18¢
282 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
nodes in the clades in FiGureE |: extant conifers; extant conifers + Lebachiaceae;
or extant conifers + Lebachiaceae + Cordaitales.
Other characters can be used to establish monophyly and outgroup rela-
tionships when fossil gymnosperms are used for comparison. Extant conifers
can be distinguished from the fossil Lebachia by at least two characters. One
is the cone scale (character 100-2), a highly modified fertile short shoot (Florin,
1951; Taylor, 1981; Stewart, 1983; Meyen, 1984; Crane, 1985). Crane (1985)
stated that the ovulate fertile short shoot—or “‘scale’’— of extant conifers differs
from that of the Lebachiaceae in that the shoot apex is not differentiated and
that there is no phyllotactic spiral in parts of the former. There is still consid-
erable discussion as to exactly what it represents: for example, short shoot alone
or short shoot plus sterile scale (Guédés & Dupuy, 1974; Jain, 1976). However,
the exact nature of the structure does not affect my argument as long as part
of the scale is a short shoot.
The second character 1s palynological. Pollen of modern conifers is char-
acterized by distal germination, whereas that of the Lebachiaceae does not have
a thin area on the distal surface, thus indicating proximal germination (Mapes
& Rothwell, 1984). This character shows homoplasy; Millay and Taylor (1976)
have shown that the shift from proximal to distal germination also occurred
in the Callistophytaceae and the Cordaitales.
If Cordaites is considered as the outgroup to conifers (Doyle & Donoghue,
1986), a number of derived characters support monophyly of the Lebachia-
ceae + extant conifers. The pollen cones (character 49) of the Lebachiaceae
and modern conifers are simple or uniaxial; those of Cordaites are compound.
Conifer leaves—‘‘microphylls” (character 27)—are rather small and usually
single veined (except in the Araucariaceae and a few species of the Podocar-
paceae); the leaves of the Cordaitales, Ginkgo, and the cycads are rather large
and many veined. The Lebachiaceae (except a few species of genera such as
Ernestiodendron) and extant conifers have bilaterally flattened ovulate short
shoots (or scales); the Cordaitales have radially symmetrical fertile ovulate
short shoots (Florin, 1951; Taylor, 1981; Rothwell, 1982; Stewart, 1983).
Ovule orientation (character 114) is a difficult character to employ because
it is variable in some groups. The ovule is erect in Ginkgo, the cycads, Ephedra
L., and Gnetum. The most recent interpretation for the Voltziales is that most
have inverted ovules (Clement-Westerhof, 1984; Mapes & Rothwell, 1984).
Crane (1985) also suggested resin canals as a synapomorphy for Lebachia
and extant conifers. Resin canals do occur in nearly all conifers and taxads,
although in many different plant parts (i.e., xylem, roots, leaves, seed coats);
this may suggest different origins (homoplasy). Mucilage canals have been
described for Ginkgo and may be similar to resin canals in conifers. Studies of
resin-duct development and resin chemistry may help our understanding of
these characters.
FAMILY-LEVEL ANALYSIS
In this analysis the characters chosen for the representative genera were
consistent (with minor exceptions) within the family but varied across the
1987] HART, CONIFERS 283
Abies Ketel Plarix Tsuga Cedrus Larix Ptsuga Catha Pinus Picea
mms 24
mee 61
wee 1()2'
me 97
wes 124"
39*
55-2
102'
10'
71
120
4,6, 19, 22, 23,
28, 32,37, 39, 41, 42,
43, 50, 69, 74, 78, 87,
89,116, 122
Figure 4. Cladistic relationships of Pinaceae. (Catha = C ‘athaya; Ketel = pe ona:
eee = aa larix, Ptsuga = Pseudotsuga; * = reversal; ' = one parallelism;
character evolved twice; ”’ = character evolved three times; "’ = character evolved four
imes.)
families. Thus the characters were the important consideration, the genera being
chosen merely to represent them. FiGure 2 shows the results of the family-
level analysis, which employed 22 characters and representatives of the seven
as Sciadopitys (included because it differs in so many characters from the
Taxodiaceae, in which it is normally placed, that it has sometimes been put
in other families—e.g., Pinaceae, Saxton, 1913; Sciadopityaceae Hayata, Hay-
ata, 1932). The consistency index 1s .711.
Four additional trees, each with one extra step (consistency index of .659),
were generated (see FiGurE 3). In all of these, the Taxaceae and the Cepha-
lotaxaceae came out as sister taxa, as did the Taxodiaceae and the Cupres-
saceae. Sciadopitys is most often the outgroup to the Cupressaceae and the
Taxodiaceae and is placed there in the subsequent family-level analysis. The
family Pinaceae is most often the outgroup to all living families of conifers.
284 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
The placement of Sciadopitys, the Podocarpaceae, and the Araucariaceae is
variable.
PINACEAE. Ten genera and 48 characters were used in the cladistic analysis of
the Pinaceae (results shown in FiGure 4). The consistency index is .600. Mem-
bers of this family are distinguished by seven synapomorphies restricted to
them: 6 phloem fibers absent), 41 (leaf transfusion-tissue tracheids all around
vascular bundle), 43 (biflavonoids absent), 69 (sperm cells without cell walls),
74 (ventral-canal cells without walls (nuclei only), 78 (megaspore membrane
thin at micropylar end), and 89 (proembryo four-tiered). Several other char-
acters (e.g., resin ducts, character 19), initially scored as derived within the
Pinaceae, are derived at the family level but show subsequent loss in different
lineages. There were numerous other synapomorphies (e.g., character 39) show-
ing homoplasy within conifers that are evidently derived at the family level.
PopocarPaceae. Fifteen genera and 24 characters were used in the analysis of
the Podocarpaceae (results presented in FiGure 5). The consistency index is
.500, rather low. Only two unique synapomorphies seem to unite the Podo-
carpaceae: the binucleate embryonal cell of the proembryo (90), and the epi-
matium (105, but missing in two taxa). Additional apomorphies are found in
other conifers (28-2; 119) or are only found in most Podocarpaceae (e.g., 48);
the algorithm has interpreted them as being derived at the family level but
subsequently lost within the family.
TAXODIACEAE-CUPRESSACEAE. Thirty-one genera and 53 characters of the Cu-
pressaceae and the Taxodiaceae (including Sciadopitys) were analyzed (see
Ficures 6 and 7). The consistency index is .544. Sciadopitys is even more
clearly separated from the Taxodiaceae-Cupressaceae than the family-level
analysis indicated, with 12 synapomorphies separating them. It can be seen
that the Taxodiaceae, even exclusive of Sciadopitys, are paraphyletic. There
are several monophyletic groupings within the Taxodiaceae, including Sequoia
Endl. and Sequoiadendron Buchholz, Metasequoia Miki, Taxodium, and Glyp-
tostrobus Endl.; and Taiwania Hayata, Cryptomeria D. Don, and Cunning-
hamia R. Br. ex Rich.
Several synapomorphies define the Cupressaceae as a monophyletic group
within the Taxodiaceae (see FiGuRE 6). Within the Cupressaceae, there is di-
vision of northern and southern taxa (FiGuRE 7). The analysis indicates that
northern Cupressaceae are paraphyletic although there are several monophy-
letic groupings, including Microbiota Komarov and Platycladus Spach, Thuja
L. and Thujopsis Sieb. & Zucc., Fokienia A. Henry & H. Thomas and Caloce-
drus Kurz, and Juniperus L., Chamaecyparis Spach, and Cupressus. However,
it should be remembered that these hypotheses of relationships are tenuous
since few characters were utilized in the analysis. The southern taxa, including
the African Tetraclinis Masters, form a monophyletic group. This group divides
into an unresolved quadrachotomy: Diselma J. D. Hooker, Fitzroya J. D.
Hooker, and Pi/gerodendron Florin; Austrocedrus Florin & Boutelje, Libocedrus
Endl., and Papuacedrus L.; Neocallitropsis Florin; and Widdringtonia Endl.,
Callitris Vent., and Actinostrobus Miq.
Phyll Saxeg Podo Prumn Decus Acmop_ Dacrd Falca Dacryc Para Haloc Lepid Lagar Microc Micros
37-5 F110" — 117°
ime 107° + 7
— 110'
114-2"
90-1
Ficure 5. Cladistic relationships of Podocarpaceae. (Acmop = Acmopyle, Dacryc = Dacrycarpus, Dacrd = Dacrydium, Decus = Decusso-
carpus; Falca = Falcatifolium, Haloc = Halocarpus, Lagar = Lagarostrobus, Lepid = Lepidothamnus, Microc = Microcachrys; Micros =
Microstrobos, Para = Parasitaxus, Phyll = Phyllocladus, Podo = Podocarpus, Prumn = Prumnopitys; Saxeg = Saxegothaea; * = reversal; ' =
one parallelism; ” = character evolved twice.
SUgadINOO “LYUVWH [L861
S8C
286 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Sciad Athro Sequ Sequd Glypt Metas Taxod Taiwa Crypt Cunni — Cupres
86-4
a8" 37-2"
95' ne
121' mtr 30-2
37*
p= 46
wom 39-2" mpm 62-2
ume 84-2 fe er =m 114
mm 86-4 ba 59"°
mem 97°
ome 37-2"
mp 38-1
meee 50-3
mee 58-2 22'
p= 47' 4"
Z 103"
11
20'
44
54
59
9
=e 10'
= 28-2
== 62-1
=— 71
—p= 77
=e 82
mp 84
=e 86-3
—— 103'
p= 123-2
28-3
37-1
58-3
60
66
67
86-2
97
IGURE
rotax. s; Cryp t= Cryptomeria; Cunni = haa Cupres = Co Gist = =
Cipisarohas Metas = Ve Sciad = Sciadopitys, Sequ = Sequoia; Sequd =
Sequoiadendron, Taiwa = Taiwania, Taxod = Taxodium: * = reversal; ' = one paral-
lelism; " = character evolved twice; '” = character evolved three times.)
TAXACEAE. Five genera and 16 characters were used in the analysis (see FIGURE
8); the consistency index is .857. This family can be recognized at least by the
uniaxial or “‘simple”’ seed “cone” (99). Characters such as the aril (117) are
also found in other families.
ARAUCARIACEAE., This family comprises only two genera (Agathis Salisb. and
Araucaria) and as such does not require a phylogenetic analysis. It is defined
by at least ten apomorphies (FIGURE 8)
1987] HART, CONIFERS 287
Micro Platy Thuja Thujo Fokie Caloc Junip Chama Cupre Tetra Disel Fitzr Pilge Austr Liboc Papua Neoca Widdr Calli Actin
98"
wie 67"
a a v3)
“
eT ee
—™ 119" — 97"
FIGURE 7.
Austrocedrus, Calli = Callitris,; Caloc = Calocedrus; Chama = Chak aecyparis: cae =
Cupressus; Disel = pein ee Fitzroya, Fokie = Fokienia, Junip = Juniperus;
Liboc = Libocedrus, Micro = Microbiota, Neoca = Neocallitropsis; Papua = Papuacedrus,
Pilge = Pilgerodendron, Platy = Platycladus, Tetra = Tetraclinis, Thujo = Thujopsis;
Widdr = Widdringtonia; * = reversal; ' = one parallelism; ” = character evolved twice;
” = character ee three times.)
DISCUSSION
This cladistic analysis of conifers provides explicit criteria for establishing
phylogenetic relationships and classifications based on multiple character sets,
facilitates the understanding of the evolution of characters, illustrates the dis-
tinction between character-state polarity and taxonomic polarity, is helpful in
understanding evolution and biogeography of the group, demonstrates the use-
fulness of fossil gymnosperms as outgroups, and focuses attention to gaps in
knowledge requiring further research.
CLASSIFICATION
The classification of conifers, especially with regard to their relationships
with taxads and other taxa lacking ‘“‘evident” cones, has been much discussed.
The results of this analysis strongly support the monophyly of conifers and
taxads. Traditional approaches to conifer systematics (e.g., Sinnott, 1913; Aase,
1915; Thomson, 1940; Florin, 1951; C. N. Miller, 1976, 1982, 1985) have
288 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Ament Torre Pstax Taxus Autax Agathis Araucaria
53 — i 49!
33 5
62-3
13 58-1
20 68-2
22 83
91
9 92
48 ibe
50 79
99 80
117 16
119 on
FiGurE8. Cladistic relationships of Taxaceae (left) and Araucariaceae (right). (Ament =
Amentotaxus, Autax = Austrotaxus; Pstax = Pseudotaxus; Torre = Torreya, * = reversal,
‘= one parallelism.)
tended to emphasize ovulate cone structure. This study has uncovered em-
bryological, palynological, and anatomical features that also provide bases
for the recognition of conifers as a monophyletic group (see Ficures 1, 2).
The placement of the Taxaceae has been controversial in the past (see, for
example, Chamberlain, 1935; Florin, 1948b, 1951; Takhtajan, 1953; Sporne,
1965). The simple, uniaxial cone—in contrast to the biaxial one found in all
other conifers—is unique to this family, and Florin (1948b, 1951) championed
the separation of the Taxaceae from the rest of the conifers based on this
character alone. He found a similar cone in the Jurassic Paleotaxus jurassica
Florin and concluded that, since this structure is old and thus primitive, the
Taxaceae should therefore be elevated to the rank of Taxales, coordinate with
the Coniferales. In this cladistic analysis the taxads clearly fall out as a sister
group to the Cephalotaxaceae, well within the rest of the conifer families (see
FiGure 2), all of which have biaxial cones. Embryologically, the Taxaceae have
patterns of development similar to those of other conifers—a reduced number
of divisions in embryogenesis and a tiered proembryo. In this analysis the most
parsimonious explanation of the distribution of character states suggests that
the uniaxial ovulate cone is derived from a compound, biaxial one. Florin’s
reason for elevating the Taxaceae is apparently unjustified: although uniaxial
cones apparently similar to those of Taxus are found in the Jurassic, numerous
earlier gymnosperms had biaxial cones.
A close relationship between the Taxodiaceae and the Cupressaceae has been
recognized (e.g., Saxton, 1913; Eckenwalder, 1976; Stewart, 1983), although
1987] HART, CONIFERS 289
an isolated position for Sciadopitys (which is placed in the Taxodiaceae) has
also been suggested (Velenovsky, 1905; Seward, 1919; Florin, 1922; Hayata,
1932: Eckenwalder, 1976; Schlarbaum & Tsuchiya, 1985). The results from
this cladistic analysis support these general conclusions since the monophyly
of the Taxodiaceae (minus Sciadopitys) + Cupressaceae is supported by many
characters (see FiGurE 6). However, the Taxodiaceae as currently recognized
are not monophyletic but paraphyletic, the Cupressaceae form a monophyletic
grouping within that family. Thus, if one chooses to recognize the Cupressaceae
as presently circumscribed at the family rank, then the Taxodiaceae cannot be
recognized, and many clades within the current Taxodiaceae will have to be
elevated to family ranking. A possible solution is to recognize the entire Tax-
odiaceae-Cupressaceae clade as the Cupressaceae, which has nomenclatural
priority (Eckenwalder, 1976).
The monophyly of the Pinaceae is well established (see FiGurE 4), with at
least ten unique synapomorphies. Within the Pinaceae, grouping of genera is
uncertain, as has been suggested by previous workers (e.g., Van Tieghem, 1869;
Jeffrey, 1905; Pilger, 1926; Gaussen, 1966), who have each emphasized different
characters in suggesting relationships. Van Tieghem (1869), for example, di-
vided the family into two groups, those with short shoots and those without
them. My results do not support his division of the family. In my analysis
short shoots have evolved three times: in the lineage giving rise to Pinus,
Cathaya Chun & Kuang, and Larix Link; in Cedrus Trew; and in Pseudolarix
Gordon. Inspection of the morphology of the short shoots suggests differences
between them (Thomson, 1914). Those of Cedrus, Larix, and Pseudolarix are
persistent, and the leaves fall separately on an annual basis or in the second
to fifth year. In Pinus the short shoots are deciduous as an entire unit in the
second to twentieth (rarely to the forty-fifth) year, they produce a fixed number
of needles in a single season, and they are axillary to a scale. In the other genera
of Pinaceae, the needles are not fixed in number, and the short shoots are not
deciduous or axillary to a scale. In Cathaya the short shoots are poorly de-
veloped. However, even acknowledging the differences between short shoots
within the Pinaceae does not tell if they represent the same character or sep-
arately evolved, nonhomologous ones. Phylogenetic hypotheses can assist in
answering such questions: this analysis suggests that short shoots have evolved
three different times and so may not be homologous, yet that the morphological
variation noted by Thomson (1914) may not be relevant in suggesting different
evolutionary origins. Alternatively, if the information given by Thomson 1s
used to record the character, short shoots may have evolved at least four times!
Barnard (1926) claimed that some shoot dimorphism is common in conifers—
another suggestion that short shoots are a weak phylogenetic character.
The grouping of the Pinaceae into two lineages is based on a few characters:
the presence of resin ducts in the seeds (character 120) and of cleavage poly-
embryony (97) supports monophyly of Abies Miller, Pseudolarix, Keteleeria
Carriére, Cedrus, and Tsuga Carriére; resin ducts in the secondary wood (17)
and leaves with endodermis having thickened Casparian strips (39) support
monophyly of Cathaya, Pinus, Larix, Pseudotsuga Carriere, and Picea Dietr.
Singh (1978) listed embryological characters of the Podocarpaceae in addition
290 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
to those used in this analysis; for example, densely staining cytoplasm sur-
rounding the archegonium (character 81). This character, however, needs fur-
ther investigation to verify its use as a character state. De Laubenfels (1962)
suggested that the presence of two cotyledons, each with two vascular bundles,
is a feature unique to the Podocarpaceae. However, the use of this character
does not stand up to cladistic reasoning. The fact that members of the outgroup
comprising Ginkgo, the cycads, and the Gnetales have two cotyledons—and
those of Ginkgo have two vascular bundles—might suggest that this is a prim-
itive character within the Podocarpaceae. The morphological heterogeneity of
the Podocarpaceae is underscored by the variation in chromosome numbers,
which is extreme when compared to that within other conifer families (Sax &
Sax, 1933; Hair & Beuzenberg, 1958; Khoshoo, 1961; Mehra, 1968). Given
the high levels of homoplasy, the groupings of genera within the Podocarpaceae
(FiGurReE 5) must thus be very tentative, and additional research is clearly needed
to confirm them.
Although the Podocarpaceae are usually considered a natural group, Keng
(1973, 1974, 1975) has elevated Phyllocladus to family ranking, suggesting that
the phylloclade of Phyllocladus was a very ancient structure that linked conifers
with progymnosperms. For this to be the case, Phyllocladus would have to fall
out not only as separate from the rest of the Podocarpaceae, but also as splitting
off first in the family-level analysis. This is clearly not the case (see FiGures 2
(Podocarpaceae), 5). Phyllocladus is not only a terminal taxon within the Podo-
carpaceae, but the Podocarpaceae in which it belongs split off after the basal
Pinaceae (FIGURE 2; compare FiGurRE 3).
How does one evaluate a cladogram? A significant quantity of homoplasy
(the amount of parallelisms, convergence, and reversals in character states)
seriously weakens cladistic hypotheses. One measure of homoplasy is the con-
sistency index, which is the minimum range of character-state changes in the
data divided by the actual length of the tree—or the sum of character-state or
patristic changes along all branches. Fractions close to unity indicate a clado-
gram with little homoplasy (Kluge & Farris, 1969). In this study it varied from
.500 to .857, a modestly good figure compared to that in some studies (for
example, .40 in Rodman eft a/l., 1984). There may be several factors—both
artificial and real—that explain the relatively low levels of homoplasy in this
study. Comparing homoplasy indices among different taxonomic groups may
lead to divergent values due to different sizes of data matrices. The greater the
number of taxa and characters, the greater the amount of homoplasy. Thus,
the consistency index for the Cupressaceae-Taxodiaceae analysis, with 31 taxa
and 53 characters, was .544, while that for the Taxaceae analysis, with 5 taxa
and 15 characters, was .857.
There may also be biological reasons why the homoplasy values are com-
paratively low in this study. In groups like conifers, in which great gaps exist
between taxa due to extinction, character states may be comparatively dis-
tinctive, while in some more recent angiosperm groups characters may show
nearly continuous variation, with character-state delimitation correspondingly
uncertain.
Phylogenetic analyses using multiple sets of characters taken from all aspects
1987] HART, CONIFERS 791
of the plant demonstrate the value of not relying on any particular subset of
characters, such as cone structure. We also see, not surprisingly, the importance
of looking beyond the readily visible morphological features. Many of the
phylogenetically useful characters are anatomical, embryological, palynologi-
cal, or chemical. For example, apomorphies for the Pinaceae include p-type
plastids, absence of biflavonoids, arrangement of transfusion-tissue tracheids,
absence of phloem fibers, lack of cell walls in ventral-canal nuclei, thinning at
the micropylar end of the megaspore membrane, and four-tiered proembryo.
But the converse position—that gross morphological characters are not useful
as phylogenetic markers—cannot be maintained. Saxton (1913) and Ecken-
walder (1976) downplayed the value of decussate phyllotaxy that characterize
Cupressaceae, but for different reasons. Saxton (1913) believed that external
morphological characters respond to ‘conditions of living” and are therefore
poor indicators of phylogeny. Although there is some merit in what Saxton
says, a case can be made for the functional nature of just about any structure.
It is best to exclude notions of adaptation and/or function from phylogenetic
analysis, at least in the initial stages. This is not to say that phylogenies based
on characters that seem adaptive should not be questioned.
Eckenwalder (1976) dismissed decussate phyllotaxy as not being a useful
character for the Cupressaceae since it reportedly occurs elsewhere. However,
there are two problems with this position. First, some of Eckenwalder’s ex-
amples of decussate phyllotaxy are not really decussate, but bijugate or spiral
opposite—e.g., Metasequoia and the Taxaceae (Morley, 1948; De Laubenfels,
1953; Greguss, 1955). Second, while perfectly decussate leaves have indeed
evolved elsewhere (e.g., in the Cheirolepidiaceae Takht. (Alvin, 1982) and in
Microcachrys tetragona J. D. Hooker), the usefulness of this character, although
perhaps weakened, cannot be altogether discounted.
UNDERSTANDING THE EVOLUTION OF PARTICULAR CHARACTERS
Cladograms facilitate the understanding of the evolution of particular char-
acters. Florin (1951) argued for a separation of conifers and taxads based on
the single terminal ovule of the latter, which he claimed did not evolve by
reduction from a bract and ovuliferous short-shoot system. The results of this
cladistic analysis suggests, on the contrary, that the ovule structure of the taxads
evolved from the biaxial cone of the conifers. Indeed, Harris (1976) suggested
a possible scenario. An example is the peltate, perisporangiate microsporophyll
of some Taxaceae (Taxus, Pseudotaxus), which has been likened to the spo-
rangiophore of the Cordaitales (Dupler, 1919). Outgroup analysis indicates that
this unique taxad microsporophyll is derived from the bisporangiate, hypo-
sporangiate microsporophyll of other conifers.
“PRIMITIVE”? CHARACTERS VS. ““PRIMITIVE” TAXA
The cladistic results illustrate what to many is a contradiction: the presence
of both specialized and generalized (or primitive) traits within particular taxa,
or heterobathmy (Stevens, 1986). As mentioned above, much early discussion
centered on which of the modern groups of conifers is the most primitive. In
292 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
cladistic reasoning, living taxa are not viewed as primitive or advanced; only
individual characters are advanced or primitive with respect to their condition
in related taxa. Cladograms themselves simply represent the sequence of di-
vergence of lineages. Thus the occurrence of so many derived characters in an
apparently basal clade such as the Pinaceae may seem to be a contradiction,
but it is not unexpected. The cladistic interpretation of the relative age of the
Pinaceae is not inconsistent with the fossil record, which indicates that the
group is very old (C. N. Miller, 1976, 1982; Meyen, 1984).
RInnEcnce ADLIN
The distribution of conifers—both fossil and extant —has long been of interest
to biogeographers. Conifers have been divided into northern and southern
“groups.” Florin (1940, 1963) found that the southern conifer floras were
different from the northern ones as early as the late Carboniferous and Permian
periods.
Li (1953b) discussed the high diversity of extant conifers in the Pacific Basin
and showed that in both Northern and Southern hemispheres, the majority of
relict, endemic, or disjunct genera are concentrated in moist, mountainous
regions with warm temperatures bordering the eastern and western parts of the
Pacific.
In accounting for the distribution of conifers, biogeographers have drawn
upon various explanations: migration and dispersal from centers of origin,
extinction, and continental drift (Florin, 1963).
Seeking centers of origin was a common endeavor for conifer biogeographers,
as it was for other specialists. Brown (1869) concluded that each genus had
arisen out of the center in which the greatest number of species is found.
Conifers were commonly believed to have originated in northern polar regions.
Koch (1927) suggested a European origin for them.
In explaining the disjunct distribution patterns of conifers, biogeographers
generally have suggested that long-distance dispersal has not been as frequent
as in angiosperms. This is expected, given the relatively large size of most
conifer seeds. However, the fleshy propagules of many conifers (e.g., Podocar-
paceae, Taxaceae, Juniperus) are likely candidates for long-distance dispersal,
since birds are known to eat them (Givnish, 1980). Land bridges and connec-
tions have been hypothesized to get conifers from one continent to another.
Florin (1963) postulated that the migration of conifers has occurred in or along
mountain belts during the Paleozoic, Mesozoic, and Cenozoic eras. Continental
drift has often been employed to explain conifer distribution, especially in the
Southern Hemisphere (Florin, 1963; Aubréville, 1973; Page & Clifford, 1981).
Whatever cause for these distribution patterns of conifers one chooses, the
explanation will be influenced—if not determined — by cladistic relationships.
As an example, consider some of the southern Cupressaceae (FIGURE 7). Several
groups show Gondwanaland distributions: Pilgerodendron, F; itzrova (both South
America), and Dise/ma (Tasmania); Austrocedrus (South America), Libocedrus
(New Zealand, New Caledonia), and Papuacedrus (New Guinea); and Callitris,
Actinostrobus (both Australia), and Widdringtonia (southern Africa). Of these,
1987] HART, CONIFERS 293
the first two groupings are somewhat tenuous since they are supported by few
characters, but the clade of Widdringtonia, Callitris, and Actinostrobus is sup-
ported by several. A likely explanation is that the common ancestor of these
genera inhabited Gondwanaland, and with subsequent continental drift these
lineages became recognizable. Florin (1963) contended that the conifers divided
very early into northern and southern groups. The Araucariaceae, the Podo-
carpaceae, Athrotaxis D. Don, Paranocladus Florin, Walkomiella Florin, and
Buriadia A. C. Stewart & B. Sahni constituted the southern group, while the
rest of the conifers constituted the northern one. My cladistic analysis does not
support the contention that modern evolutionary distributions reflect that early
distribution of two groups. It does suggest multiple Gondwanaland distribu-
tions—two in the Taxodiaceae-Cupressaceae clade and one in the Taxaceae.
Many conifer groups (e.g., Araucarites C. Presl, Athrotaxites Unger, and Po-
docarpus, fide Krassilov, 1974) had both northern and southern distributions,
relative to the Tethys Sea, in the Mesozoic. Extinction, perhaps due to changing
climates, may also account for some of the disjunctions, especially in the
Northern Hemisphere.
ROLE OF FOSSILS
Many botanists (e.g., Stevens, 1980, 1984) and some zoologists (e.g., Pat-
terson, 1982) are reluctant to use fossils in polarizing character states. Stevens
(1980, p. 342) stated “*. . the imperfections of the fossil record cast doubt on
this method of giving evolutionary polarity to a morphocline.”” However, the
relevance of fossils depends upon the group being studied (Crane & Manchester,
1982). It may also depend upon the level of grouping in which a systematist
is interested: for example, fossils may be of importance in assessing relation-
ships of conifers to other gymnosperms, or among genera of conifers, but less
useful for species of Podocarpus.
The use of fossils in phylogenetic reconstruction may be questioned some-
what differently: are fossils automatically to be considered ancestors, are they
merely another organism, or are they special outgroups, to be given special
consideration? The answer to the first query should be obvious. Despite re-
peated claims by paleontologists to have discovered the “‘ancestor”’ for partic-
ular groups, it is extremely doubtful that ancestors for many groups will ever
be determined with any certainty.
The answer to the second will be determined by the quality and quantity of
the characters shown by the fossils. Fossils may help greatly in the understand-
ing of characters. Thus Florin (1951) was perfectly justified in discussing the
evolution of cone scales in modern coniferous taxa from short shoots of fossils,
because these characters are well represented in the fossil record.
Should fossil outgroups be given special status—that is, greater importance
than living outgroups? Here there can be no easy solution. 4 priori, fossil
outgroups cannot be given special status over living outgroups. However, it all
depends on the group being studied. Well-represented fossil groups may be
weighted more than isolated living outgroups, or vice versa. Fossil represen-
tatives have been crucial in the phylogenetic analysis of conifers. The use of
294 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
fossil groups like the Lebachiaceae, the Cordaitales, and others puts the cladistic
analysis of living conifers on a much firmer footing than if only other living
gymnosperms were employed for outgroup comparison.
Fossils also tell us something of past distributions. The relictual nature of
many genera of conifers is borne out in studies such as Chaney’s (1951). Se-
quoia, for example, once had a far greater distribution than it does now. Florin
(1940, 1963) used fossil evidence to plot former distributions of conifers on a
global basis. This type of information would never be known from the study
of living taxa.
Despite these manifest benefits of the fossil record, numerous characters are
not readily observable from fossils. Many paleobotanists will be dependent
upon the more enriched data sets available only from living plants.
New RESEARCH
This study has attempted to demonstrate the potential of cladistic analysis
in phylogenetic reconstruction; Hennig’s work (e.g., 1950, 1966) is now taking
root in systematic botany. While much of the current direction in cladistics is
methodological, the basis of phylogenetic hypotheses and evolutionary sce-
narios 1s careful research on the organisms—their characters and character
states. This analysis was possible only because of the careful work of the classical
morphologists— biologists who were greatly motivated by discovering patterns
of evolution (e.g., Thomson, 1905, 1940; Coulter, 1909; Coulter & Chamber-
lain, 1917; Buchholz, 1918, 1920, 1933, 1939, 1941; Chamberlain, 1935). Since
the purpose of this study was to bring together and critically analyze current
information, future research utilizing new techniques is needed to confirm (or
modify) some of the preliminary conclusions presented above.
This future work must develop in two directions. First, new and more com-
plete information is needed. Anatomical analyses have already proven useful
in elucidating phylogenetic relationships, and character analyses using new
techniques should be given priority. Especially needed are more studies of
reproductive biology—such as microgametophyte and megagametophyte de-
velopment, embryology, and palynology—which have already contributed many
characters useful in understanding the phylogeny of conifers (Thomson, 1905;
Buchholz, 1941; Lurzer, 1956; J. Doyle, 1957; Ueno, 1960; Chowdhurry, 1962:
Dogra, 1966, 1978; Pettitt, 1966, 1977; Singh, 1978; Haines & Prakash, 1980).
In particular, studies are needed of the poorly understood tropical and south-
temperate genera in the Podocarpaceae, Cupressaceae, and Araucariaceae, but
many northern taxa, especially those in groups that are not economically im-
portant, also need investigation. A fresh look at characters studied decades ago,
such as the megaspore membrane (Thomson, 1905), is necessary. New ana-
tomical techniques such as ultrathin sectioning and scanning and transmission
electron microscopy can contribute much to character discovery and analysis
and ultimately to phylogenetic reconstruction. We can also look for important
results from biochemical and molecular research (Praeger et al., 1976; Praeger
& Wilson, 1978; Cronin & Sarich, 1980; Sibley & Ahlquist, 1984), but the use
of this approach is not without criticism with respect to inherent assumptions
1987] HART, CONIFERS 295
of the constancy of molecular evolution (‘molecular clock’) and to whether
these kinds of data are amenable to tree construction (Farris et al., 1982; Farris,
1985).
Second, once the information is collected, character states must be analyzed
very carefully before they are incorporated into cladistic analyses. There is
substantial character variation in any group of organisms that is not suitable
for cladistic analysis due to continuous variation or incomplete surveys. AS
mentioned above, careful attention must be given to the gnition of character
states. Polarization of character states may be impossible due to their unknown
status in outgroups. After construction of a cladogram, a second stage of
character evaluation may be necessary in the weighting of functionally corre-
lated characters.
Assumptions of computer programs also need to be addressed. The under-
lying assumption of Swofford’s PAUP program used in this analysis is unre-
stricted parsimony. Characters may be lost, regained, and perhaps lost again.
Unlimited reversals, especially of complicated characters, may be unlikely in
evolution. We might look to the next generation of computer programs to
address this problem.
Third, new paleobotanical information is needed. Much of the past digging
has been conducted near major research institutions in northern regions. It is
not surprising that most fossil conifers—such as Lebachia—are northern in
distribution. No doubt there are as-yet-undiscovered fossils in southern regions
that will cast light on early conifer evolution. Eventually, fossil and modern
taxa will be included in the same analysis.
ACKNOWLEDGMENTS
The author would like to thank the Atkins Garden Fund for support for this
work. Appreciation is also offered to Robert Price and Charles Miller for sharing
data on character-state distribution and some preliminary hypotheses of phy-
logenetic relationships. Constructive criticism of this paper was kindly offered
by W. B. Critchfield, C. N. Miller, P. F. Stevens, A. H. Knoll, and P. B.
Tomlinson.
LITERATURE CITED
Aase, H. C. 1915. ee anatomy of the megasporophylls of conifers. Bot. Gaz.
See 60: 2 13.
ALMEIDA, M. T., & F. A. ee 1984. A simple method for establishing taxonomic
characters ie measurement data. Taxon 33: 405-409.
AxvIn, K. L. 1982. pean biology, structure, and paleoecology. Rev. Pa-
lacobot. Palynol. 37: 71-98.
ARNOLD, E. N. 1981. See phylogenies at low taxonomic levels. Z. Zool. Syst.
35.
AuBREVILLE, A. 1973. Distribution des coniféres dans la Pangée; essais. Adansonia 13:
125-133.
Bartey, I. W. 1909. The structure of the wood in the Pineae. Bot. Gaz. (Crawfordsville)
48: 47-55.
296 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Bairp, A. M. 1937. The suspensor and embryo of Actinostrobus. J. Roy. Soc. W.
Australia 23: 89-95,
ae The life history of Callitris. Phytomorphology 3: 258-284.
BANNAN M. 1934. Origin and cellular character of xylem rays in gymnosperms.
Bot. poe (Crawfordsville) 96: 260-281.
BARNARD, C. 1926. Preliminary note on branch fall in the Coniferales. Proc. Linn. Soc.
New South Wales 51: 114-128.
Baucn, J., W. Liese, & R. ScHuttze. 1972. The morphological variability of the
ordered pit membranes in gymnosperms. Wood Sci. Technol. 6: 165-184.
BEHNKE, H. D. 1974. Sieve-element plastids of Gymnospermae: their ultrastructure in
relation to systematics. Plant Syst. Evol. 123: 1-12.
BERTRAND, M. C. E. 1879. Sur les teguments séminaux. Des végétaux phanérogames
. 6: 57-92.
BHARADWAJ, D. C. 1963. The organization in pollen grains of some early conifers.
Palaeobotanist 12: 18-2
BouTELigE, J. B. 1955. The wood anatomy of Libocedrus Endl. s. lat., and Fitzroya
. D. Hook. Acta Horti Berg. 17: 177-216.
ee K. 1983. Angiosperms and phylogenetic systematics—some problems and
mples. Abh. Verh. Naturwiss. Vereins Hamburg 26: 343-354.
Bona A. 1849. Tableau des genres des végetaux fossiles considérés sous le point
e de leur classification botanique et de leut géologique. L. Martinet,
— R. 1869. On the ier er distribution of the Coniferae and Gnetaceae.
ns. Roy. Soc. Edinburgh 10: 175-196.
eee G. 1953. Embryogeny ae New Zealand species of the genus Podocarpus
sect. ee Phytomorphology 3: 295-306.
Bucuuotz, J. T. 1918. Suspensor and early embryogeny of Pinus. Bot. Gaz. (Craw-
fordsville) 66: 185-228,
1920. Embryo oo and polyembryony in relation to the phylogeny
of conifers, Amer. J. Bot. 7: 145.
1933. The classification ms Coniferales. Trans. Illinois State Acad. Sci. 25: 112,
. 1939. The ee of Sequoia sempervirens with a comparison of the
sequoias. Amer. J. Bot. 26: 248-257.
. 1941, Sia eae of the Podocarpaceae. Bot. Gaz. (Crawfordsville) 103: 1-37.
& N. Gray. 1948. A taxonomic revision of Podocarpus I. J. Arnold Arbor. 29:
49-76,
BuRLINGAME, L. L. 1915. The origin and relationships of the araucarians. I. Bot. Gaz.
(Crawfordsville) 60: 1-26.
Butts, D., & J. T. BucHHotz. 1940. Cotyledon numbers in conifers. Trans. [linois
State Acad. Sci. 33: 58-62.
canine L. 1890. Die Gymnospermen. Eine morphologisch-phylogenetische Stu-
e. Abh. K6nigl. Bohm. Ges. Wiss. VII. 4: 1-148.
Pa gs 1935. Gymnosperms, structure and function. University of Chicago
Press, Chicago.
CHANEY, R. W. 1951. A revision of fossil Sequoia and Taxodium in western North
America based on the recent discovery of Metasequoia. Trans. Amer. Philos. Soc.
40: 171-2
Cuen, Z. K., & F. H. WANG. 1984. On the systematic position of Amentotaxus from
its ambryological investigation. Acta Phytotax. Sin. 22: 269-276.
CHOWDHURRY, C, R. 1962. The embryogeny of conifers: a review. Phytomorphology
12: 313-338.
Cuu, C.C., & CS. Sun. 1981. Chromosome numbers and morphology in Cathaya.
Acta Phytotax. Sin. 19: 444-446.
1987] HART, CONIFERS 20y,
CLEMENT-WESTERHOF, J. A. 1984. Aspects of Permian palaeobotany and palynology.
IV. The conifer Ortiseia Florin from the Val Gardena Formation of the Dolomites
and the Vicentinian Alps (Italy) with special reference to a revised concept of the
Walchiaceae (Géppert) Schimper. Rev. Palaeobot. Palynol. 41: 51-166.
Compton, R. H. 1922. A systematic account of the plants collected in New Caledonia
and the Isle of Pines by Mr. R. H. Compton, M.A., in 1914. II. Gymnosperms. J.
Linn. Soc., Bot. 45: 421-434.
Core, E. L. 1955. Plant taxonomy. Prentice-Hall, Englewood Cliffs, New Jersey.
CouLTER, J. M. 1909. Evolutionary tendencies among gymnosperms. Bot. Gaz. (Craw-
fordsville) 48: 81-97.
— oe 1917. Morphology of gymnosperms. University of Chi-
cago Press, Chicag
CRANE, P. R. 1985. Pi issenslie analysis of seed plants and the origin of angiosperms.
Ann. Missouri Bot. Gard. 72: 716-793.
& S. R. MANCHESTER. 1982. An extinct juglandaceous fruit from the Upper
Palaeocene of southern England. J. Linn. Soc., Bot. 85: 89-101.
CRANWELL, L. M. 1940. Pollen grains of the New Zealand conifers. New Zealand J.
Sci. eee 22: 1-17
Crisct, J. V., & T. F. STUESSY. 1980. Determining primitive character states for phy-
logenctic ieee Syst. Bot. 5: 112-135.
CRONIN, J. E., & V. M. SaricH. 1980. Tupaiid mas ae ae phylogeny: the macro-
a evidence. Pp. 293-312 in W. Luckett, ed., Comparative biology and
evolutionary relationships of tree shrews. Plenum. Press, New York.
Da.umore, W., A. B. JAcKson, & S. G. HARRISON. 1966. A handbook of Coniferae
and Ginkgoaceae. Edward Arnold, London
Doak OC. G.-1935, eee of foliar types, dwarf shoots and cone scales of Pinus.
Illinois Biol. Monogr. 13: 1-106.
Docra, P. D. 1964. a mechanisms in gymnosperms. Pp. 142-175 in
P. K. K. Nair, ed., Recent advances in palynology. National Botanic Gardens,
Lucknow.
1966. Embryogeny of the Taxodiaceae. Phytomorphology 16: 125-141.
. 1978. Morphology, development and nomenclature of conifer embryo. /bid.
28: 307-322.
Dov te, J. 1945. Developmental lines in pollination mechanisms in the Coniferales.
Sci. Proc. Roy. Dublin Soc. 24: 43-62.
1954. Development in Podocarpus nivalis in relation to other podocarps. III.
General conclusions. /bid. 26: 347-
Aspects and problems of conifer embryology. Advancem. Sci. 54: 1-11.
& S. J. BRENNAN. 1971. Cleavage polyembryony in conifers and taxads—a
survey. I. Podocarps, taxads, and taxodioids. Sci. Proc. Roy. Dublin Soc. 4A: 57-88.
& ———. 1972. Cleavage polyembryony in conifers and taxads—a survey I].
Cupressaceae, Pinaceae, and conclusion. /bid. 137-158
NE. 1943. Pollination in Tsuga pattoniana and in species of Abies and
Picea. Sci. Proc. Roy. Dublin Soc. 23: 57-70.
—— & W. J. Loosy. 1939. Embryogeny in Saxegothaea and its relation to other
podocarps. Sci. Proc. Roy. Dublin Soc. 22: 127-147.
& M. EARY. 1935a. Pollination in Tsuga, Cedrus, Pseudotsuga, and Larix.
Sci. Proc. Roy. Dublin Soc. 21: 191-204.
ee 1935b. Pollination in Saxegothaea. [bid. 181-190.
W. SAXTON. a Contributions to the life history of Fitzroya. Proc. Roy.
Irish Acad. B. 41: 191-21
Doyte, J. A., & M. J. DonoGuue. 1986. Seed plant phylogeny and origin of angio-
sperms: an experimental andistié approach. Bot. Rev. (Lancaster) 52: 321-431.
Dupter, A. W. 1919. Staminate strobilus of Taxus canadensis. Bot. Gaz. (Crawfords-
ville) 68: 345-366.
298 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
—. 1920. Ovuliferous structures of Taxus canadensis. [bid. 69: 492-520.
Eames, A. J. 1913. The morphology of Agathis australis. Ann. Bot. 27: 1-38.
ECKENWALDER, J. E. 1976. Re-evaluation of Cupressaceae and Taxodiaceae: a proposed
erger. Madrono 23: 237-256.
prowen A. W. 1889. Coniferae. Jn: A. ENGLER & K. PRANTL, eds., Nat. Pflanzenfam.
: 28-116.
eae C G. 1950. A further contribution to the life history of Pherosphaera. Proc.
Linn. Soc. New South Wales 75: 320-333.
ERDTMAN, G. 1965. Pollen and spore morphology/plant taxonomy. Gymnospermae,
Bryophyta. Almqvist & Wiksell, Stockholm.
ErRpDTMAN, H. 1963. Some aspects of et ciaane Pp. 88-125 in T. Swain, ed.,
Chemical plant taxonomy. Academic Press, Lon
— &T. Norin. 1966. The chemistry of the nie Cupressales. Fortschr. Chem.
rgan. Naturst. 24: 206-287.
Esau, K. 1969. The phloem. Handbuch der Pflanzenanatomie. Band 5, Teil 2. Born-
traeger, Berlin.
FARRIS, - S. 1985. Distance data revisited. Cladistics 1: 67-85.
. G. Kiuce, & M. F. Mickevicn. 1982. Immunological distance and the
eae relationships of the Rana boylii species group. Syst. Zool. 31: 479-491.
FLorin, R. 1922. On the geological history of the Sciadopitineae. Svensk Bot. Tidskr.
—270.
1940. The Tertiary fossil conifers of south Chile and their phytogeographical
significance, with a review of the fossil conifers of southern lands. Kongl. Svenska
Vetenskapsakad. Handl. II. 19: 1-107.
——-— 48a. On Nothotaxus, a new genus of the Taxaceae from eastern China. Acta
Horti Berg. 14: 385-395.
. 1948b. On - morphology and relationships of the Taxaceae. Bot. Gaz. (Craw-
ie 110: 31-39.
1951. Evolution in cordaites and conifers. Acta Horti Berg. 15: 285-388.
—. 1954. The female reproductive organs of conifers and taxads. Biol. Rev. 29:
367-389.
——. 1955. The systematics of the gymnosperms. Pp. 323-403 in E. L. KEssEL, ed.,
A century of a in the natural sciences, 1853-1953. California Academy of
Sciences, San Fran
58. On ce taxads and conifers from northeastern Europe and eastern
Greenland. Acta Horti Berg. 17: 257-402
. 1963. The distribution of conifer and taxad genera in time and space. /bid. 20:
121-312.
& J.B. Bourevye. 1954. External morphology and epidermal structure of leaves
in the genus Libocedrus, s. lat. Acta Horti Berg. 17: 7-37.
Fitory, W. S. 1936. Chromosome numbers and phylogeny in the gymnosperms. J.
Arnold Arbor. 17: 83-89.
Foster, A. S., & E. M. Girrorp. 1974. Comparative morphology of vascular plants.
W.H. Freeman and Co., San Francisco.
GaAussEN, H. 1944. Les lanes tears actuelles et fossiles. Les Cycadales. Trav. Lab.
Forest. Toulouse II. 1(fasc. 2): 1-104.
1950. Les gymnospermes actuelles et fossiles. Les Coniférales. /bid. Il. 1(fasc.
4): 1-248.
1966. Les gymnospermes actuelles et fossiles. Genres Pseudolarix, Keteleeria,
Larix, Pseudotsuga, Pitiytes, Picea, Cathaya, Tsuga. Ibid. 11. 1(fasc. 8): 481-
Gerry, E. 1916. The distribution of the “‘bars of Sanio” in the Coniferales. Ann. Bot.
2119-124.
GivnisH, T. J. 1980. Ecological constraints on the evolution of breeding systems in
seed plants: dioecy and dispersal in the gymnosperms. Evolution 34: 959-972.
1987] HART, CONIFERS 299
Grecuss, P. 1955. nee of living gymnosperms on the basis of xylotomy.
Akadémiai Kiad6, Bu
. 1972. Xylotomy ae te living connie: Akadémiai Kiad6, Budape
GriFFITH, M.M. 1952. The growth of the shoot apex in ieee Amer.
J. Bot. 39: 253-263
—. 1971. Transfusion tissue in leaves of Cephalotaxus. Phytomorphology 21:
86-92.
Guépés, M., & P. Dupuy. 1974. Morphology of the seed-scale complex in Picea abies
(L.) Karst. ee J. Linn. Soc., Bot. 68: 127-141.
Haines, R. J., & N. PRAKASH. 1980. Proembryo and suspensor elongation in Araucaria
Juss. Austral. J. Bot. 28: 511-522.
Harr, J. B., & E. J. BEUZENBERG. 1958. Chromosomal evolution in the Podocarpaceae.
Nature 181: 1584-1586.
Hate, J. D. 1923. The bars of rims of Sanio. Bot. Gaz. (Crawfordsville) 76: 241-256.
Han, W. 1984. A scanning electron microscope observation of the leaves in some
conifers. Acta Bot. Sin. 26: 376-380.
HARBORNE, J. gia Comparative biochemistry of the flavonoids. Academic Press,
New Yor
Harris, T. +i “1976. The Mesozoic gymnosperms. Rev. Palaeobot. Palynol. 21: 119-
134.
Hart, J. A. 1985. Peripheral isolation a ne origin of diversity in Lepechinia sect.
age Cae Syst. Bot. 10: 6.
Bate B. . The Taxodiaceae ae - divided into several distinct families,
, the Le eens Cryptomeriaceae, Taiwaniaceae and the Cunninghami-
ee and further Tetraclinis should represent a distinct family, the Tetraclinaceae.
Bot. Mag. (Tokyo) 46: 24-27.
Hecut, M. K., & J. EDWARDS. ge The Cena of phylogenetic inference above
the species level. Pp. 3-51 in M. K. Hecut, P. C. Goopy, & B. M. Hecut, eds.,
Major patterns in vertebrate es Plenum, New
HEGNAUER, R. 1962. Chemotaxonomie der Pflanzen. Band I. Thallophyten, Bryophy-
ten, Pteridophyten und Gymnospermen. Birkhauser Verlag, ral and Stuttgart.
Henpy, M. D., & D. Penny. 1982. Branch and bound al
evolutionary trees. Math. Biosci. 59: 277- 290.
HeEnniG, W. 1950. Grundziige einer Theorie der phylogenet Syst tik. Deutsch-
er Zentralverlag, Berlin.
: 66. Phylogenetic systematics. University oe pane Press, Urban
HERZFELD, S. 1914. Die weibliche Koniferenbliite. Oesterr. Bot. Zeitschr. 64: "321 —358.
Hixt, T. G., & E. pE FrRAine. 1906. On the seedling nee of gymnosperms. Ann.
Bot. 20: 471-473.
& 1908. On the seedling structure of gymnosperms. I. [bid. 22: 689-
Ws
—— &
221s
1909a. On the seedling structure of gymnosperms. I]. /bid. 23: 189-
& ———. 1909b. On the seedling structure of gymnosperms. HI. Cycadaceae.
Ibid. 433-458.
Ho, R. H., & O. Szixiar. 1973. Fine structure of the pollen surface of some Taxodiaceae
and Cupressaceae species. Rev. Palaeobot. Palynol. 15: 17-26.
Ho.pen, R. 1913. R ee ae in Coniferales. Bot. Gaz. (Crawfordsville) 55: 56-64.
Hu, Y. S., & F. H. Wana. 1984. Anatomical studies of Cathaya (Pinaceae). Amer. J.
Bot. 1: 12471 35:
. J. Yao. 1981. Transfusion tissue of gymnosperm leaves. J. Linn. Soc.,
qo.
300 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
JAcKMAN, V. H. 1960. The shoot apex of some New Zealand gymnosperms. Phyto-
morphology 10: 145-157
JAIN, K. K. 1975. Evolution of wood structure in Pinaceae. Israel J. Bot. 25: 28-33.
——. 1976. Morphology of the female cone in Pinaceae. Phytomorphology 26: 169-
199.
ia ri E.C. 1905. The pee ae anatomy of the Coniferales. II. The Abietinae.
m. Boston Soc. Nat. Hist. 6: I-
——., "1917. The anatomy of ee plants. University of Chicago Press, Chicago.
. 1926. The comparative anatomy and phylogeny of the Coniferales. Mem,
Boston Soc. Nat. Hist. 5: 441-459.
JOHANSEN, D. A. 1950. Plant embryology of the Spermatophyta. Chronica Botanica
Co., Waltham, Massachusetts.
Jounson, M. A. 1951. The shoot apex in gymnosperms. Phytomorphology 1; 188-
204
KaeIser, M. 1954. Microstructure of wood of Podocarpus. Phytomorphology 4: 39-47.
Kausik, S. B. 1976. A contribution to foliar anatomy of Agathis dammara, with a
discussion on the transfusion tissue and stomatal structure. Phytomorphology 26:
262-273
_S. BHATTACHARYA. 1977. Comparative foliar anatomy of selected gym-
nosperms: leaf structure in relation to leaf form in Coniferales and Taxales. Phy-
tomorphology 27: 146-160.
Kena, H. 1973. On the family Phyllocladaceae. Taiwania 18: 142-145.
1974. The phylloclade of Phy seas and its possible bearing on the branch
systems of progymnosperms. Ann. Bot. n.s. 38: 757-764.
1975. A new scheme of seer sa of . conifers. Taxon 24: 289-292.
KuHosHoo, T. N. 1961. Chromosome numbers in gymnosperms. Silvae Genet. 10: 1-9.
Kiuce, A.G., &J.S. Farris. 1969. Quantitative ee and the evolution ofanurans.
Syst. Zool. 18: 1-32
Kocu, F. 1927. Zur Frage der fossilen und rezenten Verbreitung der Koniferen. Mitt.
Deutsch. Dendrol. Ges. 38: 182-184.
Konar, R. N., & Y. P. OBEROI. ee Recent work on Sag ed structures of living
conifers and taxads—a review. Bot. Rev. (Lancaster) 35: 89-116.
KrassiLov, V. A. 1974. Peon from the Upper anes of eastern Asia and
its bearing on the theory of conifer evolution. Palaeontology 17: 365-370.
LauBENFELS, D. J. bE. 1953. The external morphology of coniferous leaves. Phyto-
morphology 3: 1-20.
1962. The primitiveness of polycotyledony considered with special reference
to the cotyledonary condition in Podocarpaceae. /bid. 12: 296-300.
1965. The relationships of Fitzroya cupressoides (Molina) Johnston and Di-
selina archeri J. D. Hooker based on morphological considerations. /bid. 15: 414—
419
: 1969. A revision of the Malesian and Pacific rainforest conifers. I. Podocar-
paceae. J. Arnold Arbor. 50: 274-369.
Lawson, A. A. 1907. The gametophytes and embryo of the Cupressineae, with special
reference to Libocedrus decurrens. Ann. Bot. 21: 281-302.
1923. The life history of Microcachrys tetragona Hooker. Proc. Linn. Soc. New
South Wales 48: 177-193.
Li, H. L. 1952. The genus Amentotaxus. J. Arnold Arbor. 33: 192-1
1953a. A reclassification of Libocedrus and Cupressaceae. Ibid. 34: 17-36.
——.. 1953b. Present distribution and habitats of the conifers and taxads. Evolution
7: 245-261
Liu, T.S. 1971. A monograph of the genus Abies. Department of Forestry, College of
Agriculture, National Taiwan University, Taipei.
1987] HART, CONIFERS 301
& H.J.Su. 1983. Biosystematic studies on Taiwania and num erl
of the ae of Taxodiaceae. Taiwan Museum, Taipei.
Loosy, W. J., & J. Doyie. 1944. gees and early embryogeny in Podocarpus
anaiiis Sci. Proc. Roy. Dublin Soc. 23: 2 0.
Lorova, L. I. 1975. On the correlation of os sania features of the wood and
phloem in the Pinaceae. Vestn. Moskovsk. Das Ser. 6, Biol. 1: 41-51
Lurzer, E. V. 1956. Megasporenmembranen bei einigen Cupressaceen. Grana Palynol.
1: 70-7
70-78.
are W. P., M. J. DonoGuue, & D. R. Mappison. 1984. Outgroup analysis and
ane Syst. Zool. 33: 83-103.
Ne ., & H. SinGH. 1967. The female gametophyte of gymnosperms. Biol.
0.
Mapes, G., & G. W. RoTHWELL. 1984. Permineralized ovulate cones of Lebachia from
late Palaeozoic limestones of Kansas. Palaeontology 27: 69-94.
MenRA, P. N. 1968. Cytogenetical evolution of conifers. Indian J. Genet. Pl. Breed.
28: 97-111.
T. N. KHosHoo. 1956. Cytology of conifers. I. J. Genet. 54: 165-180.
MEYEN, a V. 1984. Basic features of gymnosperm systematics and phylogeny as evi-
denced by the fossil record. Bot. Rev. (Lancaster) 50: 1-112.
Mitiay, M. A., & D. A. EaGert. 1974. sai athe re ane in the Pa-
leozoic seed fern family Callistrophytaceae. Amer. J. Bot. 61: 1067-
& TayLor. 1974. Morphological adic of Paleozoic ae pollen.
a uoen 147: 75- 79.
& 76 A puma trends in fossil gymnosperm pollen. Rev. Pa-
laeobot. Seren 21: 65-91.
Mitter, C.N. 1976. Early Ses in the Pinaceae. Rev. Palaeobot. Palynol. 21: 101-
jig
1982. Current status of Paleozoic and Mesozoic conifers. /bid. 37: 99-114.
1985. Pityostrobus dase a new species of pinaceous cones from the Late
Cretaceous of New Jersey. Amer. J. Bot. 72: 520-529.
Miter, H. J. 1973. The wood of. pees J. Arnold Arbor. 54: 111-119.
Mortey, T. 1948. On leaf arrangement in Metasequoia re ee Natl.
U.S. A. 34: 574-578.
Mose.ey, M. F. 1943. Contributions to the life history, morphology, and phylogeny
of Widdringtonia cupressoides. Lloydia 6: 109-1
Ntima, O. O. 1968. The araucarias. Fast growing timber trees of the lowland tropics.
No. 3. Commonwealth Forestry Institute, Department of Forestry, University of
Oxford.
Owens, J. N., & M. Motper. 1975. Pollination, female gametophyte, and embryo and
seed development in yellow cedar (Chamaecyparis nootkatensis). Canad. J. Bot. 53:
186-199
: . Sexual reproduction in western red cedar (Thuja plicata).
Canad. J. Forest Res. 7: 605-613.
Pace, C. N.. & H. T. Currrorp. 1981. Ecological biogeography of Australian conifers
and ferns. Pp. 472-498 in A. Keast, ed., Ecological biogeography of Australia. W.
Junk, The Hague, Boston, and London
PATTERSON, C. 1982. Morphological characters and homology. Pp. 21-74 in K. A.
Y & E. A. Fripay, eds., Problems of phylogenetic reconstruction. renen
Press, London.
PATTON, a T. 1927. Anatomy of Australian coniferous timbers. Proc. Roy. Soc. Vic-
toria 40: 2-16.
302 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Peirce, A. S. 1936. Anatomical interrelationships of the Taxodiaceae. Trop. Woods
46: 1-14.
. Systematic anatomy of the woods of the Cupressaceae. Ibid. 49: 5-21.
PentiaLiow, D.P. 1907. A manual of North American gymnosperms. Atheneum Press,
Boston.
PETTITT, 1 ML. 1966. Anew interpretation of the structures of the megaspore membrane
in some gymnospermous ovules. J. Linn. Soc., Bot. 59: 253-263.
19 The megaspore wall in gymnosperms: ultrastructure in some zooido-
gamous forms. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 195: 497-515
Puituips, W. W. J. 1941. The identification of coniferous woods by their microscopic
structure. J. Linn. Soc., Bot. $2: 259-320.
Piccer, R. 1903. Taxaceae. Jn: A. ENGLER, ed., Pflanzenr. IV. 5(Heft 18).
1926. Coniferae. Jn: A. ENGLER & K. PRANTL, eds., Nat. Pflanzenfam. ed. 2.
07.
S. a Pitcat. 1974. Shoot apical organization of some gymnosperms. Phy-
tomorphology 24: 68-74
Poot, D. J. W. 1929. On the anatomy of araucarian wood. Receuil Trav. Bot. Néerl.
482-620.
PraeGer, E. M., D. Fowier, & A. C. Witson. 1976. Rates of evolution in conifers
(Pinaceae). Evolution 30: 637-649.
— & A.C. Witson. 1978. Construction of phylogenetic trees for proteins and
nucleic acids: empirical evaluation of alternative matrix methods. J. Molec. Evol.
129-142.
Quinn, C. J. 1964. Gametophyte development and embryogeny in the Podocarpaceae.
I. Podocarpus sect. Dacrycarpus. Phytomorphology 14: 342-351.
1966. Gametophyte development in Podocarpaceae. IV. Dacrydium co-
lensoi General conclusions. /bid. 16: 199-211.
1970. a boundaries in the Se Proc. Linn. Soc. New South
Wales 94: 166-17
. 1982. ow of Dacrydium Sol. ex Lamb. emend. de Laub. (Podocarpa-
ee Austral. J. Bot. 30: 311-320.
REMANE, A. 1952. Die Grundlagen des natiirlichen ys sicms, der vergleichenden Anat-
omie und der Phylogenetik. Geest und Portig, Leip
Reyre, Y. 1968. La sculpture de l’exine des pollens gymnospermes et des chla-
mydospermes et set utilisation dans identification des pollens fossiles. Pollen &
Spores 10: ce
RopMan, J. E., ee Ou , R. R. NAKAMURA, J. McCLAMMER, JR.,
BLEDSOE. fon A eR analysis and revised er aor ps of Centrospermae.
Syst. Bot. 9: 297-323.
RoTHWELL, G. W. 1982. New interpretation of the earliest conifers. Rev. Palaeobot.
SAHNI, B. 1920. On certain archaic features in the seed of Taxus baccata with remarks
on the antiquity of the Taxineae. Amer. J. Bot. 34: 118-133.
SATTLER, R. 1984. Homology: a eee challenge. Syst. Bot. 9: 382-394.
SAx, K., & H. J. Sax. 1933. Chromosome number and morphology in the conifers. J.
Amold Arbor. 14: 356-374
Saxton, W. T. 1913. The classification of conifers. New Phytol. 12: 242-262.
1934. Not ifers VII. The morphology of Austrotaxus spicata Compton.
Ann. Bot. 48: 411-427.
SCHLARBAUM, S. E., & T. TsucutyA. 1985. Karyological derivation of Sciadopitys
verticillata Sieb. et Zucc. from a pro-taxodiaceous ancestor. Bot. Gaz. (Crawfords-
ville) 146: 264-267
1987] HART, CONIFERS 303
Sewarp, A. C. 1919. Fossil plants. Vol. 4. Cambridge University Press, Cambridge,
England.
Suaw, G. R. 1914. The genus Pinus. Arnold Arbor. Publ. No. 5. Houghton Mifflin
Co., Boston.
SIBLEY, Cc G., & J. E. Axtourst. 1984. The phylogeny of hominoid primates, as
indicated by DNA-DNA hybridization. J. Molec. Evol. 20: 2-15
Sicpa, J. 1984. An international census of the Coniferae, I. Phytologia Mem. 7: 1-79
Sincu, H. 1961. The life-history and systematic position of Cephalotaxus drupacea
Sieb. & Zucc. Phytomorphology 11: 153-197.
78. Embryology of gymnosperms. Briider Borntraeger, Berlin
& J. CHATTERJEE. 1963. A contribution to the life history of Cryptomeria
japonica D. Don. Phytomorphology 13: 428-445.
Sinnott, W. W. 1913. The morphology of the reproductive structures in the Podo-
carpineae. Ann. Bot. 27: 39-82
Sporne, K. R. 1965. The morphology of gymnosperms. Hutchinson, London
Steppins, G.L. 1948. The chromosomes and relationships of Metasequoia and Sequoia.
STERLING, C. 1963. Structure of the male gametophyte in gymnosperms. Biol. Rev. 38:
167-203.
Stevens, P. F. 1980. Evolutionary polarity of character states. Ann. Rev. Ecol. Syst.
11: 333-358.
1984. Homology and phylogeny: morphology and systematics. Syst. Bot. 9:
395-409.
_ 1986. Evolutionary classifications in botany, 1960-1985. J. Arnold Arbor. 67:
313-339
STEWART, W. N. 1983. Paleobotany and the evolution of plants. Cambridge University
Stipp, B. M., & K. Cosentino. 1976. Nucellangium: gametophytic structure and re-
lationship to Cordaites. Bot. Gaz. (Crawfordsville) 137: 242-249,
STRASBURGER, E. ne Die Coniferen und die Gnetaceen. Eine morphologische Studie.
Fischer, Jen
1878. Tiber Befruchtung und Zelltheilung. H. Dabis
1879. Die Angiospermen und die Gymnospermen. G.F a , Jena.
a, M. 1953. Studies on the germination of the pollen Bae in conifers. Jap.
14: 13-21.
eda A. L. 1953. Phylogenetic principles of the system of higher plants. Bot.
Rev. (Lancaster) 19: 1-45.
Tayior, T. N. 1981. Paleobotany, an introduction to fossil plant biology. McGraw-
Hill, New York.
W.N. Stewart. 1964. The Paleozoic seed Mitrospermum in American coal
balls. Palacontogaphicn a, B. 115: 51-58.
TEGNER, J. 1965. acrydium—anatomy and taxonomy. Bot. Not. 118: 450-452.
: 67. merce and taxonomy in the Podocarpaceae. [bid. 120: 504-506.
Tuomson, R. B. 1905. The megaspore-membrane of the gymnosperms. Studies, Univ.
Toronto, Biol. Ser. 4: 1-64.
14. The spur Shoot of the pines. Bot. Gaz. (Crawfordsville) 17: 362-386.
1940. The structure of the cone in the Coniferae. Bot. Rev. (Lancaster) 6:
73-84.
& H. B. Sirron. 1926. Resin canals in the Canadian spruce (Picea canadensis
(Mill.) B.S.P.)—an anatomical study especially in relation to traumatic effects an
their bearing on phylogeny. Philos. Trans., Ser. B. 214: 63-111.
panereye P. van. 1869. Anatomie compar€ée dela fleur femelle et du fruit des Cycadées,
es Coniféres et des Gnétacées. Ann. Sci. Nat. Bot. V. 10: 269-304
RR aes P. B. 1984. Homology: an empirical view. Syst. Bot. 9: 374-381.
304 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
TURRILL, W. B. 1959. ene Pp. 494-518 in Vistas in botany. Vol. 1. Per-
amon Press, Lon
UENo, J. 1960. Studies o on pollen grains of Gymnospermae, concluding remarks to the
relationships between Coniferae. J. Inst. Polytechn. Osaka City Univ., Ser. D., Biol.
11: 109-136.
VasiL, V., & R. K. Sanni. 1964. Morphology and embryology of Taxodium mucro-
natum Tenore. Phytomorphology 14: 369-384.
VELENOVSKY, J. 1905. Vergleichende Morphologie der Pflanzen. Fr. Rivnae, Prague.
Wana, F. H., Z. K. CHen, & Y. S. Hu. 1979. On the systematic position of Taxaceae
from the embryological and anatomical studies. Acta Phytotax. Sin. 17(3): 1-7.
, S.C. Lee, & Z. K. Cen. 1980. The embryogeny of Taiwania in comparison
with that of other ea of Taxodiaceae. Acta Phytotax. Sin. 18: 129-137.
Watrous, L. E., & Q. D. WHEELER. 1981. The outgroup comparison method of char-
acter analysis. Syst. Zool. 30: I-11
WiLpE, M. 1975. A new iiterpretaiion of the microsporangiate cones in Cephalotax-
aceae and Taxaceae. Phytomorphology 25: 434-450,
WILEY, . O. 1981. Phylogenetics: the theory and practice of phylogenetic systematics.
Wiley and Sons, New York.
Wen R. P. 1935. Pollen grains; their structure, capa eae and significance
in science and medicine. McGraw-Hill, New York a don
Yao, B. J., & ¥.S. Hu. 1982. Comparative anatomy of ea leaves. Acta Phytotax.
Zikiicenan. Ww. 1930. Die Phylogenie der Pflanzen. G. Fischer, Jena.
APPENDIX. Character states used in the phylogenetic
analysis of coniferous taxa.
BRANCHING AND GROWTH PATTERNS. |, Higher-order branches spiral / opposite (Morley,
1948; Dallimore et a/., 1966). 2, Short shoots absent f en (Barnard, 1926: Doak,
1935; Morley, 1948; Stebbins, 1948: Dallimore et a/., 1966). 3, Branches not annually
deciduous / annually deciduous (Morley, 1948; ne 1948: Eckenwalder, 1976).
ANATOMY. 4, Sieve-element plastids starch accumulating / protein accum ulating (Behnke,
1974)
STEM ANATOMY. 5, Stem tip without / with tunica corpus (Johnson, 1951: Griffith, 1952:
Jackman, 1960; Pillai, 1963; Sporne, 1965; Pillai & Pillai, 1974). 6, Phloem fibers present/
absent (Esau, 1969)
Woop ANATOMY. 7, Phloem-fiber sclereids absent / present (Lotova, 1975). 8, Phloem
mucilage absent / present bea 1975). 9, Xylem parenchyma absent / present (Bailey,
1909; Phillips, 1941; Gre , 1955; Sporne, 1965; Te — 1965, 1967; H. J. Miller,
1973; Chu & Sun, 1981). 0. “End or transverse walls of wood parenchyma (as seen in
tangential section) smooth / nodular or pitted (Peirce, cn 1937; Phillips, 1941: Bou-
telje, 1955). 11, Horizontal walls of wood parenchyma (as seen in radial section) smooth /
nodular or pitted (Greguss, 1955). 12, Bordered pits of tracheids alternate, multiseriate,
hexagonal in outline / uniseriate (Phillips, 1941; Florin, 1951; Gr reguss, 1955; Sporne,
1965; Stewart, 1983). 13, Spiral thickenings on longitudinal tracheid walls (early wood)
absent / present (Compton, 1922; Phillips, 1941; Greguss, 1955, 1972: Stewart, 1983).
14, Spiral thickenings on transverse tracheid walls absent / present (Greguss, 1972; Hu
& Wang, 1984). 15, Bordered pits with / without torus (Bauch e¢ al., 1972). 16, Crassulae
*The descriptor to the left of the slash (/) indicates the primitive condition, the one to the right
the derived condition. oo characters a slash (/) is used for ordered characters. a vertical
line (|) for unordered o
1987] HART, CONIFERS 305
(bars of Sanio) present / absent (Jeffrey, 1905; Gerry, 1916; Hale, 1923; Chamberlain,
1935: Phillips, 1941). 17, Resin ducts in secondary wood absent / present (Jeffrey, 1905;
Chamberlain, 1935; Jain, 1975; Taylor, 1981; Hu & Wang, 1984). 18, Traumatic resin
ducts absent / present (Bailey, 1909; Phillips, 1941). 19, Resin ducts in rays present /
absent (Patton, 1927; Phillips, 1941; Hu & Wang, 1984). 20, Horizontal walls of wood
rays smooth / thickened, nodular or with simple pits (Bannan, 1934; Phillips, 1941;
Boutelje, 1955; Greguss, 1955). 21, Tangential walls of wood rays smooth / thickened,
nodular (Greguss, 1955). 22, Indentations on horizontal walls of ray parenchyma absent /
present (Phillips, 1941; Kaeiser, oT SS 1955). 23, Ray tracheids absent / present
(Holden, 1913; Phillips, 1941). 24, Ray tracheids smooth walled / dentate (Phillips,
1941). 25, Cross-field pits cupressoid or taxoid (round) / piciform ae slits) (Phillips,
1941). 26, Tracheids not resinous / resinous (Patton, 1927; Pool, 192
Leaves. 27, Leaves large / small. 28, Leaves falcate in profile and tetragonal in cross
section / (1) linear or lanceolate and bifacially flattened | (2) eee | (3) bilaterally
flattened | (4) needlelike | (5) double (fused?) (De Laubenfels, 1953). 29, Leaves single,
spread out on branch / (1) in fascicles, spirally arranged on short ee | (2) helically
arranged on short shoots (Thomson, 1914). 30, Leaf phyllotaxy spiral / (1) spiral op-
posite (bijugate) | (2) decussate | (3) ternate (3-whorled) (De Laubenfels, 1953). 31,
Seedling phyllotaxy whorled / opposite (De Laubenfels, 1953, 1965), 32, Leaf attachment
decurrent / (1) with stalklike a | (2) with shield-shaped attachment (De Lau-
benfels, 1953; Liu, 1971). 33, Mature foliage leaves monomorphic / dimorphic (facial
and lateral leaves) (De uae 1953). 34, Lateral margins Paes, leaves (in flattened
branches with dimorphic leaves) free / fused. 35, Leaf bases distinctly decurrent / fused
(De Laubenfels, 1953). 36, Leaves persistent / annually deciduous (Dallimore et al.,
1966). 37, Apical meristems without modified leaves / (1) shorter leaves interrupting
epistomatic (Florin, 1951; Florin & Boutelje, 1954). 39, Leaves with endodermis (vas-
cular sheath) not having / having thickened Casparian strips (Yao & Hu, 1982). 40,
Mesophyll parenchyma smooth / plicate (Kausik & Bhattacharya, 1977; Yao & Hu,
1982: Han, 1984). 41, Tracheids of leaf transfusion tissue lateral to the vascular bundle /
all around vascular punele (mostly on abaxial side) (Griffith, 1971; Kausik, 1976; Kausik
& Bhattacharya, 1977; Hu & Yao, 1981). 42, Vascular bundles of leaf 1 / (1) 2 /
(2) more than 2 (Chamberlain. 1935: Kausik & Bhattacharya, 1977; Stewart, 1983).
CHEMISTRY. 43, Biflavonoids present / absent (Hegnauer, 1962; Harborne, nee 44,
Nootkatin absent / present (H. Erdtman, 1963; H. Erdtman & Norin, 1966). 45, Hi-
nokinflavone absent / present (H. Erdtman, 1963; Harborne, 1967). 46, ae
absent / present (H. Erdtman, 1963; H. Erdtman & Norin, 1966). 47, Leaf wax estolid /
nonestolid (Hegnauer, 1962).
SEX DISTRIBUTION. 48, Plants monoecious / dioecious (Chamberlain, 1935; Florin, 1948b;
Li, 1952; Greguss, 1955: Singh, 1961; Dallimore er a/., 1966; Ntima, 1968; Givnish,
1980).
MICROSPORANGIATE STROBILUS. 49, Microsporangiate strobili compound / simple (Stew-
art, 1983). 50, Microsporangiate strobili terminal / axillary. 51, Microsporangiate strobili
single at ends of leafy shoots / (1) grouped in clusters | (2) grouped in racemes or panicles.
52. Microsporophylls spiral / decussate (whorled). 53, Microsporophylls open (laminar),
ce ee perisporangiate (Thomson, 1905; Dupler, 1919; Chamberlain,
1935: Ueno, 1960; Wilde, 1975). 54, Microsporangia 2 / more than 2 (Saxton, 1934;
SCENE 1935: Florin, 1951; Ueno, 1960). 55, Nierosporaudial dehiscence longi-
tudinal / (1) oblique / (2) transverse (Liu, 1971).
ICROGAMETOPHYTE. 56, Prepollen / pollen (Mapes & Rothwell, 1984). 57, Pollen-tetrad
formation simultaneous (tetrahedral) / successive (bilateral) (Ueno, 1960). 58, Pollen
306 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
with shallow functional germination furrow / (1) with harmomegathus | (2) with func-
tionless germ furrow | (3) with pore (Wodehouse, 1935; Ueno, 1960; G. Erdtman, 1965).
59, Pollen without / with papilla germination (Elliot, 1950; Takeuchi, 1953; Ueno, 1960;
Ho & Sziklai, 1973). 60, Pollen grains with comfit perine absent / present (Ueno, 1960).
61, Pollen sexine tegillate / (1) rough corrugate | (2) granular | (3) roughened (Wodehouse,
1935; Ueno, 1960). 62, Pollen-sexine ultrastructure simple or absent / (1) compound /
(2) double / (3) roughened (Wodehouse, 1935; Ueno, 1960; Reyre, 1968). 63, Pollen
without / with annular thickenings (Ueno, 1960). 64, Pollen without / with triradiate
streaks (Ueno, 1960). 65, Pollen winged (monosaccate: bilateral or bisaccate) / (1) wing-
_—
1948; Florin, 1951; Ueno, 1960; Bharadwaj, 1963; Sporne, 1965; Millay & Taylor, 1974;
Singh, 1978). 66, Pollen intine thin / thick (Ueno, 1960; Singh, 1961; Liu & Su, 1983).
67, Pollen multi- or binucleate / uninucleate at pollination (Singh & Chatterjee, 1963;
Vasil & Sahni, 1964). 68, Pollen grains containing | or 2 / (1) 0 | (2) many prothallial
cells (Chamberlain, 1935; Wodehouse, 1935; Elliot, 1950; Ueno, 1960; Sterling, 1963;
Konar & Oberoi, 1969; Millay & Eggert, 1974; Singh, 1978). 69, Sperm nuclei with /
without cell walls (Chamberlain, 1935; Singh, 1978). 70, Sperm cells unequal / equal
(Burlingame, 1915; Ueno, 1960; Sterling, 1963; Owens & Molder, 1975; Wang, Chen,
& Hu, 1979).
MEGAGAMETOPHYTE AND EMBRYO. 71, Pollination drop present / absent (J. Doyle, 1945:
Dogra, 1964; Singh, 1978). 72, Pollen germination on nucellus / on scales (Dogra, 1964:
Singh, 1978). 73, Micropyle symmetrical / asymmetric (J. Doyle & O’Leary, 1935a,
1935b; J. Doyle & Kane, 1943; Looby & Doyle, 1944; J. Doyle, 1945; Dogra, 1964:
Singh, 1978). 74, Ventral-canal cell with distinct cell wall / with no wall, but having
nuclei (Lawson, 1907; Chamberlain, 1935; Owens & Molder, 1975). 75, Alveoli open
on area adjacent to central vacuole / closed by cell walls (Lawson, 1923). 76, Megaga-
metophyte without / with layer of peripheral cells (Saxton, 1913: Maheshwari & Singh,
1967; Singh, 1978). 77, Megaspore membrane thick, double / thin (Thomson, 1905:
Lawson, 1907; Quinn, 1966; Owens & Molder, 1975; Stidd & Cosentino, 1976; Singh,
1978). 78, Megaspore membrane of uniform thickness / thin at micropylar end (Thom-
son, 1905). 79, Megaspore membrane suberized / not suberized (Thomson, 1905). 80,
Tapetum primary / secondary (Thomson, 1905; Saxton, 1913; Singh, 1978). 81, Arche-
gonia not surrounded / surrounded by densely cytoplasmic tissue (Singh, 1978). 82,
Archegonia separate / grouped together to form complexes (Lawson, 1907: Chamberlain,
1935; Maheshwari & Singh, 1967; Owens & Molder, 1975, 1980; Singh, 1978; Wang,
Lee, & Chen, 1980). 83, Archegonia separated by vegetative cells / arranged in ring
(Eames, 1913; Eckenwalder, 1976). 84, Archegonia apical (at micropylar end) / (1) lateral
(at middle ee | (2) lateral (at chalazal end of gametophyte) (Saxton, 1913:
Moseley, 1943; Florin, 1951; Maheshwari & Singh, 1967; Konar & Oberoi, 1969: Foster
& Gifford, 1974: Singh, 1978). 85, Archegonial jacket Sey he ea (Singh, 1978). 86,
Proembryo with free nuclear divisions many / (1) 5 or 4 / (2) 3 / (3) 2/ (4) 0 (Eames,
1913; J. Doyle & Saxton, 1933; J. Doyle, 1954; Chowdhurry, 1962: Sporne, 1965: Chen
& Wang, 1984). 87, Proembryo with secondary / primary type of wall formation (Dogra,
1966). 88, Proembryo nontiered / (1) with upper, suspensor, and embryonal tiers / (2)
nontiered (reduced) (Moseley, 1943; Chowdhurry, 1962; Foster & Gifford, 1974: Dogra,
1978; Haines & Prakash, 1980). 89, Proembryo 3- / 4-tiered (Dogra, 1978; Singh, 1978).
90, Proembryo with embryonal cells uninucleate / binucleate (Saxton, 1913; J. Doyle &
Looby, 1939; Buchholz, 1941; Elliot, 1950; Brownlie, 1953; J. Doyle, 1954; Chowdhurry,
1962; Quinn, 1964, 1966, 1970). 91, Proembryo basal / central (Haines & Prakash,
1980). 92, Proembryo with irregular shape / with spherical shape of free nuclear embryo
nd curved planes of upper, SuRPEDs Or and embryonal tiers of cellular phase (Haines &
Prakash, 1980). 93, Proembryo with development of primary suspensor from suspensor /
from upper tier (Dogra, 1978). 94, Suspensor anchorage of proembryo not within / within
archegonium (Haines & Prakash, 1980). 95, Prosuspensor present / absent (Baird, 1937,
1953; Johansen, 1950). 96, Proembryo not completely filling / completely filling arche-
1987] HART, CONIFERS 307
onium (Moseley, 1943). 97, Polyembryony simple / cleavage (J. Doyle, 1957; J. Doyle
& Brennan, 1971, 1972; Singh, 1978).
OVULATE STROBILUS. 98, Cone terminal on leafy branches / axillary on short, leafy shoots
Ovulate strobilus compound / simple (Dupler, 1920; Li, 1952; Sporne, 1965). 100,
symmetr ical / (1) bilaterally flattened / (2) “scales”
(Taylor, 1981; Mapes & Rothwell, 1984; Meyen, 1984). 101, Bract- scale complex free /
fused (Sporne, 1965). 102, Cone bract not keeled / keeled (C. N. Miller, 1985). 103,
Cone scales flat / peltate (Chamberlain, 1935; Li, 1953a; Sporne, 1965; Foster & Gifford,
1974). 104, Cone scales imbricate, thin / valvate, thickened (Li, 1953a). 105, Cone scales
woody / modified into an epimatium (Sinnott, 1913). 106, Epimatium fully covering
seeds / (1) half covering seeds / (2) lacking (Sinnott, 1913, Herzfeld, 1914; Aase, 1915;
Chamberlain, 1935; Florin, 1951, 1958). 107, Epimatium not fused / fused to seed coat
(Quinn, 1982). 108, Bracts not fleshy / fleshy (De Laubenfels, 1969; Quinn, 1982). 109,
Bracts free / fused (De Laubenfels, 1969; Quinn, 1982). 110, Receptacle not warty /
warty (De Laubenfels, 1969). 111, Cone scales persistent / deciduous (Chamberlain,
1935; Liu, 1971). 112, Cones pendulous / upright at maturity (Liu, 1971). 113, Uniaxial
seeds arranged singly on primary shoots of unlimited / limited growth (Florin, 1948a,
1948b, 1954)
OVULES AND SEEDS. 114, Ovules inverted / (1) semi-erect / (2) erect (Stebbins, Be
Stewart, 1983; Clement- Westerhof, 1984; Mapes & Rothwell, 1984; Miller, 1985). 1
Number of ovules per cone scale: | / 2 or more (Clement-Westerhof, 1984). 116, oe
storage product: starch / oils (Hegnauer, 1962). 117, Seed without / with aril (Florin,
1951, 1958; Sporne, 1965; Foster & Gifford, 1974; Quinn, 1982). 118, Aril not developed
by intercalary growth, not fused to seed / partly developed by intercalary growth, fused
to seed coat (Florin, 1948a, 1948b). 119, Seeds winged / not winged (Taylor & Stewart,
1964; De Laubenfels, 1965; Dallimore et a/., 1966; Singh, 1978; Rothwell, 1982). 120,
Resin ducts in seed coat absent / present (Price, pers. comm.). 121, Number of coty-
ledons: 2 / more than 2 (Hill & De Fraine, 1906, 1908, 1909a, 1909b: Buchholz, 1920;
Butts & Buchholz, 1940; De Laubenfels, 1962). 122, Seeds maturing in 2 / 1 year(s)
(Singh, 1978).
CytoLocy. 123, Chromosome number: 12 / (1) 10 | (2) 11 (Sax & Sax, 1933; Flory,
1936; Mehra & Khoshoo, 1956).
GRETHER, MIMOSA 309
TAXONOMIC AND NOMENCLATURAL NOTES ON THE
GENUS MIMOSA (LEGUMINOSAE)
ROSAURA GRETHER!
These notes result from studies concerning the revision of Mimosa species
occurring in the state of Oaxaca, Mexico. They comprise revised synonymies,
typifications, a new combination, and a new name and are based on study of
type collections and field observations.
The following taxonomic and nomenclatural notes are based on a study of
those species of Mimosa L. occurring in the state of Oaxaca, Mexico. Exami-
nation of type specimens and of additional material from Oaxaca, other parts
of Mexico, and Central and South America, as well as fa observations in
Mexico, supports the synonymies and changes proposed her
This paper formalizes and validates synonymies, i eRe Henan a new
combination, and a new name before publication of ‘‘Leguminosas de Oaxaca,”
now in preparation, which will include keys, descriptions, and geographic dis-
tributions for the genus.
The following species, in alphabetical order, are known to occur in Oaxaca.
Mimosa acantholoba (Humb. & Bonpl. ex Willd.) Poiret in Lam. Encycl. Méth.
Bot. Suppl. 1: 83. 1810
Acacia acantholoba Humb. & Bonpl. ex Willd. Sp. Pl. 4: 1089. 1806. Type: America
eae Humboldt & Bonpland 3800 (holotype, B-Willd., IDC 7440. 1391: IL.
3!; iso
Mimosa ee Robinson, Proc. ne Acad. Arts 36: 472. 1901. Neomimosa
eurycarpoides (Robinson) Britton & R N. Amer. FI. 23: 172. 1928. Type: Mexico,
Sinaloa, near Colomas, 21 July 1897, Ms 1805 (holotype, us! (fragments, GH!;
photo and fragments, Ny!)).
Mimosa colimensis Robinson, Proc. ea Soc. Nat. Hist. 258. 1904. Neomimosa
dla ae Britton & Rose, N. Amer. Fl. 23: 172. 1928. Type: Mexico,
r Colima, Aug. 1897, aan 128 (holotype, (fragments, Ny!); iso-
type eae
ene russellii Britton & Rose, N. Amer. Fl. 23: 173. 1928. Type: Mexico
Sinaloa, vicinity of Rosario, 14 April 1910, Rose, Standley, & P.G. Russell 14555
(holotype, us! (photo, MEXxU!)).
The original description of Mimosa eurycarpoides was based on a flowering
specimen (with an associated unattached fruit, probably of Acacia farnesiana,
as indicated by Robinson (1904)). Mimosa colimensis was also based on flow-
ering material; Neomimosa russellii, on a fruiting specimen.
‘Departamento de Biologia, Division de C.B.S., U niversidad Autonoma Metropolitana—Iztapalapa,
Apdo. Postal 55-535, 09340 México, D.F., Me
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 309-322. July, 1987.
310 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Because all the type specimens of the synonyms are incomplete, I have
collected material with flowers and fruits in type localities or nearby (Sinaloa,
0.5 km NW de Rosario, R. Grether 810, MEXU, UAMIZ; Colima, 8.5 km SE de
Colima, R. Grether 875, MEXU, UAMIz) and have examined many specimens
from the states of Sinaloa, Nayarit, ene Colima, Michoacan, Guerrero, and
Oaxaca in the field and/or the herbari
Concerning the inflorescence, Robinson (1904) remarked that Mimosa co-
limensis differs somewhat from M. eurycarpoides in the oval form of the young
heads; however, examination of the type specimens shows that both of them
have subglobose young heads, although they look globose to almost globose
when mature. Flower characteristics of both types are also the same: calyx
campanulate, glabrous, one third to one half of corolla length; corolla five-
lobed, glabrous, 2—2.5 mm long; stamens ten.
Although I could not gather flowering specimens of Neomimosa russellii in
the type locality, remnants of flowers show the corolla to be five-lobed and
glabrous, and fruits of the population growing there clearly correspond to
Mimosa acantholoba.
Form, pubescence, and size of stipules and leaflets are essentially the same
in type specimens of Mimosa eurycarpoides, M. colimensis, and Neomimosa
russellii, as are the number of pinnae and leaflets.
My analysis of the original description and the microfiche of the type spec-
imen of Acacia acantholoba, as well as my examination of several specimens
from Nicaragua, Ecuador, and Peru, leads me to the conclusion that all these
names have been used for a single widely distributed American species.
The fruits of Mimosa acantholoba vary in form and in the density of bristles
(Grether, 1984): the valves can be elliptic to oblong and completely glabrous
to setose, even in a single population.
Mimosa adenantheroides (Martens & Galeotti) Bentham, London J. Bot. 5:
88. 1846
Acacia adenantheroides Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 10(2):
312. 1843. Type: Mexico, Oaxaca, mountains of Sola de Vega and Yolotepec, S of
Oaxaca, 1840, Galeotti 3208 (holotype, BR (fide Rudd, 1984); isotype, K! (photos,
MEXUI, us!
ee cylindrflora Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 10(2): 313.
43. Type: Mexico, Oaxaca, Don Dominguillo, 1840, Galeotti 3207 (holotype, BR
ae MEXU!, Us!)).
Mimosa remota Bentham, London J. Bot. 5: 88. ae Type: Mexico, Oaxaca, Cor-
dillera, 1840, Galeotti 3240 (holotype, BR; isotype, K
Mimosa gomezii Britton & Rose, N. Amer. FI. 23: 159. aoe Type: Mexico, Oaxaca,
valley of Oaxaca, 20 Sept. 1894, Nelson 1479 (holotype, us! (fragments, K!, photo
and fragments, Ny!); isotype, GH!).
The type specimens of Acacia adenantheroides, A. cylindriflora, and Mimosa
remota are in flower, while that of M4. gomezii has both fruits and flowers
(although the spikes are very short in the latter).
I consider Mimosa adenantheroides to be a single variable species because
all the types were collected in the state of Oaxaca and examination of numerous
1987] GRETHER, MIMOSA 311
specimens from Oaxaca, including material collected near the type locality of
Acacia adenantheroides (Distrito Sola de Vega, La Cumbre, 18 km SW de Sola
de Vega, M. Sousa et al. 10509, MExu!, UAMiz!; Distrito de Juquila, 22 km E
de Juquila, 4 km W de Yolotepec, 7. Sousa et al. 10545, MExU!, UAMiIz!), and
from the states of Jalisco, Michoacan, México, Puebla, Morelos, Guerrero, and
Chiapas, indicates much variation in populations with respect to size and
number of pinnae and leaflets, length of spikes, and number of corolla lobes
(four or five) and stamens (eight to ten), as well as to size of the legume and
density of glandular dots and prickles at its margin.
The genus has been widely collected in Oaxaca, and no other closely related
species that could be confused with Mimosa adenantheroides has been found.
Mimosa camporum Bentham, J. Bot. (Hooker) 2: 130. 1840. Type: British
Guiana, June 1839, Schomburgk 725 (holotype, BR; isotypes, F!, G, K,
M, NY!, us!, W).
Mimosa flavescens Splitg. Tijdschr. Natuurl. Gesch. Physiol. 9: 110. 1842. Type: Sur-
inam, Splitgerber s.n. (isotypes, K (photo, A!), w
Mimosa aeschynomenes Bentham, Bot. Voy. Sulphur, 89. 1844. Type: [Nicaragua,]
Realejo, 1841, Hinds s.n. (holotype, BM; isotype, K!).
Mimosa pusilla Bentham, Bot. Voy. sees 90. 1844. Type: [Nicaragua,] Realejo,
1842, Hinds s.n. (holotype, BM; isot K!).
Mimosa flaviseta Bentham, London J. Bot ‘5: 90. 1846. Type: Surinam, 1843, Hostman
813 (holotype, BM; isotypes, GH!, K (photo, A!), NY!).
Mimosa martensis Britton & Rose in Britton & Killip, Ann. New York Acad. Sci. 35:
152. 1936. Type: Colombia, Santa Marta, 1898-1899, H. H. Smith 714 (holotype,
NyY!; isotype, us!).
Bentham (1875) considered Mimosa flaviseta, M. aeschynomenes, and M.
flavescens as synonyms of M. camporum and mentioned (p. 436) M. pusilla as
‘possibly a small slender variety of 4. camporum.”
Robinson’s (1898) description of Mimosa camporum was based on two
specimens from Mexico (Rose 3116 (us!), from Acaponeta [Nayarit], and 3295
(F!, K!, us!), from Tepic [Nayarit], however this corresponds to M. occidentalis
Britton & Rose, mainly in the large oval heads 2.5 cm in diameter. In fact,
Britton and Rose selected Rose 3295 as the type of M. occidentalis, described
in N. Amer. FI. 23: 162. 1928.
Iam here placing Mimosa pusilla and M. martensis in the synonymy of M.
camporum, because stipule, leaflet, bracteole, flower, and fruit characters are
those of /. camporum. Even though size and density of pubescence have been
indicated as differences between M. pusilla, M. martensis, and M. camporum,
examination of type specimens and other material from Mexico (states of
Guerrero, Oaxaca, Veracruz, Tabasco, and Chiapas), Nicaragua (near Realejo,
QDersted 4323, F!), Costa Rica, and Venezuela shows variation in size and density
of hispidity, even in specimens from the same locality.
Mimosa ervendbergii A. Gray, Proc. Amer. Acad. Arts 5: 178. 1862. TyPE
Mexico, Veracruz, Prov. Huasteca, near Tantoyuca, 1858, ect ee
2, p.p. (holotype, GH!; isotypes, K! (photo, MEXU!), Us!).
je JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Mimosa costaricensis Bentham, Trans. Linn. Soc. London 30: 423. 1875. Type: Costa
Rica, Aguacate, Bersted 15 (lectotype, here designated, k! (photo and fragments,
us!)).
Mimosa mexiquitensis Britton, N. Amer. Fl. 23: 168. 1928. Type: Mexico, Chiapas,
exiquito, Sept. 1913, Purpus 6816 (holotype, Ny!; isotypes, GH!, Mo!, Us!).
Neomimosa donnell- smithii Britton & Rose, N. Amer. FI. 23: 173. 1928. Mimosa
donnell-smithi (Britton & Rose) Standley & Steyerm. Publ. Field Mus. Nat. Hist.,
Bot. Ser. 23: 163. 1944. Type: Guatemala, Departamento Alta Verapaz, Cubilquitz,
1902, Von Tuerckheim 8197 (holotype, us!).
Mimosa scalpens Standley, Publ. Carnegie Inst. Wash. 461: 58. 1935. Type: British
oe vicinity of Jacinto Hills, 4 May 1934, Schipp 1306 (holotype, F!; isotypes,
GH!, MO!, NY!),
Mimosa ervendbergii was based on a mixed collection of flowering material,
as indicated by Robinson (1898): the specimen on the left correponds to this
species, and the one on the right to A/. invisa Martius. Robinson considered
M. costaricensis to be a synonym of M. ervendbergii, and I have confirmed the
correctness of that decision by examining the type specimens. Gersted 15 is
here selected as the lectotype of M/. costaricensis.
Mimosa mexiquitensis also corresponds to the same species; examination of
flowering and fruiting material from Chiapas, in addition to the type specimen,
leads me to this conclusion
The type of Neomimosa donnell-smithii, a fruiting specimen, has remnants
of flowers that clearly match the same structures in Mimosa ervendbergii (calyx
long ciliate, one third to one half of corolla length; corolla glabrous, four-lobed
stamens eight). It 1s interesting to note that Standley and Steyermark transferred
Neomimosa donnell-smithti as Mimosa donnell-smithii in 1944; the same au-
thors included that species in the Flora of Guatemala (1946), although pointing
out (p. 56) that “‘we have seen no representation of this species.”’ In the same
publication they considered WV. sca/pens from Belize, described by Standley in
1935, to be a different species occurring in Guatemala, even though the two
are, in fact, the same taxon.
The original description of Mimosa scalpens indicates pentamerous flowers,
and that of M. ervendbergii tetramerous ones; however, variation in the number
of corolla lobes (four or five) and stamens (eight to ten) has commonly been
observed in the species. Although corolla-lobe number is a good character for
many species of \/imosa, it varies (four or five) in several species of the genus.
Considering the characters that distinguish Mimosa ervendbergii (calyx lobes
long-ciliate, very conspicuous in bud: corolla four- or five-lobed, glabrous;
stamens eight to ten; legume articulated, stipitate, glabrous, apex rostrate, mar-
gins prickly; twigs angled, densely tomentose; stipules filiform, tomentose;
leaflets puberulous above, tomentose below, with a prominent excentric nerve)
and having seen all of them in the type specimens and in additional herbarium
material from Mexico (states of Veracruz, Puebla, Oaxaca, Tabasco, and Chia-
pas), Guatemala, Nicaragua, and Costa Rica (Monte Aguacate, | 1/47, Bersted
4463, F!, topotype of VW. costaricensis), as well as in field observations made
mainly in the states of Chiapas and Oaxaca, I conclude that all these names
have been used for one taxon, the correct name of which is M. ervendbergii A.
Gray.
1987] GRETHER, MIMOSA 2 Oe,
Mimosa hexandra Micheli, Mém. Soc. Phys. Genéve 30(pt. 2, 7): 91. t. 27.
889
Mimosa bimucronata (DC.) Kuntze subsp. hexandra (Micheli) aioe Repert. Spec.
Nov. Regni Veg. 9: 3. 1910, and var. intermedia Hassler, ibid. M a bimucronata
(DC.) Kuntze var. hexandra (Micheli) J. F. Macbr. Contr. Gray Herb. 59: 12. 1919.
Type: Paraguay, bords du Mbay, prés de Paraguari, Oct. 1882, Balansa 4422 (ho-
lotype, G (photo, us!); isotypes, B (photo, us!), F!, Ny!, P).
Mimosa vepres Lindman, Bih. Kongl. Svenska Vetensk. -Akad. Hand. 24(3,7): 46. fig.
2: ee Hs Paraguay, Colonia Risso, 30 _ 1893, Lindman A2263 (holotype,
s, fide Barneby, pers. comm.; isotypes, GH!, us!).
ea coroncoro Sd & Dugand, Caldasia 31 1): 33. 1944. Type: Colombia, De-
partamento Atlantico, entre Palmar de Varela y Ponedera, Finca “El Paraiso,” Aug.
1943, Dugand & Fae ail 3461 (lectotype, co. 16064; isolectotypes, A!, co. 16065
s!).
The original description of Mimosa coroncoro indicates Dugand & Jaramillo
346] (COL) as the type; however, Forero and Ruiz (1983) lectotypified the
species because there are two specimens of that collection at CoL; they selected
coL 16064 as lectotype and cot /6065 as isolectotype.
I am placing Mimosa coroncoro in the synonymy of M. hexandra main|
because the corolla is three-lobed and there are six stamens, characteristics
rarely encountered in the genus. The legume is also very distinctive: exami-
nation of Dugand 3132 (us!; from Finca “El Paraiso,” entre Palmar de Varela
y Ponedera, Departamento Atlantico, Colombia) shows that it has a persistent
margin, even though the authors of the original description indicated that this
was not the case.
This is the first report of Mimosa hexandra in Mexico; observation of fruits
in the field (Isthmus of Tehuantepec, state of Oaxaca) confirms the presence
of persistent margins.
Macbride considered the species to be a variety of Mimosa bimucronata;
however, the very distinctive, completely sessile fruit with very thick coriaceous
valves, the predominantly trimerous flowers, and the fewer (six to twenty),
thicker-textured leaflets (all characters observed in material from Mexico, Co-
lombia, Venezuela, Brazil, and Paraguay) clearly distinguish M. hexandra from
the related M. bimucronata.
Mimosa lacerata Rose, Contr. U. S. Natl. Herb. 5: 141. 1897.
Acanthopteron laceratum (Rose) Britton, N. Amer. Fl. 23: 179. 1928. Type: Mexico,
Puebla, vicinity of Piaxtla, 24 Nov. 1894, Ne/son 2008 (lectotype, here designated,
us!; isolectotype, Ny!).
Mimosopsis glutinosa Britton & Rose, N. Amer. Fl. 23: 178. 1928; not Mimosa glu-
tinosa Malme, Ark. Bot. 23(13): 51. 1931. Type: Mexico, Puebla, near San Luis
Tultitlanapa, July 1908, Purpus 3174 (holotype, us!; isotypes, F!, GH!, Mo!).
Mimosa biuncifera Bentham var. horrida Miranda, Anales Inst. Biol. Univ. Nac.
12: 610. 1941. Type: Mexico, Puebla, cerro NW de Matamoros, 22 March
1941, F. Miranda 1410 (lectotype, here designated, MEXU!).
Two syntypes of Mimosa lacerata, Nelson 2008 (ny!, us!) and Pringle 6247
(F!, GH!, K!, MEXU!), were originally cited; no lectotype has been chosen. I hereby
314 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
select the specimen collected by Nelson because it represents the taxon more
completely.
Britton (in Britton & Rose, 1928, p. 179) transferred the species to the
monospecific genus Acanthopteron, he considered the “legume with marginal
wings deeply irregularly cleft into flat, rigid spinous teeth” to be a generic
character; however, all other characters are those of Mimosa. Mimosa lacerata
is not the only species with lacerate margins of the legume; for example, the
legumes of MM. bahamensis Bentham also have such a margin, even though it
is not closely related to MW. lacerata. Therefore, I cannot consider Acanthopteron
a distinct genus.
Britton and Rose (op. cit.) described Mimosopsis glutinosa based on a fruiting
specimen; the type specimen has abnormal and immature fruits of M. /acerata.
In addition, field observations in the states of Puebla and Oaxaca have dem-
onstrated that some individuals growing in very eroded areas are depauperate,
with the lacerate margin of the legume not well developed; however, such
plants can always be recognized as M. /acerata because of other vegetative and
flower characters, as well as the somewhat lacerate margin and the glandular
dots of the fruit.
Miranda (1941) described Mimosa biuncifera var. horrida, pointing out that
he had not found any other character besides the prickles to distinguish the
plant collected in Matamoros from the typical 7. biuncifera. Miranda did not
cite specimens in the original description, but he mentioned (p. 611) “la planta
de Matamoros” in the protologue and annotated the specimen F. Miranda
1410 (mexu!) as M. biuncifera var. horrida. Considering all aspects of the
protologue and having found the specimen collected and annotated by the
author, I hereby propose Miranda 1410 as lectotype. Examination of that
specimen shows tetramerous flowers with puberulous corollas, as well as the
twinned, laterally compressed, very broad prickles typical of M. /acerata.
Fruiting material collected in Matamoros, Puebla (Miranda 2279, MExv!)
also corresponds to Mimosa lacerata. In addition, herbarium and field obser-
vations, mainly in the states of Puebla and Oaxaca, show that flowering Mimosa
lacerata (e.g., Miranda 1410) has rose to purple heads and rose stamens, and
it can thus be clearly distinguished from M. biuncifera, which has pentamerous
flowers with pubescent corollas, white to slightly rose heads, and white stamens.
Therefore, I consider M. biuncifera var. horrida to be conspecific with M.
lacerata.
Mimosa lactiflua Del. ex Bentham, Trans. Linn. Soc. London 30: 393. 1875;
Martius, Fl. Brasil. 15(2): 311. 1876, descr. ampl. Type: habitat in America
Meridionali (an Brasilia?), ex specimine olim in Horto Monspeliensi
culto, in Herb. D.C. asservato, 1836, 307b (holotype, G-pc!).
Mimosa mixtecana Brandegee, Univ. Calif. Publ. Bot. 3: 379. 1909. Type: Mexico
Puebla, vicinity of San Luis Tultitlanapa, near Oaxaca, May-July 1908, ene 2673
eee la here designated, us! (photo, MEXxu!); isolectotypes, F!, GH!, Mo!, Ny!, UC
(photo, MEXU!
Ay cae Rie Britton & Rose, N. Amer. Fl. 23: 153. 1928. Type: Mexico, Oaxaca,
1987] GRETHER, MIMOSA 315
Distrito de Tlacolula, Cerro de la Carbonera, Matatlan, June 1906, Conzatti &
Vazquez 1482 (holotype, us! (photo and fragments, NyY!); isotype, GH!).
Concerning the occurrence of Mimosa lactiflua in Mexico, Bentham (1875,
pp. 393, 394) stated, ‘““Delile’s specimens were from the Botanical Garden of
Montpellier, supposed to be of American, perhaps Brazilian, origin. In the
Berlin herbarium there is a specimen from Mexico, Ehrenberg, which agrees
with the detailed description I had made (now inserted in the Flora Brasiliensis),
except that the leaflets are under instead of over '/ in. long.”
I (Grether, 1978) cited the species as occurring only in the state of Oaxaca,
Mexico. Since that time, however, numerous specimens from the states of
Morelos, Puebla, Guerrero, and Oaxaca have been examined that clearly cor-
respond to Mimosa lactiflua. Personal communication with R. C. Barneby and
a review of the holdings of different herbaria have yielded no evidence that
this species occurs in Brazil. Besides, I have not seen it in material examined
from Central America.
The holotype of Mimosa lactiflua, seen when it was on loan to Ny from G-pc,
is a flowering specimen characterized mainly by its glabrous, tetramerous flow-
ers and its oblong-lanceolate to elliptic or ovate, glabrous, glaucous leaflets
with prominent reticulate nerves beneath. The lectotype and isolectotypes of
M. mixtecana (Purpus 2673) are flowering and fruiting specimens, also with
glabrous, tetramerous flowers and glabrous, glaucous leaflets, very variable in
shape and size as M. lactiflua. The type of M. vazquezii shows the same flower
characters and variable, elliptic to ovate leaflets. In spite of the uncertain origin
of the specimen cultivated at Montpellier, the holotype of M. lactiflua is a good
specimen, and this is the oldest and correct name for the species.
Mimosa pe cea Mém. Soc. Phys. Genéve 34(3): 277. ¢. 22. 1903.
Type: Mexico, Michoacan, pied du Volcan de Jorullo, 13 April 1898,
Langlassé 99 (haletpe. G; 1sotypes, F!, K! (photo, MEXxv!)).
Mimosa conzatti Britton & Rose, N. Amer. FI. 23: 153. 1928. Type: Mexico, Oaxaca
Distrito del Centro, Cerro San Antonio, 6 aePl 1908, Conzatti 2239 (holotype, cu!
(photo and fragments, Ny!, Us!); isotype, F
Mimosa langlassei was described from a flowering specimen, and the de-
scription of M. conzattii was based on a fruiting one; however, examination
of material of the latter at F, GH, Ny, and us shows remnants of flowers at the
base of fruits; these flowers are tetramerous and the corolla lobes are puberulous,
Flowering and fruiting material of Mimosa langlassei (Michoacan, 18kmN
de La Huacana, cerca del Volcan El Jorullo, R. Grether 1117, MEXU, UAMIZ)
was collected near the type locality; the pubescent and slightly setose valves of
the legume agree with the fruits of 4. conzattii, and the flowers are tetramerous
and puberulous. I also visited the type locality of M. conzattii,; unfortunately,
the area is quite disturbed, and the species is no longer growing there.
Concerning typification of Mimosa conzattli, there 1s a note on the GH, Ny,
316 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
and us sheets of Conzatti 2239 saying “ex herb. Field Mus.” I have studied
the Field Museum specimen distributed by Conzatti as ‘Acacia’, this was
annotated in 1910 by Greenman, who identified it as 4. xanti Gray and sent
fragments to B. L. Robinson (Gu), who verified the identification. Britton and
Rose saw only the specimen of the same number at GH and took even smaller
fragments from it. These were deposited at Ny by Britton and at us by Rose,
and each was mounted with a photograph of the undivided Gu specimen. At
the suggestion of D. H. Nicolson (us), I now recognize the GH specimen as the
holotype of M4. conzattii, the specimens at Ny and us as fragments of the
holotype, and the specimen at F as an isotype.
Examination of additional material from Michoacan, Guerrero, Oaxaca,
Puebla, and Chiapas confirms that it is a single species, Mimosa langlassei.
Mimosa mellii Britton & Rose, N. Amer. Fl. 23: 155. 1928. Type: Mexico,
Oaxaca, near Chivela, 18 Jan. 1927, Mell 2 (holotype, us! (photos,
MEXU!, UAMIZ!
Mimosa chiapensis Britton, N. Amer. FI. 23: 154. 1928. Type: Mexico, Chiapas, river
bottom, erat Monserrate, May 1925, Purpus 10313 (holotype, Ny! (photos
MEX iz!); isotype, Us!).
Mirosa Mola Britton & Rose, N. Amer. FI. 23: 155. 1928. Type: Mexico, Chiapas,
near Los Pinos, 12 Dec. 1906, C. B. Doyle 56 (holotype, us! (photos, MEXU!, UAMIZ!;
fragments, NY!)).
Mimosa oaxacana Britton & Rose, N. Amer. Fl. 23: 155. 1928. Type: Mexico, Oa-
xaca, between Guichocovi and Lagunas, 27 June 1895, Ne/son 2746 (holotype, us!
(photos, MExU!, UAMiz!; fragments, NY!)).
Mimosa mellii, M. chiapensis, M. doylei, and M. oaxacana were all described
by Britton and Rose in the same publication. Although the original descriptions
show some differences (mainly in pubescence of leaflets and corolla lobes), all
of them correspond to a single species.
There are several bases for this conclusion. Types of Mimosa mellii and M.
doylei are fruiting specimens with remnants of flowers; fruits of both are sessile
and slightly setose, and they clearly correspond to the same taxon. The type
specimen of M. doylei has few, tetramerous flowers, like those of M/. mellii—
not sideae ncaiy a as quoted in the original description. The type of M. doylei
has puberul orolla lobes and leaflets, while the type of M. mellii has glabrous
to renee scenes corolla lobes and completely glabrous leaflets.
The types of Mimosa oaxacana and M. chiapensis are flowering specimens;
both have tetramerous flowers, as well as puberulous corolla lobes and leaflets.
I have visited the type locality of Mimosa mellii and have collected a topotype
of that species (Oaxaca, Chivela, R. Grether 1363, MEXU, UAMIZ); I have also
examined other topotypes (Mel/ s.n., Aug. 1928, us!, and Dec. 1928, Ny!). The
type locality of MM. oaxacana (between Guichocovi and Lagunas) is south of
Chivela in the same region. There are several collections from this area, al-
though it is difficult to state which could be considered as a topotype. I located
Hacienda Monserrate through Sousa’s (1969) publication on Purpus’s botanical
collections in Mexico; despite a thorough search of this locality and the vicinity,
I could not find ©. chiapensis there. However, I did collect additional material
southwest of the type locality (Chiapas, Municipio Cintalapa, 9.5 km NW de
1987] GRETHER, MIMOSA 317
Rizo de Oro, camino a Colonia Rodolfo Figueroa, cerca del limite con Oaxaca,
R. Grether 1758, MEXU, UAMIZ). I could not locate the type locality of M. doylei
(Los Pinos) on present or old maps, or by asking local people in Chiapas.
Field observations—as well as examination of type specimens, topotypes of
Mimosa mellii, and additional flowering and fruiting material from Oaxaca
and Chiapas—permit me to state that variation in pubescence of leaflets is
probably due to the stage of leaf development: flowering specimens (May to
July) generally have puberulous leaflets, although some populations show vari-
ation from puberulous to glabrate leaflets even in a single individual, and some
others have leaflets always glabrous. In fruiting specimens (December to Jan-
uary) the leaflets are generally glabrous, although they are puberulous to gla-
brous in a few of them. However, the linear-oblong, strongly reticulate-nerved
leaflets are constant in all flowering and fruiting material examined. The corolla
lobes of M. mellii also vary in pubescence: 1n some individuals they are glabrous
and in others puberulous; in some variation is from puberulous to glabrous
even on a single plant. In addition, the legume varies from slightly setose to
completely glabrous.
Despite the differences mentioned above, it is not possible to distinguish
several species or varieties. Also, the geographic distribution of this taxon is
apparently restricted to the Isthmus of Tehuantepec (Distrito de Juchitan),
Oaxaca, and the adjacent region of Chiapas (Municipio de Cintalapa and Mu-
nicipio de Arriaga), at altitudes between 150 and 1000 m
I have selected Mimosa mellii as the name for the species, considering that
its type is the best and most complete specimen (with mature fruits and rem-
nants of flowers).
Mimosa mollis Bentham, J. Bot. (Hooker) 4: 408. 1842. Type: Mexico, Puebla,
Acatlan, 1834, Andrieux 400 (holotype, k; isotypes, G (photos, F!, MEXU!),
OXF (photo, MExU!), w (photo, F!)).
Mimosa herincquiana Micheli, Mém. Soc. Phys. Genéve 34(3): 276. 1903. Tye:
Mexico, Guerrero, Cariote [Canhén] del Zopilote, 27 May 1899, Langlassé 1040
(holotype, G; isotypes, F!, GH!, K! (photo, MEXU!), Us!).
The types of Mimosa mollis and M. herincquiana are flowering specimens;
the original descriptions show differences only in numbers of pinnae (four or
five vs. seven or eight, respectively) and leaflets (six to ten vs. seven or eight).
Examination of photographs of types, specimens collected near the type
locality of Mimosa mollis (Puebla, 11 km SE de Acatlan de Osorio, M. Sousa
8210, MEXU!), isotypes, and the topotype of M. herincquiana (Guerrero, Canon
del Zopilote, 36 km N de Zumpango del Rio, R. Grether 1143, MEXU, UAMIZ),
as well as additional flowering and fruiting material from Puebla, Guerrero,
and Oaxaca, indicates the similarity of the two taxa, which I consider synon-
ymous.
There are four to ten pinnae and six to twelve leaflets. The tomentose branch-
lets and stipules, the villous oblong to elliptic leaflets, the villous pentamerous
flowers, and the tomentose, unarmed, sessile fruits are distinctive characters
of the species.
318 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Mimosa orthocarpa Spruce ex Bentham, Trans. Linn. Soc. London 30: 437.
1875; Martius, Fl. Brasil. 15(2): 380. 1876, descr. ampl. SyNryPes: Bra-
zil, “habitat prope Santarem provinciae Paraensis,” Riedel s.n. (kK); ad
Lacum Quiriquiry, Prov. Para, 1850, Spruce 518 (k, herb. Bentham; Ny
neg. 1897!).
ssa glandulosa Bong. ex Bentham, Trans. Linn. Soc. London 30: 437. 1875, nomen
1udum.
Mimosa calderonii Britton & Rose, N. Amer. Fl. 23: 167. 1928. Type: El Salvador,
El Angel, Oct. 1923, S. Calder6n 1842 (holotype, us! (photo and fragments, Ny!);
ae GI Hl).
Two syntypes of Mimosa orthocarpa, Riedel s.n. and ““Sello”’ 518 were orig-
inally cited. According to Barneby (pers. comm.), ‘‘Sello”” must be an error for
Spruce, considering that Sello was never on the Amazon. Bentham attributed
the epithet to Spruce, and the specimen at K in Bentham’s herbarium is labeled
Spruce 518.
I have examined a photograph from s (F neg. 1350) and a specimen at Ny,
both labeled “Spruce s.n., ad ripas fluminis das Trombétas et lacus Quiriquiry,
Prov. Para, Dec., 1849.” Barneby has examined other specimens labeled Spruce
s.n. at K (herb. Hooker), LE, and w. The specimens Spruce s.n. could be from
the same collection as Spruce 518, but they have different collection dates (Dec.
1849, and 1850, respectively). According to Urban (1906), Spruce was at Qui-
riquiry in December, 1849. If that is so, then the date on Bentham’s sheet
could be an error; however, there is no doubt that the specimens Spruce s.n.
and Spruce 518 are conspecific.
Bentham (1875) considered Mimosa glandulosa to be a synonym of M.
orthocarpa, based on the specimen named by Bongard. After examining two
specimens originally named M. glandulosa Bong. (Santarem, Nov. 1828, Riedel
37, A, K, Riedel 1560, Le), Barneby (pers. comm.) confirmed that . glandulosa
Bong. ex Bentham is a nomen nudum and a synonym of M. orthocarpa.
Barneby’s and my examinations of type specimens of Mimosa orthocarpa
and M. calderonii, of additional material from Mexico (states of Guerrero,
Oaxaca, Veracruz, Tabasco, and Chiapas), Colombia, Venezuela, and Brazil,
as well as my study of original descriptions, support this synonymy.
I have not chosen a lectotype of Mimosa orthocarpa because I have seen
a photograph only of Spruce 518.
Mimosa polyantha Bentham, J. Bot. (Hooker) 4: 410. 1842. Type: Mexico,
Puebla, Acatlan, Andrieux 397 (holotype, kK; isotype, w (photos, F!,
MEXU!)).
Mimosa ails a ee oi Soc. Nat. Hist. 31: 260. 1904. Typ
Me pe CTO: mountains above Iguala, 5 Oct. 1900, Pringle 8408 pocece
GH!; 1sotyp , MEX a ve NyY!, vet),
ae stipitata ees Proc. Boston Soc. Nat. Hist. 31: 261. 1904. Type: Mexico,
ee rero, On mountains above Iguala, 5 Oct. 1900, Pringle 8406 (holotype, GH!:
sotypes, F!, K!, MEXU!, Ny!, Us!).
Tree setigera Britton & Rose, N. Amer. FI. 23: 160. 1928. Type: Mexico, Sinaloa,
1987] GRETHER, MIMOSA 319
vicinity of Rosario, 14 April 1910, Rose, Standley, & Russell 14553 (holotype, us!;
isotypes, GH!, NY!).
The fruits of Mimosa polyantha were unknown to Bentham; however, Rob-
inson (1898) described them, and he assumed material with oblong legumes
abruptly acuminate at each end, hispid on the margins, and with valves having
short, spreading setae to be typical, based on the specimens Pringle 4635
(mExu!), Rose 1475, and Palmer s.n.
I have examined topotypes of Mimosa a vantha (Puebla, 4 km SE de Acatlan
de Osorio, R. Grether 735, MEXU, UAM 1 km SE de Acatlan, Té/lez 1086,
MEXU!; Acatlan, F. Miranda 2971, on and the legumes correspond to
Robinson’s description of them.
Mimosa Stipitata and M. polyanthoides were collected in the same place. |
have visited the type locality and vicinity and have observed variation in
number of pinnae and leaflets, as well as in density of setae and length of the
stipe of the legume, even in the same population (Guerrero: 22 km W de
Iguala, camino a Teloloapan, R. Grether 1132, MEXU, UAMIZ; 6 km W de
Xalostoc, camino a Teloloapan, R. Grether 1133, MEXU, UAMIZ).
Mimosa setigera was based on a specimen with setose legumes; however,
examination of material from Rosario, Sinaloa, and vicinity (Sinaloa: 16 km
SE de Escuinapa, R. Grether 1099, MEXU, UAMIz; between Agua Caliente and
Rosario, Rudd 2099, mexu!, Rudd 3000, MExv!) also shows variation in valves
(from setose to glabrous) and differences in the length of the legume stipe.
Differences in number of pinnae and leaflets depend on the season, because
flowering specimens have immature leaves, while fruiting material has mature
and old ones. Flower characters are constant for all material examined from
Sonora, Sinaloa, Michoacan, Guerrero, and Oaxaca, as well as from Puebla,
Morelos, and Veracruz.
Mimosa pueblensis R. Grether, nomen novum
eee filipes Britton & Rose, N. Amer. FI. 23: 177. 1928. Mimosa filipes (Britton
Gentry, Brittonia 6: 315. 1948, not Martius, Herb. FI. Brasil. 132. 1837.
TYPE: Me xico, Puebla, vicumty of San Luis Tultitlanapa, July 1908, Purpus 3175
(holotype, us!; isotypes, GH!, Mo!).
I am proposing a new name for the species because the epithet fi/ipes used
by Britton and Rose was not available. It had been used by Martius for a
different Brazilian Mimosa, making the Britton and Rose name a later hom-
onym.
Mimosa pueblensis is known only from the states of Puebla and Oaxaca,
Mexico; although Britton and Rose mentioned the state of Morelos, I have not
seen material from there.
The species is characterized mainly by its slender, puberulous peduncles with
red glandular dots, axillary, solitary or in clusters of two to six (to ten); 1ts deep
purple buds and flowers; its deeply five- (rarely four-)lobed, glabrous to pu-
berulous corolla; and its sessile, glabrous legume, with red glandular dots more
conspicuous in young fruits, and the margin sparingly prickly or unarmed.
Britton and Rose (1928) cited Purpus 3175 as the type of Mimosopsis filipes;
320 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
however, the original description was based on two specimens, Purpus 3175
(flower) and Purpus s.n. (flower and fruit). Both were annotated as type, and
both were collected in the same locality on the same date; there 1s no doubt
that they are conspecific.
Mimosa rhodocarpa (Britton & Rose) R. Grether, comb. nov.
Mimosopsis rhodocarpa Britton & Rose, N. Amer. Fl. 23: 175. 1928. Type: Mexico,
Michoacan, Patamban, Jan. 1903, Nelson 6550 (holotype, us!; isotype, GH!).
A new combination is necessary to transfer the species to the genus Mimosa.
Mimosopsis Britton & Rose 1s an artificial genus: the only distinctive character
is the unsegmented legume; all other vegetative and flower characters are those
of Mimosa
The species is distinguished by its oblong, glabrous, rather thick leaflets with
ciliate margins; its five-lobed, glabrous calyx about half as long as the corolla
and with a ciliate margin; its five-lobed, glabrous, purple corolla; and its sessile,
reddish, puberulous to glabrous, shiny, reticulate, completely unarmed legume
3-4.5 cm by 8-10 mm. Mimosa rhodocarpa has the broadest fruit of all the
related Mexican species.
Mimosa rhodocarpa is known from the states of Zacatecas, Jalisco, Michoa-
can, México, Hidalgo, Puebla, Guerrero, and Oaxaca.
Mimosa ursina Martius, Flora 21(2), Beibl. 4: 56. 1838. Type: Brazil, Prov.
Bahiensis, inter Conceicao et Arrayal da Feira de S. Anna in desertis,
II-llI, 1819, Martius s.n. (holotype, Mm).
Mimosa paucisperma Britton & Rose, N. Amer. FI. 23: . 1928. Type: Mexico,
Chiapas, ear Arriaga, Sept. 1923, Pere 9306 ea uc (photo and frag-
ments, Ny!, us!)).
Barneby examined the holotype of Mimosa ursina at m, and he and I ex-
amined photographs and fragments (branchlets, leaves, flowers, and fruits) of
the type collection of M. paucisperma at Ny and us; the characters of the type
material are in accord. In addition, the original descriptions of both species
are complete, and all characters, including those of habitat (in savannas and
flooded places) clearly agree.
Our review of additional material from Brazil, Honduras, El Salvador, and
southern Mexico (states of Oaxaca, Tabasco, and Chiapas), including a topo-
type of Mimosa paucisperma (Chiapas, alrededores de Arriaga, salida de la
carretera a Tapachula, R. Grether 1783, MEXU, UAMIZ), indicates that it com-
prises only one species.
Detailed observation of herbarium specimens and fresh material shows some
flower characters not considered in the original descriptions of Mimosa ursina
and M. paucisperma, including a glabrous, four-lobed corolla, four stamens,
and a widened stigma.
Mimosa watsonii Robinson, Proc. Amer. Acad. Arts 36: 473. 1901. Type:
Guatemala, eastern portion of Vera Paz and Chiquimula, 1885, Watson
323 (lectotype, here designated, Gu!; isolectotype, us).
1987] GRETHER, MIMOSA 321
Mimosa recordii Britton & Rose, N. Amer. Fl. 23: 170. 1928. Type: British Honduras,
Stann Creek District, Middlesex, 19 Jan. 1926, Record s.n. (holotype, us!; isotype,
NyY!).
Mimosa rekoana Britton, N. Amer. Fl. 23: 170. 1928. Type: Mexico, Oaxaca, Cafetal
eee as Espino), 20 Nov. 1917, Reko 3610 (holotype, us! (fragments, Ny!);
isotype, u!).
Mimosa ee Britton, N. Amer. Fl. 23: 169. 1928. Type: Honduras, Department
of Atlantida, vicinity of Tela, 14 Dec. 1927-15 March 1928, Standley 54698 (ho-
lotype, NY!; isotypes, A!, F!, Us!).
Robinson described Mimosa watsonii from flowering and fruiting material
(Watson 185 and Watson 323, respectively), I am here selecting Watson goo
as lectotype because fruiting material is more distinctive of the species than
the flowering specimen.
The species was originally characterized by its leaves with two pairs of pinnae,
the lower pinnae bearing one or two pairs of leaflets and the upper ones with
two or three pairs; terminal leaflets up to 5 cm long; tetramerous flowers with
a four-lobed corolla and eight stamens; and 5 cm by 7-10 mm, articulate,
glabrous and finely papillose pods unarmed except for a few scattered, minute,
recurved spines on the tomentulose replum.
Mimosa recordii was described by Britton and Rose; M. rekoana and M.
resinifera by Britton. The descriptions of these three taxa were based on flow-
ering material and were published in North American Flora.
Standley and Steyermark (1946) included Mimosa watsonil, M. resinifera,
and M. recordii in the Flora of Guatemala. The authors considered M. rekoana
to be a synonym of M. recordii and described its legume, which is like that of
M. watsonii. The amplified description of M. resinifera given by Standley and
Steyermark also agrees with that of M. watsonii, even though the fruit was not
described.
I have examined type specimens and additional material from Mexico (states
of Guerrero, Oaxaca, Veracruz, Tabasco, and Chiapas), Guatemala, Belize, and
Costa Rica. Although the presence of resinous dots on the lower surface of the
leaflets was cited as a distinguishing character for Mimosa resinifera, these are
present in the other three type specimens, as well as in all additional material
examined. Other constant leaf characters include the cupular gland at the petiole
base, some cylindrical glands along primary and secondary leaf rachides, and
the pubescence and reticulate nerves of the leaflets. Great variation has been
observed in the number of pinnae and leaflets: from two pairs of pinnae with
one to three pairs of leaflets, as Mimosa watsonii was originally described, to
two or three pinnae with two to five leaflets per pinna, to two to four pinnae
with four to seven leaflets, to five or six pinnae with four to nine leaflets. The
leaflets also vary from 2.5 to 12 cm in length, and from 1.5 to 6 cm in width.
Intermediate combinations are frequent and make it difficult to delimit several
taxa.
I have analyzed geographic eee altitudinal range, vegetation types
where the species grows, and flowering and fruiting times but have not found
it possible to delimit subspecific re oe the accumulated data.
The flowers are arranged in large panicles of white heads and the corolla is
four- (rarely 5-)lobed, glabrous, and with few or no resinous dots on the lobes
a2 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
in all types and additional material examined. The fruits are sessile or very
slightly stipitate, with five to eleven segments, glabrous, and with resinous dots
on the valves on the type specimen of Mimosa watsonii, as well as on additional
material. Because flower and fruit characters are exactly the same for the four
taxa originally described, and there are several constant leaf characters, I con-
sider these four taxa to be a single species, Mimosa watsonii.
ACKNOWLEDGMENTS
I wish to express my appreciation to Mario Sousa, Departamento de Botan-
ica, Instituto de Biologia, UNAM, for his constant assistance and for critically
reviewing the manuscript; Richard S. Cowan, then of the Department of Bot-
any, Smithsonian Institution, for his valuable suggestions during my visits to
us; and Bernice G. Schubert, Arnold Arboretum, Harvard University, and
Rupert C. Barneby, New York Botanical Garden, for their invaluable aid and
their critical reviews of the manuscript.
This study was partially supported by the Consejo Nacional de Ciencia y
Tecnologia, México: Convenio CONACyT:BID-UAMI (Grant PCECBNA-
000914).
LITERATURE CITED
BeNnTHAM, G. 1875. Mimosa. In: Revision of the suborder Mimoseae. Trans. Linn.
Soc. London 30: 388-441.
1876. Mimosa. In: C. F. P. von Martius, FI. Brasil. 15(2): 294-390.
Britton, N.L.,& J. N. Rose. 1928. are Neomimosa, Mimosopsis, Acanthopteron.
In: Mimosaceae. N. Amer. FI. 23: 144-179.
Forero, E., & R. Ruiz. 1983. Tipos : Leguminosae-Mimosoideae en el Herbario
Nacional Colombiano. Mutisia 57: 1-6.
GReTHER, R. 1978. A general review of the genus Mimosa (Leguminosae) in Mexico.
Bull. Int. Group Study Mimosoideae 6: 45-50.
. Notes on the genus Mimosa in ey ce Ibid. 12: 43-48.
Mian, F. 1941. Estudios sobre la vegetacin de México 1: La vegetacion de los
cerros del sur de la Meseta del Anahuac. El cuajiotal. ie Inst. Biol. Univ. Nac.
Mexico 12: 569-614
— B.L. 1898. Revision of the North American and Mexican species of Mimosa.
c. Amer. Acad. Arts 33: 305-331.
——. 1904. Notes on eee in Mexico and Central America. Proc. Boston
c. Nat. Hist. 31: 258, 259.
ee v. E. 1984. Identity of sone Mexican Acacia and ne poe Cad as
and Galeotti. Anales Inst. Biol. Univ. Nac. México, Ser. Bot. 47-53: 4.
Sousa, M. 1969. Las colecciones botanicas de C. A. Purpus en “México, ae 1898-
1925. Univ. Calif. Publ. Bot. 51: 1-36
STANDLEY, P. C., & J. A. STEYERMARK. 1946, Mimosa. In: Flora of Guatemala. Field-
lana, Bot. 24(5): 52-64.
Ursan, I. 1906. ea collectorum botanicorum. /n: C. F. P. von Martius.
Fl. Brasil. 1(1): 1
1987] WEITZMAN, FREZIERA 323
TAXONOMIC STUDIES IN FREZIERA (THEACEAE),
WITH NOTES ON REPRODUCTIVE BIOLOGY
ANNA L. WEITZMAN!
Three new species of Freziera, one each from Venezuela (from the Guayana
Highland), Colombia, and Ecuador (both from the Andes), are described, il-
lustrated, and compared to related species. The monotypic genus Patascoya
is reduced to synonymy in Freziera, and the appropriate combination is made.
Observations of herbarium specimens and natural populations suggest an un-
equal sex ratio in this dioecious genus, with carpellate plants predominating.
This is the reverse of the situation in most tropical forest trees.
Freziera Willd. is a Neotropical genus of trees mostly distributed in cloud
forests in northwestern South America. It is easily recognized by its alternate,
distichous leaves and its axillary clusters of flowers. Most species grow at high
altitudes, close to the upper limit of cloud forests. A few species grow at lower
altitudes in moist coastal regions in Colombia, Panama, and Venezuela.
Species of Freziera are trees 5 to 15 (to 35) m tall, or rarely shrubs. The
leaves of all species are alternate and distichous. Flowers are axillary and
solitary or in racemose fascicles of two to seven (to 15). The pedicel of each
flower is subtended by a single bract (or by two bractlike structures in some
species with exclusively solitary flowers). As in most Theaceae, each flower has
two bracteoles; in Freziera they are nearly always apical on the pedicel and
often appear to be part of the calyx, since they are attached to the floral
receptacle and may be quite sepaloid in appearance (FIGURES Ic, 2d). The
corolla of all species of Freziera is urceolate and thickened above. The thick-
ening is made up of sclereids, although in the field the corolla looks and feels
quite waxy. The petals spread only at the tips, and the opening is, as far as I
have observed, | mm or less in diameter. The stamens and stigma are well
within the flowers. The pollen grains of Freziera are small, averaging ca. 10
um in diameter, and copious. The fruits are berries, which are nearly always
immature on herbarium specimens. Although carpellate plants usually have
everything from buds to large green fruits, I did not find mature fruits (re-
portedly blue or black) in the field. Seeds from the largest green fruits fail to
germinate, implying that they are immature
Species of Freziera occur in cloud and Hoi coastal forests at elevations up
to 3500 m, and the genus is distributed in the West Indies (Cuba, J amaica,
and the Lesser Antilles), southern Mexico, Central America, and much of South
‘Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138. Present
address: Department of Botany, Smithsonian Institution, Washington, D. C. 20560, and Gressitt
Center for Entomological Research, Bishop Museum.
© President and Fellows of Harvard College
Journal of the Arnold Arboretum 68: 323- 334, ca 1987.
324 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
America (the Guayana Highland, the Venezuelan coastal cordillera, and the
Andes south to central Bolivia).
Morphologically, Freziera has gynodioecious flowers; however, all species
for which there are sufficient data are functionally dioecious. Carpellate plants
have flowers with staminodia and a functional gynoecium. The flowers in
staminate plants have functional stamens and usually have what appears to be
a functional gynoecium but nearly always fall off soon after anthesis. Only three
collections of one species (F. chrysophylla Bonpl.) have so far been observed
to be hermaphroditic. These specimens have flowers with nearly 100 percent
viable pollen (tested with cotton blue in lactophenol) on the same branch as
fruits. The flowers may be truly hermaphroditic, or the flower type may have
changed as the branch matured.
In herbaria there are far more specimens of Freziera representing carpellate
plants than staminate ones. Of 448 collections of about 29 species, 303 (67.6%)
are carpellate, 102 (22.8%) are staminate, and 43 (9.6%) either are sterile or
have buds too young for determination if anthers would develop or not. In 86
specimens of Freziera candicans Tul., the ratio is 62:20:4 (72.1:23.3:4.6
percent), respectively, and in 49 of Freziera canescens Bonpl., 40:5: 4 (81.6:
10.2: 8.2 percent). This unequal sex ratio in collections may be due to collection
artifact or to unequal sex ratios in natural populations. I suspect that one reason
for the preponderance of carpellate specimens may be that botanists, in trying
to collect what appear to be better specimens, select those with buds, flowers,
and fruits (i.e., specimens from carpellate plants), rather than ones with only
buds and flowers. Since staminate and carpellate flowers are externally identical,
many ete a do not realize the importance of separate collections
of the two s
There ae be a truly eee sex ratio in Freziera. It is usually harder to
find staminate than carpellat ividuals in the field; in some small populations
(about eight to ten observed individuals) I was unable to find any staminate
plants at all. According to Opler and Bawa (1978, and references therein),
dioecy is more common in tropical ecosystems than in temperate ones (see
also Bawa, 1980), and dioecious plants often have sex ratios that depart from
unity. Forty-four percent of the dioecious tropical forest trees they studied had
sex ratios departing significantly from unity: of ten species, eight were biased
toward a greater number of staminate individuals, and only two were carpellate-
dominant (both were members of the Polygonaceae, a family known for car-
pellate-dominant sex ratios (Opler & Bawa, 1978)). Lloyd (1973) found that
when sex ratios are skewed, perennials tend to show an excess of staminate
plants, the reverse of the apparent situation in Freziera.
Explanations advanced for carpellate-dominant sex ratios in plants include
differential survival rates, differential reproductive maturation, and seral po-
sition (Opler & Bawa, 1978). Further field study of Freziera is necessary since
my observations of individual sex ratios are anecdotal rather than quantitative.
If the genus is really carpellate-dominant in natural populations, it is very
unusual among tropical trees.
The three new species and one new combination proposed below result from
work on a monograph of the entire genus.
1987] WEITZMAN, FREZIERA Bae,
Freziera carinata A. Weitzman, sp. nov. FIGuRE 1.
A speciebus aliis Frezierae in ramulis alatis, foliis auriculis basalibus demum
revolutibus (foliis ut videtur base abrupte attenuatis), et costis petiolis carinatis,
differt.
Small tree 2-9 m tall; mature branches terete; twigs dorsoventrally flattened,
with narrow paired wings decurrent from base of petiole keel and descending
through 2 internodes; bark dark red-brown, papillate when young, striate and
splitting with age, glabrous or occasionally very short-strigose-glabrescent; len-
ticels few, large, very narrowly to widely elliptic, appearing late; terminal bud
conduplicate-involute, (2-)4-6.3 cm long, finely strigose. Leaves with petiole
(0.1-)0.3-0.6(—-1.8) cm long, erectly winged, canaliculate above, keeled below,
glabrous; colleter(s) 1 to several in petiole base, linear or triangular, flattened,
red to black; blade elliptic or narrowly obovate, (4.1-)9.2-14.8 by (2.1-)2.9-
4.9(-6) cm, coriaceous, the base rounded, ciliolate, auriculate, with auricles
becoming revolute (base then appearing attenuate), the apex acute, short-acu-
minate, ultimately retuse, terminating in caducous, thick, conical, black seta,
the margin finely serrate, teeth (46 to) 71 to 95 (to 122) per side, with caducous,
thick, conical or slightly curved, forward-pointing, black setae inserted in the
sinuses (rarely—only in specimens from Cerro de la Neblina—with few thin
hairs surrounding base of each seta), the surface glabrous above, densely short-
strigose (rarely glabrous) below, with small papillae densely and evenly dis-
tributed above and below, and larger ones on midrib in horizontal rows above
and scattered below, the midrib flat with small central ridge above, keeled
below, the lateral veins (16 or) 17 to 24 (to 31) per side, flat to slightly rounded
above, prominently rounded below. Inflorescence axis 0.5—2.5 mm long, with
flowers | to 5, pedicel scars absent or | to 5 and contiguous; floral bract
persistent, triangular, |.1-3.1 by 0.8-1.5 mm, sclerotic, the base clasping, the
apex acute to rounded, terminating in thick, conical, black seta, the margin
entire, sometimes with several black setae and/or flaps, sometimes ciliolate,
the outer surface sparsely to densely sericeous; pedicel erect, cylindrical, 3.1-
6 by 0.7-1.1 mm, glabrous to strigose; bracteoles 2, apical on pedicel, subop-
posite, persistent, sepaloid, seemingly part of calyx, broadly to very broadly
ovate, equal or unequal, 1.4—2.4 by 1.5—2.4 mm (lower), 1.8-3 by 1.9-2.9 mm
(upper), sclerotic basally and chartaceous above, the base clasping or cordate,
the apex rounded, with terminal or subterminal (on outer surface) thick, conical,
black seta on lower (or rarely both) bracteole(s), the margin ciliolate, with basal
conical, dark setae, the outer surface sparsely strigose or centrally glabrescent.
Flowers 4.7—7 by 3.1—4.1 mm; sepals 5, broadly ovate, nearly equal, 2.2-3.8
by 2.1-3.1 mm, sclerotic basally and chartaceous above, the base cordate, the
apex rounded and often splitting, the margin membranaceous, minutely cil-
iolate, with dark or pale basal flaps, the outer surface glabrous to minutely
strigose, the inner surface glabrous; corolla urceolate, the petals 5, slightly
connate basally, ovate, nearly equal, 3-5.8 by 1.5-2.5 mm, membranaceous
in lower 4, sclerotic above, apically acute, recurved at anthesis. Staminate
flowers with stamens (14 or) 15, uniseriate, free or slightly adnate basally,
unequal, unordered, the filaments unequal, flat, linear, ca. 0.9 and ca. 1.4 mm
326 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
oe
Ficure 1. Freziera carinata: a, habit; b, undersurface of leaf; c, flower; d, petal of
carpellate flower, stamens adnate; e, gynoecium of carpellate flower; f, ovary and stamens
of staminate flower; g, seeds, side and chalazal views; h, fruit (b, f from holotype; c-e,
g, h from Maguire & Maguire 35334).
1987] WEITZMAN, FREZIERA 327
long, the anthers linear, equal, 0.8-0.9 mm long, lightly pigmented, basally
cordate, the apiculus ovate, ca. 0.1 mm long, apically rounded; gynoecium
conical, 2.4-3.2 by 1.5-1.7 mm, the ovary 3-locular, with locules ca. 1.5 mm
long, each containing ca. 12 ovules, the style tapering, the stigmatic lobes 3,
erect, 0.25-0.35 mm long, dark, minutely papillate. Carpellate flowers with
staminodes 15 (or 16), uniseriate, free, linear, flat and rarely with peripheral
flaps, equal or unequal, 0.6-1.6 mm long, apically rounded; gynoecium conical,
3.2-4.9 by 1.7-2.1 mm; the ovary (2- or) 3-locular, with locules 1.3-1.8 mm
long, each containing 16 to 30 ovules, the style tapering, the stigmatic lobes (2
or) 3, erect, 0.2-0.3 mm long, dark and minutely papillate. Immature fruits
globose, tapering abruptly into persistent style, 6.9-7.7 by 4.9-5.7 mm, green;
mature fruits unknown but reportedly blue; immature seeds (6 to) 16 to 29 per
locule, reniform, 1.2-1.4 mm long, dark red, the testa reticulate.
Type. Venezuela, Edo. Bolivar, Auyan-tepui, cumbre de la parte central oc-
cidental (divisidn occidental del cerro), vecindad del “Drizzly Camp,” sobre
piedra de arenisca, a lo largo de afluente del Rio Churtin, 1760 m, 4 May 1964,
J. Steyermark 93366 (bud, é fl’—holotype, GH; isotypes, NY, U (N.v.), US, VEN).
ADDITIONAL SPECIMENS EXAMINED. Venezuela. TERR. FED. AMAZONAS: Serrania Yutajé,
Cerro Yutajé, Rio Manapiare, 2100 m, Maguire & Maguire 35334 (@ fl, fr; Ny (3 sheets));
Cerro de la Neblina, Rio Yatta, NW head of Canon Grande, 2000 m, Maguire et al.
42322 (young fr; Ny); Cerro de la Neblina, limite Venezuela-Brasil, altiplano, 1800-2000
m, Ewel 177 (bud, young fr; My, Ny); Cerro de la Neblina, Camp VII, 5.1 km NE of
Pico Phelps, 1730-1850 m, Nee 30641] (mixed coll., bud, 6 & @ fl, fr; liquid-preserved
material Gu, duplicates yet to be distributed); Dpto. Atabapo, below Salto Los Monos
on tributary of headwaters of Rio Iguapo, 3°35'N, 65°23'W, 1500-1600 m, Liesner 18515
(bud, fruit; GH); Dpto. Atabapo, gallery forest and open area on Plateau of Huachamacari,
3°50'N, 65°25'W, 1720 m, Liesner 18073 (bud; GH). Epo. BoLivar: Disto. Cedeno,
Serrania Guanay, sector NW, en las cabeceras mas orientales del Rio Paraguaza, 5°55'N,
66°23'W, ca. 1700 m, Huber 11003 (2 bud, fl, fr; Ny); Meseta de Jaua, Cerro Sarisarifama,
cumbre, porcion NE, interior de la Sima Mayor, 4°41'N, 64°13’W, 700 m, Brewer-Carias
s.n. (6 bud; ven); Cerro Guaiquinima, cumbre, sector NE, cerca del borde, cabeceras de
brazo NE del Rio Carapo, 5°59’N, 63°25'’W, 1490-1500 m, Stevermark et al. 117329 (6
bud; MO, NY, U, VEN); Auyan-tepui, no further locality or date, Pannier & Schwabe s.n.
(2 bud, young fr; ven); Auyan-tepui, Valle Encanto, lado derecho del Salto Angel, Fo/dats
7135 (bud; ven); Auyan-tepui, plateau, central E section of NW arm, 5°56'N, 62°34'W,
1850 m, Prance & Huber 28302 (bud, ¢ fl, fr; GH); Chimanta Massif, SE- facing upper
shoulder of Apacara- -tepul, below summit, 2000-2100 m, Stevermark 75782 (2 bud, fr;
sheets)); Chimanta Massif, altiplanicie en los farallones superiores de Apacara-tepui,
sector N del Macizo, 5°12’N, 62°12’ W, ca. 2200 m, Steyermark et al. 128337 (fr; GH,
MO (2 sheets), VEN); Chimanta Massif, ere SE, amplia altiplanicie en la secci6n NE
del Acopan-tepui, en las cabeceras del Rio Yunek, 5°12’N, 62°5'W, 1950 m, Huber et
al. 10118 (bud, ¢ fl; ny); Ptari-tepui, along base of E-facing high sandstone bluffs, 2410-
2450 m, Stevermark 59937 (bud; A, F); Cerro Venamo (parte SW), cerca de los limites
con la Guayana Inglesa, a lo largo del afluente W aes el Rio Venamo, 950-1150
m, Steyermark et al. 92345 (bud, @ fl, fr; GH, K, EN); Cerro Roraima, no further
ee 2000 m, Ule 8726 (bud, 2 fl; k); Cerro Rorima, forested SW-facing quebrada
2In the specimen citations below, I state flowering condition and sex for the flower stage observed;
if no sex is indicated, floral material was insufficient for examination
328 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
near Rondén Camp, 2040-2130 m, Stevermark 58697 (bud, 2 fl, young fr; A, F); Cerro
Roraima, trail through cloud forest to summit, 5°12'N, 60°40'W, — 2600 m, Luteyn
& Aymard 9767 (bud, ¢ fl, fr; GH, NY, U, VEN), 9772 (CAS, GH, NY, VEN).
Freziera carinata 1s characterized by its winged stems and its prominently
keeled petioles and midribs. The leaf blades are auriculate, but the auricles
become revolute very early and the blades then appear attenuate. Specimens
of F. carinata are unusual: the young growth of most other species dries dark
or light brown or rarely green, while that of F. carinata usually dries orange-
brown with paler orange spots. Like most species of Freziera for which phe-
nology is known, F. carinata flowers throughout the year.
Kobuski (1941), who recognized only one species from the Guayana High-
land, apparently did not see any specimens of Freziera carinata for his mono-
graph of the genus. However, despite having seen only a photo and a leaf
fragment of the type of F. roraimensis Tul. and no material of F. guianensis
Klotzsch ex Wawra, he correctly placed the latter in synonymy under F. ro-
raimensis, stating (p. 490), ““F. roraimensis and F. guianensis were collected at
he same locality by the same collector. There Is no doubt in my mind that
only one good species exists in this locality. ...”’ Although the name Freziera
roraimensis has been used by Kobuski and subsequent workers in all deter-
minations of Guayana Highland material, that species has not been re-collected
since Schomburgk found it in November 1842 in the vicinity of Mt. Roraima.
All other known material from the Guayana Highland belongs to F. carinata.
Freziera carinata has been collected on most of the larger tepuis so far visited
except Duida. The two species now recognized from the Guayana Highland
region can be distinguished by use of the following key:
Twigs flattened; midribs and petioles strongly keeled: twigs and leaves glabrous to mi-
nutely strigose: leaf blades elliptic or setae obovate, (4.1-)9.2-14.8 by (2.1-)2.9-
4.9(-6) cm; flowers 4.7-7 by 3.1-4.1 mm. 2.0.00... 0.0 ee F. carinata.
Twigs terete; midribs and petioles sere twigs and leaves densely ade -sericeous and
short-villous, leaf blades narrowly elliptic, 8.1-9.7 by 2.6-3.6 cm; flow
Me TMI. ot vere sues ahaa Pe oes Sees a eeds Ate ati enaer Fy roraimensis.
Freziera echinata A. Weitzman, sp. nov. FIGURE 2.
A speciebus aliis Frezierae in ramulis et foliis utrinque pilis erectis densis
longis persistentibus praeditis, setis erectis pilis erectis cingentibus in margi-
nibus foliis instructis, et bracteolis sepalisque dense longe sericeis extus paginis
totis et intus versus apices acutes, differt.
Tree ca. 5 m tall; mature branches and twigs terete, dark red-brown, papillate,
conspicuously ridged below each side of leaf base, finely striate elsewhere, very
densely golden-hirsute, the hairs persistent, erect, of 2 lengths (ca. 3 and 0.5
mm), the lenticels ovate, 0.4-0.6 mm across, splitting vertically; terminal bud
conduplicate-involute, 4-6 cm long, erect-hirsute. Leaves with petiole 2-3 mm
long, erectly winged, canaliculate, hirsute above and below; blade narrowly
ovate, 10.4-12.3 by 2.6-3.5 cm, subcoriaceous, the base unequal with sides
asymmetric, truncate or rounded on long side, cuneate to truncate and revolute
on short side, the apex long-acute, terminating in caducous, thick, conical,
1987]
Se
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WEITZMAN, FREZIERA
320
Ficure 2. Freziera echinata: a, habit; b, leaf undersurface (hairs omitted); c, leaf
margin; d, flower; e, inside of outer sepal; f, stamens; g, style and stigma (from type).
black seta, the margin entire, slightly revolute, with numerous erect, articulated,
conical, black setae ringed by longer erect hairs, the upper surface sparsely
hirsute (densely so on midrib) and densely papillate, with hairs persistent, erect,
up to 3 mm long, the lower surface densely hirsute, the midrib sunken above,
prominently rounded below, the lateral veins | 1 to 13 per side, inconspicuous,
slightly sunken above, prominently rounded below. Inflorescence axis less than
1 mm long, with 3 to 5 flowers; floral bract persistent, ovate, 6.2-9.3 by 2.1-
3.3 mm, sclerotic, keeled, the base clasping, the apex acute, terminating in
330 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
thick, conical, black seta, the margins entire, the surface densely long-tomen-
tose; pedicel erect, angled, ca. 1 by | mm, densely long-tomentose; bracteoles
2, persistent, sepaloid, ovate, nearly equal, 5-6.4 by 2.6-3.3 mm, sclerotic, the
base clasping, the apex acute, the surface densely tomentose outside, tomentose
near upper margin inside. Staminate flowers 8.5-9.6 by 5-5.5 mm; sepals 5,
ovate, unequal, 6-6.8 by ca. 2.9 mm (outer), 4-5.6 by 2-2.7 mm (inner),
sclerotic, the base clasping, sometimes with dark lobes or few dark basal setae,
the apex acute, the margin ciliolate, the surface densely tomentose outside,
tomentose on upper | (outer sepals) or glabrous (inner ones) inside; corolla
urceolate, white, the petals 5, distinct, narrowly ovate, unequal, 6-7.7 by 2.2-
2.5 mm (outer), 5.5-6 by 1.6-1.7 mm (inner), membranaceous in lower 4-4,
sclerotic above, apically acuminate; stamens 15, uniseriate, free or slightly
adnate at very base, the filaments unequal, geniculate or linear, 1.1-2.2 mm
long, flattened at base, cylindrical above, the anthers |.1-1.2 mm long, con-
nective pigmented, the apiculus 0.05-0.13 mm long, apically rounded and with
terminal seta; gynoecium narrowly conical, 3.8-4.9 by ca. | mm, the ovary
3-locular, with locules ca. 1.1 mm long, each containing ca. 60 ovules, the style
1.8-2.1 mm long, the stigmatic lobes 3, erect, 0.4-0.7 mm long, dark, minutely
papillate. Carpellate flowers and fruits unknown.
Tyre. Colombia, Dpto. Cauca, Parque Nacional Munchique, km 50-55 along
road above Uribe, 2256-1875 m, 25 April 1979, J. L. Luteyn, M. Lebrén-
Luteyn, & G. Morales L. 7448 (bud, 4 fl—holotype, Ny; isotypes, AAU, CAS, COL
(n.v.), GH, MO).
Freziera echinata is characterized by long, narrow leaf blades; long, erect,
persistent hairs on both leaf surfaces and on the stems; and erect setae sur-
rounded by erect hairs on the leaf margins. The flowers have densely long-
sericeous bracts, bracteoles, and sepals that are conspicuously pointed at the
apex. The bracteoles and outer sepals are sericeous inside, a condition unknown
elsewhere in the genus. The hairs are so dense that the floral parts cannot easily
be distinguished from each other. I have not seen a flower past anthesis, but
the most developed buds have extremely long, narrow corollas and petals. Some
floral characters are only partly known since the few flowers observed have all
been at least partially eaten wherever the sclereids in the tissues are not dense,
so the stamens, the base of the petals, and the ovary are usually gone.
This species, known only from the type collection, cannot be confused with
any other. No other taxon has this erect pubescence on the twigs and leaves,
or the extremely dense, long indumentum on the flowers. Freziera chrysophylla,
which has similarly shaped leaves, differs from F. echinata in having leaves
glabrous above and densely golden sericeous below, and pedicellate flowers
with round, sericeous bracteoles and sepals. Freziera tomentosa Ruiz & Pavon,
which like F. echinata has sessile flowers, is actually more similar to F. chry-
sophylla, with leaf blades glabrous above and densely sericeous below, but has
leaf blades wider than F. echinata or F. chrysophylla and rounded, glabrous
bracteoles and sepals.
1987] WEITZMAN, FREZIERA 231
Freziera minima A. Weitzman, sp. nov. FIGURE 3a-i.
A speciebus aliis Frezierae praeter F. stuebelii (Hieron.) A. Weitzman in
habitus fruticoso foliis minutis crenatis, et a F. stuebelii in foliis reticulato-
venosis differt.
Compact shrub | m tall; mature branches terete, brown, the bark conspic-
uously striate, splitting vertically; twigs square, slightly winged, persistently
brown-sericeous, the lenticels large, round, 1-1.7 mm in diameter on older
branches, splitting horizontally and vertically; terminal bud merely condupli-
cate, 1.5-4 mm long, short-sericeous. Leaves with petiole 0.9-3.1 mm long,
with narrow, involute wings, canaliculate, sericeous above and below; blade
broadly ovate, 7-12.1 by 4.9-10.6 mm, subcoriaceous, the base equal to sub-
equal, obtuse, truncate, round, or slightly cordate, the apex acute or obtuse,
ultimately retuse, terminating in caducous, thick, conical, red to black seta, the
margin crenate, with teeth 9 to 16 per side, and caducous, thick, conical, short,
black setae inserted in the sinuses, the surfaces glabrous, but with few caducous,
short, sericeous hairs on midrib above and below, the midrib flat to prominent
above, prominently rounded below, the lateral veins 5 to 7 per side, promi-
nently rounded above and below. Flowers solitary, subtended by 2 bractlike
structures, these basal on pedicel, persistent, narrowly ovate, 1.5-2.5 by 0.7-
0.8 mm, sclerotic, keeled, the base clasping, the apex acute, terminating in
thick, conical, black seta, the margin entire, with erect, thick, conical, black
setae, the outer surface sparsely short-sericeous; pedicel erect in bud and fruit,
recurved at anthesis, cylindrical, 2—2.9 by 0.7—1 mm, ridged, strigose, bracteoles
2, apical on pedicel, opposite, persistent, broadly ovate, unequal, 2.5-2.8 by
1.9-2.5 mm (larger), 2-2.1 by 1.6-1.8 mm (smaller), smaller one sometimes
keeled, the base rounded, the apex obtuse to rounded, on smaller bracteole
always and on larger one sometimes terminating in conical, black seta, the
margin membranaceous, with caducous cilia, the outer surface sparsely strigose-
glabrescent centrally. Flowers 6.7-7.7 by 3.4-4.1 mm; sepals 5, broadly ovate,
nearly equal, 3-3.6 by 2.5-3.3 mm (outer), 2.9-3.3 by 2.5-3 mm (inner),
sclerotic basally and chartaceous above, the base broadly cordate, the apex
rounded, the margin membranaceous, with caducous cilia and dark basal setae
(outer sepals) or pale basal flaps (inner sepals), the surfaces glabrous; corolla
urceolate, the petals 5, distinct or slightly connate basally, ovate, nearly equal,
5.3-6.4 by 2.1-3.4 mm, membranaceous in lower 4—'4, sclerotic above, the
apex obtuse, recurved at anthesis. Staminate flowers with stamens 18, unise-
riate, slightly adnate basally, unequal, unordered, filaments flat; long stamens
with the filaments linear, ca. 1.8 mm long, the anthers ovate, ca. 1.1 by 0.8
mm, the apiculus ovate, 0.1 mm long, apically rounded; short stamens with
the filaments linear or geniculate, 0.9-1.1 mm long, the anthers ovate, 0.7-1
mm long, basally cordate, the apiculus ovate, 0.1-0.2 mm long, apically round-
ed; gynoecium narrowly conical, the ovary 3-locular, ca. 1.1 by 1.5 mm, with
ovules ca. 7 per locule on 2 pendulous axile placentae, the style abruptly tapering
to linear, ca. 2.2 mm long, the stigmatic lobes 3, erect, ca. 0.25 mm long, the
stigmatic surface adaxial, dark, minutely papillate. Carpellate flowers with
aoe JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
wrens |S Hreziera
ELI LQ ee ae ,
SE | stuebel
FiGure 3. ao, Freziera minima: a, habit, from below; b, shoot apex; c, undersurface
of leaf; d, flower; e, petal of staminate flower, stamens adnate; f, gynoecium of staminate
flower; g, gynoecium and staminodia of carpellate flower: h, seed; 1, fruit (a—e from type,
g-1 from Holm- Nielsen et al. 3906). j, Freziera stuebelii, undersurface of leaf (from photo
of type).
staminodes 18, uniseriate, free (1 adnate to inner petal), linear, flat, equal, ca.
1.5 mm long, apically rounded: gynoecium conical, the ovary 3-locular, ca. 1.3
by 1.7 mm, with ovules 8 to 14 per locule, the style ca. 1.9 mm long, the
stigmatic lobes 3, slightly flaring, ca. 0.9 mm long, dark, conspicuously pa-
1987] WEITZMAN, FREZIERA 333
pillate. Fruits ovoid, 6.8-7.5 by 5.1-5.5 mm, with narrow dark band just below
stigmas, 2- or 3-locular, locules splitting out of fruit separately as mericarps;
seeds 2 to 8 per locule, irregularly rounded, ca. 1.6 mm long, dark brown.
Type. Ecuador, Prov. Loja, Zamora-Chinchipe border, crest of E cordillera,
ca. 13 km E of Loja, cloud forest and stunted crest vegetation, ca. 3°58’S,
79°10'W, 2840 m, 28 Jan. 1985, J. L. Luteyn & E. Cotton 11288 (bud, 4 fl—
holotype, Ny; isotypes, AAU, CAS, GH, MO, QCA (n.v.), U).
ADDITIONAL SPECIMEN EXAMINED. Ecuador, Prov. ZAMORA-CHINCHIPE, road Loja-Za-
mora, km 14, mountain ridges with elfin forest and open bogs, 4°S, 79°09’W, 2750-2770
m, 19-20 April 1973, Holm-WNielsen et al. 3906 (bud, @ fl, fr; AAU).
Freziera minima is characterized by minute, broadly ovate, crenate leaf
blades. In appearance the foliage does not resemble that of any previously
known Freziera. Other species with leaves nearly as small are F. microphylla
Sandw. (11.5-27.5 by 7.4-14.9 mm) and F. suberosa Tul. (10.7-18.6 by 7.5-
10.4 mm), both of which are densely sericeous on the twigs and leaf under-
surfaces and have revolute and therefore apparently entire leaf margins, quite
unlike F. minima. Other species such as F. euryoides Kobuski and F. parva
Kobuski, which have relatively small leaves and are sparsely pubescent (like
F. minima), have leaves two to five times longer than those of F. minima.
Freziera minima may also be closely related to the following species (see below).
Freziera stuebelii (Hieron.) A. Weitzman, comb. nov. FIGURE 3).
Taonabo stuebelii Hieron. Bot. Jahrb. Syst. 21: 320. 1896. Type: Colombia, Cerro
Patascoy, 3300 m, Stibel Colomb. 366 (holotype, B, destroyed: photos at GH, Mo,
negative at F (no. 9738)).
Patascoya stuebelii (Hieron.) Urban, Ber. Deutsch. Bot. Ges. 14: 283. 1896.
Ternstroemia stuebelii (Hieron.) Kobuski, J. Arnold Arbor. 23: 343. 1942, as steubelii,
nomen illegit
Freziera stuebelii was collected only by Stiibel at Cerro Patascoy, Colombia.
It is known only from a photograph of the holotype, which was destroyed at
Berlin (no isotypes are known). Urban (1896) mentioned the likely relationship
of Patascoya Urban to Freziera because they both have distichous leaves,
pubescence, and relatively few stamens. The photograph of the type suggests
similarity in habit, at least, to Freziera, and the leaves are similar to those of
F. minima, both having very broadly ovate, crenate blades about | cm long
and wide. Although no flowers or fruits are visible in the photograph, according
to the descriptions provided by Hieronymus (1896), Urban (1896), and Mel-
chior (1925), the flowers agree in all characters with those of Freziera.
When placing this species in its own genus, Urban (1896) heavily emphasized
the two bractlike structures at the base of the pedicel and the position of the
ovules. In both Freziera minima and F. stuebelii the solitary flowers are sub-
tended by two bractlike structures; these are morphologically similar to the
single bracts that subtend each flower in an inflorescence of other species of
Freziera. The fact that there are two such structures in these species is not
surprising since they are the equivalent of bud scales and since branches in
334 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Freziera usually are just an extension of the inflorescence axis. In describing
Taonabo stuebelii, Hieronymus (1896), following Szyszylowicz’s (1893) keys
to the Theaceae, placed the species in Taonabo Aublet (= Ternstroemia Mutis
ex L.f.) because that genus has pendulous ovules with apical placentation and
Freziera has exclusively axile placentation. I have observed pendulous placen-
tae and ovules in several species of Freziera. For example, in functionally
staminate flowers of F. minima, there are ca. seven ovules per locule that hang
from two pendulous axile placentae, which may appear apical.
Compared to Freziera minima, F. stuebelii has leaves that are broader and
more cordate at the base, with the lateral veins apparently sunken above, very
prominent below, and bifurcate instead of reticulate (compare FiGuRE 3c and
j). Bifurcate venation is unknown in other species of Freziera. All other
aspects of leaf morphology are apparently similar to those of F. minima and
are present elsewhere in the genus. In all species of Freziera most small veins
that go to the margin end in a seta, as they do in F. stuebelii, although in the
other species the veins are reticulate.
The relatively slight differences between Freziera and Patascoya discussed
above do not warrant generic status for Patascoya. Based on the evidence at
hand, I believe that F. stuebelii belongs to Freziera and is most closely related
to F. minima.
ACKNOWLEDGMENTS
I would especially like to thank P. F. Stevens and J. L. Luteyn for helpful
comments on this manuscript and throughout my work on this genus. Dis-
cussions with K. S. Bawa and C. Sobrevila enhanced the notes on reproductive
biology. I am grateful to the curators at A, AAU, CAS, F, GH, K, MO, MY, NY, U,
us, and vEN for the loan of specimens on which this study is based. Two
reviewers suggested improvements, for which I am grateful. I also want to
thank S. A. Spongberg and E. B. Schmidt for their help with the manuscript.
LITERATURE CITED
Bawa, K. S. 1980. Evolution of dioecy in flowering plants. Ann. Rev. Ecol. Syst. 11:
15-39
Hieronymus, G. 1896. Plantae stuebelianae novae. Bot. Jahrb. Syst. 21: 321.
Kosuskl, C. E. 1941. Studies in the Theaceae, VIII. A synopsis of the genus Freziera.
J. Arnold Arbor. 22: 457-496.
Lioyp, D. G. 1973. Sex ratios in sexually dimorphic Umbelliferae. Heredity 32:
Me tcuior, H. 1925. Theaceae. Jn: H. G. A. ENGLER & K. A. E. PRANTL, Nat. Pflan-
zenfam. ed. 2. 21: 109-154
Opter, P. A., & K. S. BAwaA. 1978. Sex ratios in tropical forest trees. Evolution 32:
812-821.
sete I. von. 1893. Theaceae. Jn: H. G. A. ENGLER & K. A. E. PRANTL, eds.,
at. Pflanzenfam. III. 6: 175-192.
oe I. 1896. Patascoya, eine neue Ternstroemiaceen-Gattung. Ber. Deutsch Bot.
Ges. 14: 282, 283.
1987] MEYER & HARDIN, AESCULUS FLAVA 335
STATUS OF THE NAME AESCULUS FLAVA SOLANDER
(HIPPOCASTANACEAE)
FREDERICK G. MEYER! AND JAMES W. HARDIN?
The name for the yellow buckeye, Aesculus octandra Marshall (1785), has
been in general use since publication of Robinson and Fernald’s seventh edition
of Gray’s New Manual of Botany (1908). Earlier, the name A. flava Aiton
(1789) had been in general use for this tree of the southern Appalachian Moun-
tains of the eastern United States. More recently, it has been shown that the
name 4A. flava Sol. was effectively published in 1778 and has priority as the
oldest valid name for this well-known tree.
The yellow buckeye of the eastern United States, a well-known tree and a
characteristic component of the mixed mesophytic forests of the southern Ap-
palachians (Hardin, 1957), has been known as either Aesculus flava Aiton or
A. octandra Marshall for nearly two centuries. Aiton’s A. flava (1789) was used
fairly consistently until publication of Robinson and Fernald’s seventh edition
of Gray’s New Manual of Botany (1908), when A. octandra Marshall (1785)
was accepted as the earlier valid name because of its priority of four years over
A. flava Aiton. Since that time, Marshall’s name has been universally accepted
for this North American tree as the earliest valid epithet (Hardin, 1957). More
recently, a note published in Bean’s Trees and Shrubs Hardy in the British Isles
(1970) explained that the name 4. flava, as pointed out by B. L. Burtt, of the
Royal Botanic Garden, Edinburgh, was originally published by Daniel Solander
in Catalogus Arborum et Fruticum in Horto Edinensi Crescentium (Anony-
mous, 1778). Solander’s name precedes A. octandra Marshall by seven years,
and Aiton’s A. flava by eleven. Our aim is both to include additional details
to confirm Burtt’s observations that 4. flava Sol. is the older and correct name,
and to alert botanists, foresters, and others who might easily have overlooked
the horticultural reference in Bean (1970).
With respect to the name Aesculus flava Aiton, it is well known that Aiton’s
Hortus Kewensis (1789) was in preparation for some twenty years; the text was
written largely by Jonas Dryander, who had succeeded Daniel Solander as
librarian for Sir Joseph Banks after Solander died in 1782. Also, Dryander is
known to have used manuscript material written earlier by Solander in pre-
paring the descriptions for Hortus Kewensis, but without reference to the source
of the information. Dryander may indeed have consulted Solander’s original
manuscript notes, but the wording in Aiton (1789, p. 494) on A. flava was
'U. S. National ee Agricultural Research Service, U.S. Department of Agriculture, Wash-
ington, D. C. 2
*Department of see North Carolina State University, Raleigh, North Carolina 27695-7612.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 335-341. July, 1987.
336 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
altered: Ae. foliolis quinis, corollae laminis cordato-subrotundis; unquibus calyce
duplo longioribus. (Compare FiGure | and the description of A. flava below.)
The catalogue with the name Aesculus flava Sol. published at Edinburgh
anonymously in 1778 was (fide Morton, 1986) issued under the direction of
Dr. John Hope (1725-1786), at that time Regius Keeper of the Edinburgh
Botanic Garden; however, no author’s or editor’s name appears on the title
page of that publication. In the Edinburgh Catalogus five new species in the
genera Aesculus L. (one), Andromeda L. (one), Cornus L. (two), and Crataegus
L. (one) can be clearly attributed to Solander as the publishing author (Burtt
in Morton, 1986). Of these, only 4esculus flava has been adequately typified.
Following a brief Latin diagnosis for each of the names, we find the identifying
letter ““S” standing for Solander, the publishing author, as explained at the end
of the catalogue.
n the recent edition of his Checklist of United States Trees (Native and
Naturalized) (1979), Elbert Little rejected Aesculus flava Sol. with the argument
(pers. comm. to Hardin, September, 1979) that the name had no publishing
author. On the other hand, /ndex Kewensis (Suppl. XV, 1974) listed the name
A, flava Sol. ex [Hope]. However, it is clear that the name flava was effectively
and validly published in 1778 in the Edinburgh Cata/ogus, which was published
anonymously, and that Solander, not Hope, was the author of the name. The
name A. flava Sol., listed by Bean (1970), Spongberg (1975), and Kartesz and
Kartesz (1980), is correct as cited, although the full bibliographic citation should
be A. flava Sol. in Anonymous, Cat. Arb. Frut. Horto Edin. Cresc. 1778, 3.
1778.
HISTORICAL BACKGROUND
Matters relating to the history and typification of the name Aesculus flava
are of concern because we have only Solander’s original manuscript notes and
the brief description published in the Edinburgh Catalogus. We are without a
clue as to the source of the material seen by Solander, except that it was
cultivated and growing in England. Specimens of wild material were not avail-
able to him. What was the source of the original North American material?
Was it really the tree we know as yellow buckeye, or another species? There
are no definitive answers for these questions. Original herbarium voucher spec-
imens of this tree from the wild collected in this period have not been located
and probably do not exist. We do know that in Solander’s time material of
this species was already growing in private gardens in England, and that it was
offered by at least one nursery in the London area in 1774.
Aiton (1789) reported that Aesculus flava was in cultivation by Mr. John
Greening (d. 1770) in 1764, the earliest recorded date of introduction, and that
it was from North Carolina. We also have evidence that yellow buckeye was
in cultivation in the Vineyard Nursery of Messrs. Lewis Kennedy and James
Lee at Hammersmith, London. In their Catalogue of Plants and Seeds, issued
in 1774, the third entry under Aesculus (p. 3) is “8 Flo. flavo, Yellow Horse
Chestnut.” This clearly confirms that Lee was indeed growing the yellow buck-
eye (4. flava) in his nursery in 1774. It is possible that Solander, a friend of
1987] MEYER & HARDIN, AESCULUS FLAVA ee
James Lee, saw flowering specimens of A. flava growing in Kennedy and Lee’s
Vineyard Nursery at Hammersmith.
James Lee (1715-1795), nurseryman, author, and correspondent, although
not a well-known figure in botanical circles, made noteworthy contributions
both to horticulture and to botany (Willson, 1961).
In his early years after coming to London, Lee was employed as a gardener
first at Syon, near Kew, and later, by the Duke of Argyle, at Whitton, near
Hounslow. About 1745 James Lee entered into a partnership with Lewis Ken-
nedy (1721-1782) in a nursery called ““The Vineyard” at Hammersmith, now
the site of Olympia, the great London exhibition hall. Lee devoted the re-
mainder of his life to his nursery and to introducing rare plants from different
parts of the world. In the preface to Hortus Kewensis, Aiton (1789) mentioned
that Lee had supplied a list of plants introduced by the Duke of Argyle at
Whitton. At that time the Vineyard Nursery maintained a collector in America,
one at the Cape of Good Hope, and another in South America (Loudon, 1838).
The genus Leea Royen ex L. of the Vitaceae was named in honor of James
Lee.
INTRODUCTION OF AESCULUS FLAVA FROM NORTH AMERICA
While we are unable to pinpoint the original source of Aesculus flava in
British gardens, we know that seeds of North American plants were regularly
being sent to England from about 1735 onward. John Bartram (1699-1777),
of Philadelphia, sent no less than 145 shipments of seeds and plants to cor-
respondents in England between 1735 and 1769 (Berkeley & Berkeley, 1982).
Bartram also sent many shipments of plants to his English Quaker friend, Peter
Collinson, of Mill Hill, near London, who in turn distributed much material
to his horticultural friends and to Daniel Solander, the botanist, for identifi-
cation (Earnest, 1940; Darlington, 1967).
John Bartram (Earnest, 1940; Darlington, 1967; Berkeley & Berkeley, 1982)
visited Pittsburgh in the fall of 1761 and met Col. Henry Bouquet from Ohio,
receiving from him plant material from the Ohio River valley. Yellow buckeye
(Aesculus flava) and Ohio buckeye (4. glabra Willd.) could have been included
in this material. In the fall of 1762, Bartram was on an extended trip to the
interior of South Carolina (Wateree and Congaree rivers), to western North
Carolina, and to southwestern Virginia (Yadkin and New rivers, Natural Bridge,
Luray Caverns, Staunton, and the Shenandoah Valley). On this excursion he
collected fruits of three different “horse chestnuts” from southwestern Virginia
that were later identified by Solander as 4. hippocastanum L., A. pavia L., and
A. media, the last “not taken notice of by Dr. Linnaeus” (Berkeley & Berkeley,
1982, p. 349). The one called 4. hippocastanum was undoubtedly A. glabra,
which is interesting because 4. glabra 1s unknown in that area today, so far as
we know. Those called A. pavia and A. media, initially identified without
flowers, were probably variations of 4. flava, which could have been named
by Solander after flowering material was available in England.
Another possible source of yellow-buckeye material from the North Carolina
mountains was W. V. Turner, an Indian agent who sent plants to Sir Joseph
Banks (Joseph Ewan, pers. comm. to F. G. Meyer, September, 1980).
68
338 JOURNAL OF THE ARNOLD ARBORETUM [VOL.
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Figure |. Solander’s original manuscript ae: describing Aesculus flava.
TYPIFICATION OF AESCULUS FLAVA
A copy of Solander’s original manuscript slips (nos. 339-341) with a de-
scription of Aesculus flava was kindly supplied by the librarian of the Depa
rt-
ment of Botany at the British Museum (Natural History). Thus, for purposes
of typification, it has been possible to use Solander’s original description and
to publish more than 200 years later his complete manuscript notes for t
first time (see FiGurE 1). This is important, since the brief protologus of
he
A,
flava published in the Edinburgh Catalogus included only the first four lines
Ficure 2. Neotype of Aesculus flava Sol.
from Solander’s original manuscript (our translation of Solander’s protologue:
““calyx ovate, half the length of the upper petal claws, blade cordate-subrotund,
stamens curved”) and therefore was incomplete and is inadequate for typifi-
cation. These details allow a positive identification only of a buckeye—A. flava
or A. sylvatica Bartram. In his complete text, however, Solander describes the
yellow flowers, the relative length of calyx and upper petal claws, the extremely
dimorphic petals, and the included stamens of A. flava/sylvatica, details that
are adequate for typification. In addition, these characters definitely eliminate
A. pavia and A. glabra from consideration. Aesculus glabra was described by
Willdenow in 1809. The differences between A. flava and A. sylvatica are mainly
in habit (tree vs. shrub) and in rather subtle features of pubescence and size of
floral parts (Hardin, 1957).
Marshall’s (1785) description of Aesculus octandra was only slightly more
diagnostic, for he did indicate that it was a tree. His common name “New
River horse chestnut” would most likely have come from John Bartram (E.
Berkeley, pers. comm. to J. Hardin, February, 1982), in reference to the material
brought back from his trip of 1762 to the New River in southwestern Virginia.
Unfortunately, there is no specimen of Aesculus flava that was collected or
annotated by Solander. The earliest possibly appropriate material in the
British Museum (Natural History) was collected by James Lee at the Vineyard
Nursery of Messrs. Kennedy and Lee and has the number “74” (interpreted
as 1774) on the herbarium label. This specimen (see FiGurE 2), although
somewhat damaged after more than two centuries, contains several leaves and
a short portion of a poorly preserved inflorescence with a few flowers. We
consider it to be authentic 4. flava Sol. The leaflets are somewhat narrower
340 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
than normal, but well within the range of variation of the species. There are a
few poorly preserved stipitate glands at the base of the calyx, which definitely
identify the specimen as A. flava rather than A. sy/vatica. There is no evidence
of A. sylvatica in England prior to William Bartram’s discovery and description
of 1791. Bean (1970) indicates its cultivation (probably as 4. neglecta Lindley)
in Europe in 1826.
We hereby designate the James Lee specimen (BM), the earliest known doc-
umented material, as the neotype of Aesculus flava Sol.
DESCRIPTION OF AESCULUS FLAVA SOLANDER
Aesculus flava Sol. in Anonymous, Cat. Arb. Frut. Horto Edin. Cresc. 1778,
3. 1778. NEOTYPE: ex hort. Lee, [17]74 (BM).
Aesculus octandra Marshall, Arbust. Am. 4. 1785. Type: not s
Aesculus lutea Wangenh. Schriften Ges. Naturf. Freunde Berlin 8: 135. 1788. Type:
ot seen.
Additional synonymy is given in Hardin (1957).
Translation of Solander’s holographic description of Aesculus flava (FiGURE 1):
flava AESCULUS leaflets five; calyx ovate, half the length of the upper petal
claws; blade cordate-subrotund; stamens incurve
Raceme lax, subpendulous. Rachis and peduncles green.
Calyx [turning] from green to dull yellowish, ovate, open, half the
length of the upper petal claws.
Corolla pale sulphur yellow. Blade subrotund, subcordate, undulate:
the two inner ones inclined upward.
Filaments generally seven, subulate, apices inclined upward, shorter
than the petals, unequal, turning yellow
Style yellow, subulate (in flowers seen, little shorter than the stamens,
with those inclined).
Petioles green.
Leaves flat
Observation. Flowers without copious secretion.
ACKNOWLEDGMENTS
We are indebted to the Keeper and the Librarian of the Department of
Botany, British Museum (Natural History) for providing critical materials,
including a historic specimen and a copy of Daniel Solander’s manuscript notes,
which together were invaluable in the typification of Aesculus flava Sol. We
also wish to thank Joseph Ewan and Edmund Berkeley for their interest and
diligence in helping to solidify some of the historical aspects of our paper.
Finally, we would like to thank B. L. Burtt, T. R. Dudley, Elizabeth McClintock,
R. C. Rollins, E. E. Terrell, and R. M. Tryon for their kindness in critically
reading Our manuscript.
1987] MEYER & HARDIN, AESCULUS FLAVA 341
LITERATURE CITED
Arron, W. 1789. Hortus Kewensis. 3 vols. George Nicol, London. (A. flava cited, Vol.
9
ANoNyMous. 1778. Catalogus arborum et fruticum in Horto Edinensi crescentium,
anno 1778. Balfour and Smellie, Edinburgh. (Aesculus flava Sol., p. 3.
BEAN, W. J. 1970. Trees and shrubs hardy in the British Isles. ed. 8. (SIR GEORGE
TayLor, general ed.) 5 vols. J. Murray, pee . (Footnote with details from the
original description of Aesculus flava Sol., Vol. 1,
BERKELEY, E., & D. S. BERKELEY. 1982. The life and travels of John Bartram, from
Lake Ontario to the River St. John. University Presses of Florida, Gainesville.
(Details on John Bartram’s travels and collections, pp. 201, 210, 311-318, 349 (note
)
Burtt, B. L. 1986. The garden catalogues of 1775 and 1778. P. 43 in A. G. Morton,
John Hope, 1725-1786, Scottish botanist. Edinburgh Botanic Garden (Sibbald)
Trust, Edinburgh.
Pee W. 1967. Memorials of John Bartram and Humphry Marshall. (Intro-
duction by J. Ewan; facsimile of 1849 a Hafner Publ. Co., New York. (Letters
Soni Cie to J. Bartram, pp. 229, 2
Earnest, E. 1940. John and William eee once Pennsylvania Press, Philadelphia.
Caer of Bartram’s travels, pp. 57, 58.)
Haron, J. W. 1957. A revision of the American Hippocastanaceae. Brittonia 9: 173-
195, (piscicsion of A. flava, pp. 189-191.)
Index Kewensis. 1974. Suppl. XV. (A. flava Sol. ex [Hope], p. 4.)
Kartesz, J. T., & R. KARTEsz. 1980. A synonymized theta of the vascular flora of
the United States, Canada, and Greenland. Vol. 2, The biota of North America.
Univ. North Carolina Press, Chapel Hill. (A. flava Sol., with synonymy by Hardin,
p. 249.)
KENNEDY, L., & J. Lee. 1774. Catalogue of plants and seeds, sold by Kennedy and Lee,
ae and seedsmen at the Vineyard, Hammersmith. London. (Yellow horse chest-
nut,
LITTLE, E cL. Jr. 1979. Checklist of United States trees (native and naturalized).
Agriculture Handbook No. 541, Forest Service, United States Department of Ag-
riculture, Washington, D. C. (4. octandra and synonymy, p. 46.)
Loupon, J.C. 1838. Arboretum et fruticetum Britannicum. 8 vols. London. (Account
MarsHALL, H. 1967. Arbustum Americanum. (Introduction by J. Ewan; facsimile of
1785 ed.) Hafner Publ. Co., New York. (4. octandra, pp. 4, 5.)
Rosinson, B. L., & M. L. oe 1908. Gray’s new manual of botany. ed. 7. New
York. (A. octandra, p.
See S.A. 1975. Changes of botanical names. Pl. & Gard. 30(4): 45-47. (Name
change of A. flava Sol., p. 46.)
WILLSON, °E J. 1961. aes Lee and the ee Nursery, Hammersmith. Ham-
mersmith Local History Group, London
SANDERS, LANTANA 343
A NEW SPECIES OF LANTANA (VERBENACEAE) FROM
DOMINICA, LESSER ANTILLES
ROGER W. SANDERS!
Lantana hodgei Sanders is described from Dominica and is contrasted with
L. camara L. and L. urticifolia Miller on the basis of gross morphology, scan-
ning electron microscopy of laminar surfaces, and pollen stainability.
Studies of Lantana L. (Verbenaceae) for the Flora of the Lesser Antilles,
edited by Richard A. Howard, reveal the existence of an undescribed species
from the montane forests of the island of Dominica.
Lantana hodgei R. Sanders, sp. nov. FIGURE 1.
Differt a Lantana camara L. habitu subscandenti, trichomatibus caulium
foliorumque brevioribus sparsioribus validius appressis, petiolis longioribus,
laminis angustioribus circa duplo longioribus quam latioribus supra subtusque
nitentibus subtus subviridi-griseis, nervis secundariis nervellisque laminarum
subtus non elevatis; a L. urticifolia Miller trichomatibus laminarum remotis
non nisi nervis mediis secundariis tertiariisque insidentibus angustate conicis
antrorse geniculatis.
Subscandent shrub; main branches 2-3 m long, usually few, weak, trailing
or sprawling, usually without prickles, often scabrous with scattered appressed
hairs 0.2-0.6 mm long. Leaves with petiole 1-2.5 cm long; lamina ovate to
elliptic-lanceolate, 5-13 cm long, usually 1.7—2.5 times longer than wide, non-
rugose, the higher-order and often the secondary veins not impressed above
or keeled below, the apex usually abruptly acuminate, the base attenuate to
shortly attenuate, the margin serrate-dentate, with teeth 20 to 40 per side, 1-
2 mm long, 1-3 times longer than wide, the adaxial surface dark green, lustrous,
thinly strigillose, with hairs very sparse, restricted to midrib, secondary veins,
and center of major areoles (1 hair per areole), to 0.4 mm long (0.8 mm on
veins), often deciduous, the abaxial surface gray-green, lustrous, nearly gla-
brous, with hairs very sparse, restricted to midrib and secondary and tertiary
veins, tapering-conical, 0.1-0.5 mm long, geniculate toward base, antrorse,
strongly appressed, weak, often deciduous. Inflorescences capituliform spikes
in axils of distal leaves; peduncle 2-3 cm long; receptacle fistulose; bracts
(excluding single outer series) narrowly lanceolate, ca. 5 mm long, widest near
proximal third, deciduous in fruit, abaxially sparsely hirsute, hairs strongly
appressed. Calyx ca. 2 mm long, 2- or 3-toothed; corolla salverform, bilaterally
‘Fairchild Tropical Garden, 11935 Old Cutler Road, Miami, Florida 33156.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 343-348. July, 1987.
344 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
4-lobed, tube 5-8 mm long (when dried), limb ca. 6 by 4-5 mm, orange to red
(sometimes dull pink, according to note on 4. C. Smith 10216). Drupes 4-5
mm across, black; pyrenes obovoid, 3-4 by 3 mm, bilocular, inflated, basally
acute, distal ornamentation semicircular, shallow, oblique, not trilobed.
Type. Dominica, near Fresh Water Lake, common along road, steep slopes in
“elfin forest,” 10 March 1967, F. R. Fosberg 48269 (holotype, us!; isotypes,
F!, GH!, K [fide C. H. Stirton], Mo!, Ny!).
DISTRIBUTION AND ECOLOGY. Known only from Dominica on sunny slopes in
borders and openings of montane rainforest, 450-900 m alt. Flowering and
fruiting January to August, possibly year-round.
ADDITIONAL SPECIMENS EXAMINED. Dominica: S slope of Morne Macaque on road to
Fresh Water Lake, Ernst 1728 (us); between Laudat and Fresh Water Lake, Hodge &
Hodge 1808 (us), A. C. Smith 10216 (A, Ny, Uc, Us); Laudat, Lloyd 201 (ny), Nicolson
2102 (FTG); Springfield, Krauss 1268 (LL); Sylvania, Morne Colla Anglais, Cooper 5 (Fr,
GH, NY, US), Hodge 861] (Gu), 1038 (Gu), 1115 (GH).
EpitHET. The epithet honors Walter H. Hodge, whose extensive collections
have helped to elucidate the nature of this species.
Two other species of Lantana sect. Camara Cham., L. camara L. and L.
urticifolia Miller, occur in Dominica and the Lesser Antilles and could be
confused with L. hodgei. The three taxa are contrasted in the following key:
1. Hairs of abaxial leaf surface ep aISE, restricted mostly to midrib and secondary and
tertiary veins, tapering-conical, geniculate toward base with distal %4 parallel to lamina
or vein surface
Laminas si .6 times longer than wide; base usually truncate or cordate; adaxial
surface at maturity scabrous or strigose, more or less dull, moderate green, the
hairs scattered over entire surface, stout, usually aoe (at least the conical
bases); abaxial surface lighter yellow-green, thinly strigose on veins, the hairs
scattered to moderately abundant, stout, antrorse but with tip a above surface,
the secondary and higher-order veins keeled. ntana camara.
. Laminasca. |.7—2.5 times longer than wide; base usually oe hoc Pens
at maturity lustrous, dark green, smooth, the hairs restricted to and | 1
center of each areole, small, weak, often deciduous; abaxial ee on ee
green, almost glabrous, the hairs very sparse, weak, strongly a so higher-
order and usually secondary veins not keeled. ............... Lantana hodgei.
. Hairs of abaxial leaf surface usually abundant and crowded, at least pee crevice
between major veins and laminar surface, usually occurring on all veins including
areolar veinlets and often on noninnervated laminar tissue, filiform (or also gland
upped), straight or gently curved from basal insertion, spreading from vein surface
or erect on laminar surface. ......00.00.0 00.00. e eee ee Lantana urticifolia.
iS
ss
in)
Lantana hodgei is probably closely related to L. camara because both species
have tapering, geniculate hairs on the abaxial leaf surfaces (FiGuURE 2b, d).
Lantana camara is commonly encountered in both native and apparently
naturalized populations throughout the West Indies and northern South Amer-
ica and is a morphologically variable species. Thus, L. hodgei has been con-
sidered conspecific with L. camara in past studies (Moldenke, 1980 and in
1987] SANDERS, LANTANA 345
Wom. cuRTIN &
Fic Lantana hodgei, habit, inflorescence, and variation in leaf size and shape:
a, b, pie roe (holotype, ia C, een erg 48269 (isotype, F); d, Hodge 1115 (Gu).
Metric scales numbered in centimet
346 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
<< Sie er
Weer
—2. Scanning electron micrographs of adaxial : c, e) and abaxial (b, d, f) leaf
ene of Lantana species occurring e Dominica: a, b, L. hodgei ee 2102, FTG),
white arrows pointing to isolated hairs; c, d, L. camara (Wilbur et al. 7665, FTG); e, f,
L. colic (Hodge 859, Gu), ea basal bending of hairs primarily an artifact of
foreshortening. Largest veins shown secondary. Scale bar = 0.5 mm.
schedula). Indeed, some workers would probably submerge all taxa of Lantana
sect. Camara under L. camara, as Gibson (1970) did for the Flora of Guate-
mala. Nicolson (unpubl. ms.) calls for new approaches to augment morphology
in the delineation of lantanas in the West Indies. Extrapolating from a limited
sample of biosystematic and chromosomal studies (Sanders, 1987a, c), I believe
there 1s sufficient reason for separating L. camara from L. hodgei and other
species that have gone under the name L. camara. The structure of the abaxial
1987] SANDERS, LANTANA 347
Pollen stainability of Lantana hodgei, L. urticifolia, and their intermediates.*”
%
TAXON SPECIMEN Ne STAINABILITY
L. hodgei Cooper 5 (F) 300 85
Fosberg 48269 (us) 300 65
Hodge 1115 (c 190 81
Hodge & Hodge 1808 (Gu)4 200 3
Nicolson 2102 (FTG) 310 53
A.C. Smith 10216 (a)? 200 4
Intermediate Hodge 858 (Gu) 203 27
Hodge 860 (GH) 291 35
odge & Hodge 2592 (GH) 300 33
Shillingford 120 (Mo) 200 29
L. urticifolia Dey 69 Ae — 303 58
Hodge 858 (Ny 200 42
Hodge ae (G _ 200 35
Howard 15236 (A) (Redonda) 226 74
Lloyd NY 300 37
Stoffers 3004 (a) (Saba) 200 81
Pollen from nearly open or open corollas removed from herbarium specimens and stained in
lactophenol cotton-blue
*Collections from Bominice unless indicated otherwise.
‘Total number of pollen grains c d.
“Flowers blackened with drying aad) or infested with insect larvae.
laminar hairs divides Lantana sect. Camara into two sets of taxa—a “camara-
cohort,” with conical, geniculate hairs (FiGuRE 2b, d), and an “‘wrticifolia-
cohort * with slender, spreading hairs (Figure 2f), Each set includes one or
more morphologically distinctive, endemic, and often diploid taxa, in addition
to the more morphologically generalized (and hence overall “‘camara-like’’),
widespread, tetraploid ones (Sanders, 1986, 1987a—c). Characters with gener-
alized states in both groups of tetraploids include growth habit, leaf shape and
size, hairs of adaxial leaf surfaces (FIGURE 2c, e), bract shape and size, and
flower size. Although the chromosome number of L. hodgei is unknown, in
other characters this species exceeds the limits of variation of L. camara as
much as do the other distinctive endemics of the “camara-cohort.”
Lantana camara, as delimited here (including L. aculeata L.), is apparently
infrequent on Dominica (Dominica, | km NW of Salisbury, Wilbur et al. 7665
(F, FTG, LL, MO, US—7.V.)).
Lantana urticifolia (including L. arida Britton and L. moritziana Otto &
Dietr.) is a widespread and variable species, ranging from Mexico and Cuba
to Brazil. It is commonly encountered in Dominica in low-elevation scrub and
man-made openings on the lower slopes (Dominica: without further locality,
Imray 229 (Gu); Belle View, Hodge 857 (Gu); Fern Villa, Hodge & Hodge 2177
(Gu); Marigot, Mantipo R., Hodge 858, p.p. (NY, US); Roseau, Hodge 859 (Gu),
Lloyd 929 (Ny); between Salybia and Hatton Garden, Hodge 3201 (Gu)).
Where human disturbance has allowed Lantana hodgei and L. urticifolia to
come in contact, a spectrum of morphological intermediates between the two
348 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
are found (Dominica: Belfast, Shillingford 120 (mo); between Belle View and
Grand Bay, Hodge 860 (Gu); Marigot, Mantipo R., Hodge 858, p.p. (GH); Milton
Estate, Hodge & Hodge 2592 (Gu)). Presumably these intermediates are hybrids,
like those documented in Florida (Sanders, 1987a). The laminas of these plants
are more nearly rounded to cordate at the base, adaxially sublustrous, and
abaxially with a moderately dense mixture of filiform straight hairs and tapering
geniculately antrorse hairs. The percent stainable pollen is low compared to
that of either L. hodgei or L. urticifolia (TABLE). Indeed, the lower stainability
of pollen of L. urticifolia from Dominica compared to that from other islands
in the Lesser Antilles may suggest that Dominican L. urticifolia has undergone
widespread introgression from L. hodgei. Note especially the apparent co-
occurrence of L. urticifolia and intermediates (e.g., Hodge 858, cited above)
on that island.
ACKNOWLEDGMENTS
I am grateful to the curators of A, F, FTG, GH, LL, MO, NY, UC, and us for
loans of specimens. I used the scanning electron microscope at Florida Inter-
national University, and I am indebted to J. H. Richards, who instructed me
in its operation. I thank D. H. Nicolson for making available unmounted
specimens of Lantana from Dominica and for providing a typescript copy of
his treatment of Lantana from his unpublished Flora of Dominica. He and
C. H. Stirton reviewed drafts of the manuscript.
LITERATURE CITED
Gipson, D. N. 1970. Verbenaceae. /n: P. C. STANDLEY & L. O. WILLIAMS, Flora of
Guatemala. Fieldiana, Bot. 24(1X): 167-236.
Howarp, R. A., ed. 1974-1979. Flora of the Lesser Antilles. Vols. 1-3. Arnold Ar-
boretum of Harvard University, Jamaica Plain, Massachusetts.
Mo .penkg, H. N. 1980. sixth summary of the Verbenaceae, Avicenniaceae, Stil-
baceae, Chloanthaceae, Symphoremaceae, Nyctanthaceae, and Eriocaulaceae of the
world as to valid taxa, geographic distribution and synonymy. Phytologia Mem. 2.
Nicotson, D. H. Lantana. In: Flora of Dominica. Unpublished manuscript.
SANDERS, R. W. 1986. Biogeographic connections between Mesoamerica and the West
Indies in the distribution of Lantana (Verbenaceae) species. Jn: L. D. GOmez, Pro-
ceedings of the symposium on the biogeography of a Editorial Uni-
gee Estatal a Distancia, San José, Costa Rica. In
87a. Identity of Lantana depressa and L. vatifolia ree of Florida
and me Bahamas. Syst. Bot. 12: 44-60.
987b. Lantana sect. Camara in Hispaniola: novelties and notes. Moscosoa
5: 1n press.
1987c. Taxonomic significance of chromosome aca in Caribbean
species of Lantana (Verbenaceae). Amer. J. Bot. 74: in p
NOTE ADDED IN PROOF. A specimen documenting the occurrence of Lantana
hodgei outside of Dominica (Martinique, beyond L’Alena, Bailey & Bailey 240
(NyY)) has recently come to my attention.—R. W. S.
1987] RANJANI & KRISHNAMURTHY, MIMOSOIDEAE 349
A COMPARATIVE STUDY OF ROOT AND STEM
WOODS OF SOME MEMBERS OF THE
MIMOSOIDEAE (LEGUMINOSAE)
K. RANJANI AND K. V. KRISHNAMURTHY'!
A comparative study of the root and stem woods of 11 members of the
Mimosoideae revealed that the two woods were more alike than had been
thought. The only feature of consistent difference was the presence of a greater
amount of thinner-walled elements in root wood than in stem woo
Although structural variation in stem wood has been studied in several
arborescent plants, so far less attention has been paid to root wood (Fayle,
1968). This has mainly been due to the assumptions that the structure of root
wood is similar to that of stem wood and that root wood has only slight
economic importance. It has also been due to the difficulties in procuring
authentic root-wood samples (Cutler, 1976). We therefore undertook this com-
parative study on root and stem woods. We chose subfamily Mimosoideae for
investigation not only because of the easy availability of specimens but also
because of the lack of study on its root wood.
MATERIALS AND METHODS
Eleven species of Mimosoideae were selected for the study: Acacia arabica,
Acacia auriculiformis, Acacia leucophloea, Adenanthera pavonina, Albizzia
amara, Albizzia lebbeck, Dichrostachys cinerea, Enterolobium saman, Leu-
caena leucocephala, Pithecellobium dulce, and Prosopis spicigera. Wood sam-
ples were collected at chest height from the main stem and from the strong,
laterally spreading roots at 0.5-1 m below soil level. The collected samples
were trimmed to | cm*in such a way as to include both heartwood and sapwood,
and as many growth rings (if present) as possible. Transverse, radial-longitu-
dinal, and tangential-longitudinal sections were taken using a Bright cryostat
microtome at a thickness ranging from 15 to 30 um. The wood was pretreated
in boiling water, 10 percent hydrofluoric acid, or a glycerine-alcohol mixture
singly or in combination if there was difficulty in sectioning the wood. Sections
were stained with safranin alone or with safranin and Delafield’s haematoxylin.
In addition, macerations of the wood were prepared using Jeffrey’s fluid (Jo-
hansen, 1940); the macerated elements were also stained with safranin. For all
features recorded, 100 random measurements were made. Sample size was
accounted for using Student’s t test, and levels of significance were calculated
‘Department of Botany, Bharathidasan University, Tiruchirapalli 620 023, Tamil Nadu, India.
© President and Fellows of Harvard College, 1987.
ie of the Arnold Arboretum 68: 349-355. a 1987.
Comparison of root and stem woods of the taxa of Mimosoideae investigated.
CHARACTER*
SPECIES
1 2 S&S & S&S & F 8 9 10 11 12 13 14 15 16 17,18 19
Acacia arabica R A Dp 15 145 315 20 Ac, Cp 22 Ho 370 35 20 10 L, Sp, St 1430 (-) 680 48
Mahe D Dp 10 145 300 20 V, Ac, R, Cp 23 Ho 340 35 30 10 L, Sp, St 1200 (-) 690 47
CIP)
Acacia auriculi- R A Dp 20 75 270 4 Ac, Cp 25 Ho 125 15 70 4 L, Sp 825. (-) 4 67
formis A. Cunn. 5 9 pp 20 110 280 12 Ac, R, Cp 12 Ho 130 20 65 BL 30. a 68
ex Benthar (CF. IP)
=
Acacia leuco- A Dp 10 270 450 28 Ac, Cp 47 Ho 365 35 25 10 1655 s 15
piloee: Naa S A Dp 10 130 290 11 Ac, Cp 33 Ho 330 30 45 9 L 1310 - 47
Adenanthera R A Dp 10 180 780 10 V, Ac, Cp 30 Ho 395 30 30 13 L, St 1255 - (-) 47
pavonine S D Dp 10 130 510 9 V, Ac, Cp 26 Ho 305 30 40 15 L, St 1450 - 645 50
(cr)
Albizzia amara R D Dp 6 165 330 7 Ac, R, Cp 22 Ho 180 15 35 4 L, Sp 1070 1140 (-) 67
Boivin (CF) _
D Dp 6 140 300 10 Ac, Cp 16 Ho 230 15 60 12 L, Sp, St 1130 1150 670 62
(CF) —
Albizzia lebbeck R A Dp 5 95 210 2 Ac, Ap, Cp 7 Ho 260 70 25 23 L, Sp, St 705 1075 520 68
Bentham S A Dp 5 110 240 7 V, Ac, Cp 15 Ho 250 50 45 19 L, Sp, St 1085 1075 (-) 59
Dichrostachys D Dp 20 110 210 5 V, Ac, R, Cp 8 Ho 160 30 80 SL 955. - é 82
cinerea Wight & (CF)
D Dp 20 100 230 8 V, Ac, R, Cp 9 Ho 220 30 50 17 L 960 - s 76
(CF)
Enterolobium I Dp 10 130 240 16 Ac, Cp 20 Ho 190 30140 12 L, Sp, St 900 = ((-) 595 52
saman (Jacq. ) (IP)
Ean S I Dp 10 125 335 10 Ac, Cp 10 Ho 140 15 65 15 L 805 7 7 65
(CF )
WOLAYOPIUV GTONUV AHL JO TYNUNOL OSe
89 “10A]
Leucaena leuco- R D Dp 10 100 330 12 Ac, Cp 21 Ho 260 35 40 7 L, Sp, St 1105 950 (-)
cephala (Lam. ) (CF)
Deane D Dp 10 160 230 15 Ac, R, Cp 23 Ho 295 20 30 6 L 840i} =
(CF ,IP)
Pithecellobium BN pal, MiDpe 10, G0 4a20e <7 Vy Aa, toe 322) so 270-408 30 43~ Uetsp 1180 1270 a
dulce Bentham (IP)
D 12 140 435 17 Ac, Cp 11 Ho 210 25 60 8 |L, Sp, St 1090 935 (=
(CF, IP)
Prosopis R A Dp 14 130 295 13 Ac, Cp 20 Ho 295 20 63 12 L, Sp Fos: ie) 7
a a S A Dp 13 130 225 15 Ac, Cp 30 Ho 315 30 68 10 L, Sp 905 (-) =
*Key
pons
ANAWUEW
to characters:
Portion of plant where wood samples taken: R = root, S = stem.
Growth rings: A = absent, D = distinct, I = indistinct, CF = marked by compressed late-wood fibers, IP = marked by
initial parenchyma:
Porosity: Dp = diffuse porous
Mean number of vessels per mm? in transection.
um).
Percentage of area of transection occupied by vessels.
Nature of parenchyma: Ac = aliform confluent, ap = apotracheal diffuse, Cp = compartmented crystal, R = restricted to
side facing ae of woo = vasi icentri
Percentage of are ae transection occupied by Pieper
Nature of rays: Ho = homogeneous.
Mean height of rays in tangential-longitudinal section aN
Mean width of rays in BaQOe gb Tel enone tal Ae (um
ts
y Lays.
Type of fibers: L = libriform, Sp = septate, St = substitute (predominant type underlined).
Mean length of libriform fibers (um).
Mean length of septate fibers (um); - = absent, (-) = data unavailable due to rarity of fibers.
s (um).
Percentage of area of transection occupied by fibers.
AVACIOSOWIN ‘AHLYNNVNHSIOM ¥ INVINVa [L861
IS¢
352 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
for P = 0.01 and 0.05. Microphotographs were taken with a Nikon Labophot
microscope. Terminology is in accordance with the IAWA Multilingual Glos-
sary (International Association of Wood Anatomists, 1964).
OBSERVATIONS AND DISCUSSION
The TABLE provides the data on all qualitative and quantitative features of
the root and stem woods.
GROWTH RINGS
Although variability in growth rings has been studied in detail (Carlquist,
1980), the degree of expression of the ring within the stem and root woods of
the same plant has not yet been adequately investigated. Fayle’s (1968) state-
ment that growth-ring boundaries are better marked in the stem than in the
root 1s supported by Cutler (1976), Fahn (1982), and Zimmermann and Brown
(1971). This is the case in four of the eleven species we studied (Acacia arabica,
Acacia auriculiformis, Adenanthera pavonina, and Pithecellobium dulce) but
not for Albizzia amara, Dichrostachys cinerea, Enterolobium saman, or Leu-
caena leucocephala, growth rings were absent in the other three species inves-
tigated (Acacia leucophloea, Albizzia lebbeck, and Prosopis spicigera). The pres-
ence of growth rings and the degree of their distinction have been reported to
be highly variable even in the stem woods of the Mimosoideae (Ramesh Rao
& Purkayastha, 1972). In other words, the degree of distinction shown by
growth rings may not be directly related to the organ in which the growth ring
is present. The reason for this variability is difficult to explain since several
intrinsic and extrinsic factors (such as hormone levels, availability of carbo-
hydrates, climatic factors, and soil moisture) appear to control the expression
of growth rings.
It is generally believed that the feature or features marking the growth ring
are specific for each plant, irrespective of the organ (see Carlquist, 1980).
Although this was true of A/bizzia amara and Dichrostachys cinerea, where
compressed late-wood fibers marked the growth ring in both stem and root
woods, it was not true of other taxa, in which the growth rings of stem and
root woods were marked by quite different features (see TABLE).
VESSEL AND VESSEL ELEMENTS
Root wood has been reported to have a greater abundance of vessels and
vessel multiples per unit area than stem wood (Carlquist, 1978; Carlquist et
al., 1983, Gdmez- Vazquez & Engleman, 1983). Fayle’s (1968) results, however,
did not agree with this (see also Zimmermann & Brown, 1971). Cutler (1976),
in discussing the subject, cautioned that further research was necessary before
specific conclusions could be drawn. He made this statement because in his
study of Acer stem and root woods, he found certain samples of root wood to
have more abundant vessels than stem wood, while one sample showed no
difference in quantity. In nine of the 11 taxa we investigated, pore abundance
1987] RANJANI & KRISHNAMURTHY, MIMOSOIDEAE $55
was the same in both root and stem woods. Only in Acacia arabica and Pithe-
cellobium dulce was there a difference at the 1 percent level of significance; in
the former abundance was greater in the root wood, while in the latter the
contrary was true.
PorE DIAMETER
Presence of wider pores in root wood has been considered to be the most
consistent distinction between root and stem woods (Bhat, 1982; Carlquist,
1975, 1977, 1978; Chalk, 1983; Fahn, 1982; Fayle, 1968; G6mez-Vazquez &
Engleman, 1983; Plank, 1976; Zimmermann & Brown, 1971; Zimmermann &
Potter, 1982). Cutler (1976) was cautious enough to state that further research
into this matter was warranted in view of the number of exceptions to the
above observation. In the individuals we studied there was no significant dif-
ference even at the 5 percent level in mean pore diameter of stem and root
woods of Acacia arabica, Enterolobium saman, or Prosopis spicigera. The
difference was significant at both levels in the rest of the species, with greater
diameter being exhibited by the stem-wood vessel elements in Acacia auri-
culiformis, Albizzia lebbeck, Leucaena leucocephala, and Pithecellobium dulce
and by the root-wood vessel elements of the other four species. Thus, mean
pore diameter does not appear to be a feature of consistent difference between
root and stem woods.
VESSEL-ELEMENT LENGTH
Whether the length of vessel elements depends upon the organ 1s a question
often debated in the literature. Carlquist (1976) believed that the elements were
longer in root wood than in stem wood. This opinion was also held by Fayle
(1968), Plank (1976), and Zimmermann and Potter (1982). The data obtained
in the present study revealed that longer vessel elements were present in the
root wood of Acacia leucophloea, Leucaena leucocephala, sees FrOSODES spici-
gera, but in the stem wood of Adenanthera pavonina and Enter um saman
In all of the above, the difference in length was seers at the | percent
level. In Albizzia lebbeck the stem wood had longer elements, but the difference
was significant only at the 5 percent level. In Acacia arabica, Acacia auricu-
liformis, Albizzia amara, Dichrostachys cinerea, and Pithecellobium dulce there
was no significant difference in length of vessel elements between root and stem
woods. We therefore inferred that vessel-element length has no correlation with
the organ of the plant in which it occurs, at least in the plants we investigated.
Indeed, Carlquist (1976) himself recorded longer vessel elements in the stem
woods of Grubbia rourkei Carlq.
There was no difference between root and stem woods in qualitative features
such as vessel-element pitting, type of perforation, type of axial parenchyma,
nature of the ray, or type of fibers. We could not confirm the earlier reports
(Lebedenko, 1961, 1962; Patel, 1965; Shimaji, 1962; see also Cutler, 1976)
that xylem rays of certain plants tend to be heterogeneous in root wood but
homogeneous in stem wood.
354 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
AMOUNT OF PARENCHYMATOUS ELEMENTS
The amount of parenchymatous tissue present was considered by some earlier
workers to be a consistent difference between root and stem woods, with the
root wood tending to be more parenchymatous than the stem wood (Chalk,
1983; Esau, 1965; Fahn, 1982; Fayle, 1968; Lebedenko, 1959, 1961, 1962;
Zimmermann & Brown, 1971). However, it is not very clear whether the
increase 1s due to axial parenchyma content, ray content, or both. With respect
to rays alone, root wood was reported to have more ray content than stem
wood. This may be due to the presence of broader rays, more S per unit
area, or both. In the species we investigated, ray width in pec! de
section, calculated either in microns or in number of cells across, showed no
correlation to the organ. In some taxa the root wood had broader rays, in others
the stem wood did (see TABLE). With respect to ray abundance (number of rays
per mm? in tangential-longitudinal section), there was no consistency either.
Of the 11 species studied, only Dichrostachys cinerea and Enterolobium saman
showed greater ray abundance in root wood.
The fibers of the root wood were very much thinner walled and contained
starch grains and phenolic inclusions that were generally restricted to paren-
chyma in the stem wood. Therefore, it can be said that in all the taxa we
studied, the root wood had more thin-walled elements than the stem wood.
ACKNOWLEDGMENTS
The authors are grateful to Professor K. Periasamy for providing laboratory
facilities. The junior author (K. R.) is thankful to CSIR, New Delhi, for the
award of a Junior Research Fellowship, during the tenure of which this work
was carried out.
LITERATURE CITED
Buat, K. M. 1982. A note on cellular saan and basic density of lateral roots in
birch. Int. Assoc. Wood Anat. Bull. n.s. 3: 89-94.
Car.ouist, S. 1975. Ecological verre of xylem evolution. xi + 259 pp. Univ.
California Press, Berkeley
: . Wood eae of Roridulaceae: ecological and phylogenetic implica-
tions. Amer. J. Bot. 63: 1003-1008.
1977. Wood anatomy of Grubbiaceae. J. S. African Bot. 43: 129-144.
. 1978. Wood anatomy of Bruniaceae: correlations with ecology, phylogeny, and
organography. Aliso 9: 323-364.
980. Further concepts in ecological wood eee with comments on recent
work in wood anatomy and evolution. [bid. 9: 499-553.
,V.M. EckHart, & D.C. MICHENER. 1983. Wood anatomy of Hydrophyllaceae.
I. Eriodictyon. Aliso 10: 397-412.
CHALK, L. 1983. Roots of woody plants. Pp. 47-51 in C. R. METCALFE & L. CHALK,
Anatomy of the dicotyledons. ed. 2. Vol. |. ee Press, Oxford.
Cutter, D. F. 1976. Variations in root wood anatomy. Leiden Bot. Ser. 3: 143-156.
Esau, K. 1965. Plant anatomy. ed. 2. xvii + 767 pp. John Wiley and Sons, New York.
Faun, A. 1982. Plant anatomy. ed. 3. xi + 544 pp. Pergamon Press, Oxford.
1987] RANJANI & KRISHNAMURTHY, MIMOSOIDEAE 300
Faye, D.C. F. 1968. Radial growth in tree roots. Distribution, timing, and anatomy.
183 pp. Fac. Forest. Univ. Toronto Tech. Rep.
GOMEz- Wear z, B.G., & E. M. ENGLEMAN. 1983. Wood anatomy of Bursera longipes
and pee copallifera. Int. Assoc. Wood Anat. Bull. n.s. 4: 207-212.
INTERNATIONAL ASSOCIATION OF Woop ANATOMISTS (Committee on Nomenclature).
1964. Multilingual glossary of terms used in wood anatomy. 186 pp. Buchdruckerei
Konkordia, Winterthur.
JOHANSEN, D. A. 1940. Plant microtechnique. xi + 523 pp. McGraw-Hill Book Co.,
New Y
LEBEDENKO, L. A. 1959. The ontogeny of the wood of the roots and stems of several
representatives of Fagales. (In Russian.) Dokl. Akad. Nauk SSSR 127: 193-195.
19 Some features of the ontogeny of root and stem wood in sweet chestnut.
(In Russian.) Bjull. Moskovsk. Ob8é. Isp. Prir., Otd. Biol. 66: 66-71.
Comparative anatomical analysis of the mature wood of roots and stems
of some woody plants. (In Russian.) Trudy Inst. Lesa Drev. 51: 124-13
Pate, R.N. 1965. A comparison of the anatomy of the secondary xylem in roots and
stems. Holzforschung 19: 72-79
PLANK, S. 1976. Histologie und Verkernung des Holzes von Sambucus nigra und
Sambucus racemosa. 1. Histologie und jahreszeitliche cytologische Veranderungen.
Phyton (Horn) 17: 195-212.
RaMeEsH Rao, K., & S. K. PURKAYASTHA. 1972. Indian woods. Vol. 3. 1x + 262 pp.
Forest Research Institute, Dehra Dun.
ae K. 1962. Anatomical studies on the phylogenenc interrelationship of the genera
n the Fagaceae. Bull. Tokyo Univ. Forest 57:
Fee M. H., & C. L. Brown. 1971. ee structure and function. xii + 336
pp. Springer-Verlag, Berlin.
& D. Potter. 1982. Vessel-length distribution in branches, stem and roots of
Acer rubrum L., Int. Assoc. Wood Anat. Bull. n.s. 3: 103-109.
AL-SHEHBAZ & BATES, ARMORACIA LACUSTRIS 7
ARMORACIA LACUSTRIS (BRASSICACEAE), THE
CORRECT NAME FOR THE
NORTH AMERICAN LAKE CRESS
IHSAN A. AL-SHEHBAZ! AND VERNON BATES?
A new combination is proposed for the North American lake cress. A county
distribution map is included.
Lake or river cress is one of the most remarkable heterophyllous North
American aquatic plants. It grows in quiet waters of lakes, ponds, streams,
rivers, and springs, as well as on flood plains, mud flats, and muddy shores.
Any part of the root, stem, or leaf is capable of regenerating a new plant. The
species is widely distributed in North America east of the 95th meridian from
Wisconsin and Michigan eastward to Quebec and northwestern Vermont,
southward to Florida, westward to eastern Texas, and northward to eastern
Oklahoma, Missouri, eastern Iowa, and southeastern Minnesota (see Map).
Despite its perennial habit, its regenerating ability, and its apparent wide dis-
tribution, the species is not very common anywhere. In the northern parts of
its range, it has very rarely been collected with good fruits and seeds and appears
to regenerate and reproduce primarily asexually (La Rue, 1943).
The nomenclature of lake cress, Armoracia lacustris (which now replaces A.
aquatica), has been confused at both the specific and the generic ranks. Eaton
(see below) originally described it as a variety of horseradish (A. rusticana
Gaertner, Meyer, & Scherb., as Cochlearia armoracia L.) but later recognize
it as a distinct species of Coch/earia L. Other authors treated it as a species of
Nasturtium R. Br., Rorippa Scop., Neobeckia Greene, Radicula Moench, or
Armoracia Gaertner, Meyer, & Scherb. Under the last genus it has been known
as A. aquatica (Eaton) Wieg., but this is a later homonym of A. aquatica Kostel.
The latter is a synonym of Rorippa amphibia (L.) Besser, an entirely different
Eurasian species. Therefore, the specific epithet aquatica cannot be used for
the North American plant under the genus Armoracia. A new combination
based on Nasturtium lacustre A. Gray is proposed.
Armoracia lacustris (A. Gray) Al-Shehbaz & V. Bates, comb. nov.; based on
Nasturtium lacustre A. Gray, Gen. Pl. U. S. 1: 132. 1848. Type: same
as that of Nasturtium natans DC. var. americanum A. Gray. Gray cited
no specimens under N. /acustre but listed this varietal name as a syn-
onym
'Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138.
*Department of Biology, Memphis State University, Memphis, Tennessee 38152.
© President and Fellows of Harvard College, |
Journal of the Arnold Arboretum 68: 357-359. ce 1987.
358 JOURNAL OF THE ARNOLD ARBORETUM [vVOL. 68
VE
miles
100 200
So)
90 85 fs 5
County distribution map of Armoracia lacustris.
Cochlearia armoracia L. var. aquatica Eaton, Man. Bot. N. Amer. ed. 3. 243. 1822.
Nasturtium natans DC. var. americanum A. Gray, Ann, Lyceum Nat. Hist. New York
3: 223. 1835. Lecroryre (here designated): W. New York, Oneida Lake [4. Gray
5.n.] (GH!).
Armoracia americana (A. Gray) Hooker & Arnott, Brit. Fl. ed. 6. 28. 1850.
Rorippa americana (A. Gray) Britton, Mem. Torrey Bot. Club 5: 169. 1894.
Neobeckia aquatica (Eaton) Greene, Pittonia 3: 95. 1896.
Radicula aquatica (Eaton) Robinson, Rhodora 10: 32. 1908
Armoracia aquatica (Eaton) Wieg. Rhodora 27: 186. 1925; non A. aquatica Kostel.
Allg. Med. Pharm. Fl. 5: 1571. 1836.
Rorippa aquatica (Eaton) Palmer & Steyerm. Rhodora 40: 132. 1938.
1987] AL-SHEHBAZ & BATES, ARMORACIA LACUSTRIS 359
A few authors have questioned the placement of Armoracia lacustris and A.
rusticana in the same genus, and Rickett (1967, p. 236) stated that they ““seem
to have nothing in common except that they are both crucifers.”’ Schulz (1936)
treated the former species as a Nasturtium (sect. Rorippa (Scop.) Prantl) in the
tribe Arabideae DC. and retained the latter in Armoracia, which he placed in
the tribe Drabeae O. E. Schulz. In our opinion, both species share a number
of characters (e.g., white flowers, biseriately arranged seeds, incomplete septum,
oblong to ovate fruits, dissected lower leaves) that support their disposition in
Armoracia, aS was proposed by Wiegand (1925).
ACKNOWLEDGMENTS
We are grateful to Reed C. Rollins for a critical review of the manuscript,
to Elizabeth B. Schmidt and Stephen A. Spongberg for their editorial advice,
and to Barbara Nimblett for typing the manuscript.
LITERATURE CITED
La Rue, C. D. 1943. Regeneration in Radicula aquatica. Pap. Michigan Acad. Sci. 28:
51-61.
Rickett, H. W. 1967. Wild flowers of the United States. The southeastern states. Vol.
2, part 1. x + 322 pp. McGraw-Hill, New York.
Scuutz, O. E. 1936. Cruciferae. In: A. ENGLER & K. PRANTL, Nat. Pflanzenfam. ed.
2. 17B: 227-658.
WIEGAND, K. M. 1925. Some changes in nomenclature. Rhodora 27: 186, 187.
JOURNAL OF THE ARNOLD ARBORETUM
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Journal of the Arnold Arboretum July, 1987
CONTENTS OF VOLUME 68, NUMBER 3
A Cladistic Analysis of Conifers: Preliminary Results.
My Pe erie ede oe do bane ere eae tae eea bere ees
Taxonomic and Nomenclatural Notes on the Genus Mimosa (Le-
guminosae).
A RPE. sia sivraweree nas Were eae cae eerrs
Taxonomic Studies in Freziera (Theaceae), with Notes on Repro-
ductive Biology.
ARR er Scie eaeca sie ueedensoeee cease ees
Status of the Name Aesculus flava Solander (Hippocastanaceae).
FREDERICK G. MEYER AND JAMES W. HARDIN ................
A New Species of Lantana (Verbenaceae) from Dominica, Lesser
Antilles.
ROGER WV ARES occ bits ona ts Gas eau one eas eeeeeeateees
A Comparative Study of Root and Stem Woods of Some Members
of the Mimosoideae (Leguminosae).
Kk. RANJAN AND K..V. KRISHNAMURTHY jyscddivdessesdsaas
Armoracia lacustris (Brassicaceae), the Correct Name for the North
American Lake Cress.
IHSAN A. AL-SHEHBAZ AND VERNON BATES ..............-..--
269-307
309-322
323-334
335-341]
343-348
349-355
357-359
Volume 68, Number 2, including pages 137-268, was issued April 9, 1987.
JOURNAL OF THE
ARNOLD ARBORETUM
HARVARD UNIVERSITY VOLUME 68 NUMBER 4
ISSN 0004-2625
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JOURNAL
OF THE
ARNOLD ARBORETUM
VOLUME 68 OcToBER 1987 NUMBER 4
THE GENERA OF CYPERACEAE IN THE
SOUTHEASTERN UNITED STATES!
GORDON C. TUCKER?
CYPERACEAE A. L. de Jussieu, Gen. Pl. 26. 1789, nom. cons.
(SEDGE FAMILY)
Small to large perennial or annual herbs of aquatic or terrestrial habitats.
Roots fibrous; many species rhizomatous or stoloniferous. Plants glabrous or
Prepared for the Generic Fl ftl United States, a long-term project made Shee
by grants from the National Science Foundation and a this writing ers se BSR-8415367
(Norton G. Miller, principal investigator), under which this account wa BSR yy ie 769
(Carroll E. Wood, Jr., principal investigator). This treatment, 118th in ihe. res cutee the format
established in the first paper (Jour. Arnold Arb. 39: 296-346. 1958) and continued to the present.
The area covered by the Generic Flora ncludes North and South Carolina, Georgia, Florida, Ten-
nessee, Alabama, Mg icok Arkansas, and Louisiana. The descriptions are based primarily on the
plants of this area, with ae tion ee Saree al members of a family or genus in brackets.
ee references I did not verify are marked with asterisks.
eding the lara fore a genus, a Pea is given listing by author all familial or tribal
Pe ies pertinent to that
I have ee yed working ih ‘Norton Miller and Carroll Wood on the Generic Flora Project, ane
any
I thank t m for their interest and assistance. Thomas J. Rosatti has given helpful advice on m
occasions ae phen A. Spongberg and Elizabeth B. Schmidt improved the final manuscript with their
editorial expertise. Melin D 1 aoa initiate m es he Cyp and :
Wilbur ably supervised my graduate studies on Cyperus. Thanks a coke to the staffs of the
New York State Library especialy “Al ta Beach, eer Tibianan ), iu ssouri Botanical Garden
Library, the New York Botanical Garden Library, and the Libraries of en nold Arboretum and
the Gray Herbarium for aren access i As eee Seda : am pratt to ee curators at A,
AC, ALU, CCNL, CONN, DUKE, E, F, FSU, GA KIRI, MASS ASC , NCU, NEBC, NY,
NYS, PENN, PH, SD, SMU, UC, SL, “who | hav ve Sen specimen or Lae
access to collect s and A epitelity ere my vite Cie T. Bryson, Patricia L. Forb
aul
Goetghebeur, sone R. Guaglianone, Robert Kral, Anton A. Reznicek, alfred E. oe
Lisa A. Standley, Wm. Wayt Thomas, Marcia J. Waterway, and Karen L. Wilson have shared
i 6; Jt:;
A.S Jason R. Tucker, and Joshua D e€ e helped with field work. Vicky Martin Tucker,
my wife, has been supportive (and tolerant) of weekend coll ecting trips, week- pss research trips,
and numerous evenings given over to sedges understanding is deeply appreciated.
The illustrations were prepared by Karen - ori enbe ie: under the sapere ee Carroll Wood
or Kenneth R. Robertson, from plants collected by Carroll Wood or from specimens in the herbaria
of Harvard University (A, H).
Contribution number 543 of the New York State Science Servic
ies logical Survey, New York State Museum, The State aieion Department, Albany, New York
12230.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 361-445. October, 1987.
a02 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
scabrellate. Culms single, approximate, or caespitose, trigonous, triquetrous,
or terete, the cortex chlorenchymatous, the central region aerenchymatous or
hollow; cortical bundles with sheaths like those in the leaves. Leaves basal or
both basal and cauline; sheaths closed: blades linear to lanceolate, flat, con-
duplicate, plicate, or involute; stomata paracytic, sometimes surrounded by 1-
4 porrect or arching cuticular papillae; anatomy non-kranz or kranz, if kranz,
the bundle sheaths 2-layered (“Cyperus type’) or 3-layered (““Fimbristylis type”).
Inflorescences spicate or umbelliform [corymbose], sessile, simple, or with
second- and third- [to fifth-Jorder branching. Spikelets 1- to many-flowered,
basally subtended by a scalelike prophyll, above which may be | or more sterile
scales; flowers perfect or imperfect and monoecious (rarely dioecious), each
borne in the axil of a scale (“‘glume”’ of some authors), anemophilous (infre-
quently entomophilous); perianth absent or comprising | or 2 series of smooth
or barbed bristles, at maturity shorter to several times longer than mature
achene. Stamens (1, 2, or) 3; filaments ribbonlike or capillary; anthers broadly
ellipsoid to linear, basifixed; pollen maturing as cryptotetrads (pseudomonads),
subspheroidal, trinucleate (binucleate?) when shed. Gynoecium tricarpellate
and a 3, or bicarpellate (dorsiventrally or laterally compressed) and stig-
mas 2; styles and stigmas capillary, glabrous or glandular-pubescent; ovules
basal, anatropous, bitegmic, crassinucellar; megagametophyte (embryo sac) of
the Polygonum type. Achene trigonous or lenticular, ovoid, obovoid, or ellip-
soid, smooth, puncticulate, or papillose; endosperm mealy, with starch grains,
protein crystals, and oil droplets, filling most of the achene; embryo small;
embryogeny of the Onagrad (Juncus variation) or Asterad type; germination
epigeal. Base chromosome numbers 5, 6, 7, 8. Type GENUS: Cyperus Linnaeus.
A large family of about 80 genera and 3500 species, worldwide in distribution.
Seventeen genera occur in our area, including Carex L., with 165 species, the
largest genus of seed plants in the Southeast.
There is general agreement that the Juncaceae are the closest relatives of the
Cyperaceae (Thorne; Dahlgren & Rasmussen). Both families have tristichous
phyllotaxy, simultaneous microsporogenesis, post-reductional meiosis, non-
localized (diffuse) centromeres, anatropous ovules, and Onagrad embryogeny.
The Cyperaceae are distinguished from the Juncaceae in having conical silica
bodies in the epidermal cells, solitary ovules and basal placentation, pollen-
grain formation in which three of the meiotic products degenerate, nuclear
endosperm, and indehiscent fruits (achenes). North American Cyperaceae lack
a perianth or have one of bristles; North American Juncaceae have expanded
chartaceous tepals. This is useful regionally for distinguishing the two families,
but it cannot be used on a worldwide basis because Oreobolus R. Br. and
several other genera of Southern Hemisphere Cyperaceae also have chartaceous
tepals
Some authors (e.g., Fernald, Cronquist) have treated the Gramineae as the
closest relatives of the Cyperaceae. However, the grasses have apical placen-
tation, orthotropous ovules, distichous phyllotaxy, and open leaf sheaths, and
their affinities are with the Restionaceae and the Flagellariaceae (Thorne; Dahl-
gren & Rasmussen). Also, the grasses are chemically unlike the sedges (Har-
1987] TUCKER, CYPERACEAE 363
borne, 1971). For example, anthocyanins are common in grasses but unknown
in sedges, while aurones are common in sedges and unknown in grasses (and
in the Juncaceae).
The tribal classification was first elaborated on a worldwide basis by Nees
von Esenbeck and Kunth and has been rather stable since. Some authors
recognized tribes only; some, subfamilies and tribes; and others, subtribes also.
Two subfamilies, both distributed worldwide, are accepted in this treatment:
the Cyperoideae (Scirpoideae Pax, flowers perfect) and the Caricoideae Pax
(flowers imperfect). Included in the Cyperoideae are four tribes, of which the
Scirpeae Dumort. (including Fimbristylideae Raynal; spikelets with | or 2
sterile basal scales, numerous fertile scales spirally arranged, perianth bristles
generally present, embryos well differentiated), the Cypereae (1 or 2 sterile
basal scales, several to many fertile scales distichously arranged, perianth ab-
sent, embryos well differentiated), and the Schoeneae Dumort. (Rhynchospo-
reae Fenzl; spikelets with several sterile basal scales, fertile scales 1 or 2 (to
several), perianth bristles generally present, embryos slightly differentiated) are
represented in our area. No members of tribe Hypolytreae Fenzl (Mapanieae
Koyama) of the tropics grow in North America. Subfamily Caricoideae 1s
divided into two tribes: the Scleriae Fenzl (achenes naked, borne on a hardened
disk), represented in North America by a single genus, Sc/eria Berg.; and the
Cariceae Dumort. (achenes enclosed in a perigynium), represented in the South-
east by Cymophyllus Mack. and Carex (and also in North America by Kobresia
Willd., a circumboreal genus occurring in the northern United States and
Canada, and Uncinia Pers., an austral genus extending north to Jamaica and
Mexico).
Microsporogenesis in sedges differs markedly from that in other angiosperms.
The nucleus of the microsporocyte divides meiotically, but cytokinesis does
not follow immediately. Rather, three nuclei migrate to one end of the pollen
mother cell, where they begin to disintegrate. The fourth nucleus remains in
the center of the cell, where it divides mitotically. One of the resulting daughter
nuclei migrates to the end of the cell, joining the other three disintegrating
products of meiosis. The remaining haploid daughter nucleus divides mitoti-
cally, forming generative and tube nuclei. The generative nucleus divides again
as the exine matures, resulting in the trinucleate pollen grain characteristic of
the family. The four degenerated nuclei often remain visible as dark streaks
near the exine. The wall of the mature pollen grain is thus homologous to the
wall of the pollen mother cell. This pattern of microsporogenesis, presumably
characteristic of the entire family, has been reported in Abi/dgaardia Vahl,
Bulbostylis Kunth, Carex, Cladium P. Br., Eleocharis R. Br., Fimbristylis Vahl,
Fuirena Rottb., Scirpus L., Scleria, and Rhynchospora Vahl. In the closely
related Juncaceae cytokinesis is delayed in the pollen mother cells until each
daughter nucleus has divided a second time. Thus, the Juncaceae provide a
pattern of microsporogenesis intermediate to that in the Cyperaceae and other
monocots, and emphasizing the relationship of the Cyperaceae and the Jun-
caceae.
Embryology is nearly uniform in the Cyperaceae. Endosperm formation is
nuclear in all genera that have been investigated. Endosperm wall formation
364 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
is complete in most genera, but incomplete in Rhynchospora and Scleria. The
mature embryos of the Cyperaceae vary considerably in shape and in the
position of the cotyledon and the radicle. As a rule, each genus has its char-
acteristic type of embryo (Van der Veken). When the achenes mature, the
embryos of tribe Schoeneae are considerably less differentiated than those of
other tribes (Vanhecke).
The sedges are incompletely investigated chemically, although Cyperus is
much better known than other genera. Ethereal oils occur in the roots of three
species of Cyperus (Hegnauer). Cyanogenesis is evidently uncommon but has
been reported for three species of Cyperus and for one each of Fimbristylis and
Kyllinga Rottb. (Gibbs). This is surprising because it is widespread in the closely
related Juncaceae. Tannins occur in many sedges, having been reported in
Cyperus, Dulichium Pers., Fuirena, and Scirpus (one species each). Alkaloids
are rare; brevicarine, brevicolline, and harman occur in Carex brevicollis DC.
(Gibbs). Some terpenoids have been reported. Citral, a monoterpenoid, occurs
in species of Kyllinga (Gibbs), and several sesquiterpinoids are known from
species of Cyperus (Hegnauer). Quinones are found in both Cyperus and Fim-
bristylis (Allan et al.). Leucoanthocyanins are reported from species of Carex,
Cyperus, Dulichium, Kyllinga, and Scirpus. Anthocyanins are absent from the
family (Harborne; Harborne et a/.).
Flavonoids occur in many genera (Kukkonen, 1969; Harborne). Recently,
Harborne and collaborators have done much to expand what is known about
flavonoids in sedges. Among this class of compounds are aurones, which give
a yellowish tint to the inflorescences of many sedges. These are absent from
the Gramineae and the Juncaceae. Flavonols were present in only 15 percent
of 11 genera tested by Harborne. Flavonoid aglycones, especially quercetin and
luteolin, are widespread in the family, as are proanthocyanidins (particularly
in the leaves). Harborne and colleagues (p. 765) concluded that there are “no
dramatic correlations between flavonoid distribution and higher level classi-
fication of the Cyperaceae.”” However, certain genera or subgenera are distin-
guished chemically from closely related groups (see under Cyperus and Abild-
gaardia). Flavonoid profiles have been shown to distinguish between related
taxa in Carex and Cyperus (discussed under those genera).
Metcalfe presented much useful information on the anatomy of the Cyper-
aceae, including clear illustrations and insightful comments on the taxonomic
significance of anatomical features. Many of his descriptions were derived from
studies of ean collected in the Southeast, particularly Florida.
Devel lanatomy and morphology have received some attention (Bar-
nard), ‘The apex of spikelets in all examples studied conforms to the tunica-
corpus pattern. Periclinal division of dermatogen and hypodermal cells gives
rise to tissues that develop into the scales subtending flowers (Scirpus, Cyperus),
the carpels (in all species), the perianth bristles (Scirpus), and the perigynia
(Carex).
The first fossil remains of the Cyperaceae date from the Eocene. Fruits of
Carex, Scleria, and Scirpus are known from the Eocene and Oligocene of
Eurasia and North America; those of Dulichium and Cladium from the Oli-
1987] TUCKER, CYPERACEAE 365
gocene and Pliocene of Europe. Reports of Cyperaceae from pre-Tertiary strata
(i.e., Caricopsis Samylina) are not considered reliable (Daghlian).
REFERENCES:
ALLAN, R. D., R. L. Correti, & R. J. Wetts. A new class of quinones from certain
members Hs the family Cyperaceae. Tetrahedron Lett. 53: 4669-4672. 1969. [Cy-
peraquinones. ]
Arser, A. Water plants. xvi + 436 pp. Cambridge, England. 1920. [Review of biology
of aquatic vascular plants; Cyperaceae, 154, 416.]
ASCHERSON, P. Bemerkungen iiber das Vorkommen gefarbter Wurzeln bei den Ponte-
deriaceen, Haemodoraceen, und einigen Cyperaceen. Ber. Deutsch. Bot. Ges. 1: 498-
502. 1883. ae coloring of roots in certain species.
Bapen, J., Il], W. T. Batson, & R. STALTER. Factors affecting the distribution of
vegetation : abandoned rice fields, Georgetown Co., South Carolina. Castanea 40:
171-183. 1975. [Effects of salinity on species of Cladium, Rhynchospora, Fimbri-
stylis, and Scirpus.
BARNARD, C. Floral histogenesis in the monocotyledons. II. The Cyperaceae. Austral.
Jour. Bot. 5:115-128. 1957. [Scirpus Tabernaemontani Gmelin (as S. validus Vahl),
Cyperus Eragrostis Lam., Carex appressa R. Br
Barros, M. Ciperaceas argentinas. I. Anal. Mus. Nac. Ci. Buenos Aires 34: 425-496.
1928. [Eleocharis.] Il. Ibid. 38: 133-263. 1935. [Carex, Kyllinga, Scirpus.] Il. Ibid.
39: 253-381. 1938. [Cyperus, Lipocarpha.] IV. Ibid. 41; 323-479, 1945. [Fimbri-
stylis, PUAROSIyilS) Unie,
BATTAGLIA, | li in Hel hari fel ig (Link) Schult
Caryologia 6: 319. 332. 1954.
Beat, E. O. A manual of marsh and aquatic vascular plants of North Carolina. N.
Carolina Agr. Exper. Sta. Tech. Bull. 247. iv + 298 pp. 1977. [Cyperaceae, 93-135;
illustrations. ]
BENTHAM, G. Cyperaceae. Jn: G. BENTHAM & J. D. Hooker, Gen. Pl. 3: 1037-1073.
1883
BERGGREN, G. Atlas of seeds and small fruits of northwest European plant species. Part
2, Cyperaceae. 69 pp. + 39 pis. Lund. 1969. [Keys, descriptions, photographs of
achenes, including perigynia of Carex. ]
BLAseR, H. W. The morphology of the flowers and pale of the Cyperaceae.
141 pp. + 17 pls. Unpubl. Ph.D. Thesis, Cornell Univ.
. Studies in the morphology of the Cyperaceae. I. eats of the flowers. A.
Scirpoid genera. Am. Jour. Bot. 28: 542-551. 1941a; B. a genera.
Ibid. 832-838. 1941b; II. The prophyll. bid. a1. 53- 64. 194
BRASELTON, J. P. The ultrastructure of the non-localized eiiremee of Luzula and
Cyperus. Chromosoma 36: 89-99. 1971.
BREWBAKER, J. L. The aaa and phylogenetic significance of binucleate and
trinucleate pollen grains in the angiosperms. Am. Jour. Bot. 54: 1069-1083. 1967.
[Cyperaceae have both Rees and trinucleate genera, 1078; Fimbristylis and
Scirpus, binucleate; Carex, Cyperus, Eleocharis, Rhynchospora, and Scirpus sect.
Schoenoplectus, trinucleate.]
Brown, W. V. Variations : os associations, and origins of kranz tissue. Am.
Jour. Bot. 62: 395-402.
BURKHALTER, J. R. pence i ie vascular flora of Florida. Castanea 49: 180-186.
1984, [Cyperus ey Vahl and Scirpus deltarum Schuyler near Pensacola.]
CAROLIN, R. C., 8. W. L. Jacoss, & M. VEsK. ot ee of kranz cells in the
family Cyperaceae. Bot. Gaz. 138: 413-419.
CLARKE, C. B. New genera and species of Cyperaceae. . Kew Bull. Add. Ser. 8: 1-196.
1908. [Posthumous work; brief descriptions, synopsis of all genera and species of
the family.]
366 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
—. Illustrations of Cyperaceae. 146 pp. London. 1909.
CLIFFORD, H. T., & J. B. HARBORNE. Flavonoid pigmentation in the sedges (Cyperaceae).
Phytochemistry 8: 123-126. 1969. [Fifteen genera, 18 species.]
Cook, C. D. K. Sparganium: some old names and — types. Bot. Jahrb. 107: 269-
276. 1985. [Synonyms of Carex, Dulichium, Fuire
Cronquist, A. An integrated system of classification of qoute plants. 1262 pp. New
York. 1981.
DAGHLIAN, C. P. A review of the fossil record of monocotyledons. Bot. Rev. 47: 517-
555. 1981. [Cyperales, 535.]
DAHLGREN, R., & F. N. RASMUSSEN. Nee eae evolution. Characters and phy-
logenetic estimation. ee Biol. 16: 255-395. 1983.
Davies, J., L. G. BRIARTY, & J. O. RIELEY. Rasta of the swollen lateral roots of
the Cyperaceae. New Phytol. 72: 167-174. 1973.
Erren, L. T. Inflorescence units in the Cyperaceae. Ann. Missouri Bot. Gard. 63: 81-
112. 1976a. [Interpretation of spikelets and associated bracts and leaves with atten-
tion to homologies; many illustrations; cf. BLAser (1941a, 1941b) and KUKKONEN
(1986). ]
. The morphology of some se Brazilian species of Cyperaceae. Ibid. 113-
199. 1976b. [Eleocharis and Webster
ERDTMAN, G. Pollen morphology and et taxonomy. Angiosperms. (Corrected re-
print + new addendum.) Frontisp. + xvi + 553 pp. New York. 1966. [Cyperaceae,
141, 142; illustrations of pollen of Cladium Mariscus
EYLES, D. E., & J. L. Ropertson, Jr. A guide to the aquatic plants of the southeastern
United States. U. S. Dep. Int. Fish Wildlife Serv. Bur. Sport Fish. Wildlife Circ.
158. 151 pp. 1963. (Reprint of U. S. Publ. Health Serv. Bull. 286. 1944.
Fassett, N. C. A manual of aquatic plants (with revision appendix by E. C. OGDEN).
iv + 405 pp. Madison, Wisconsin. 1957. [Cyperaceae, 122-163.
FERNALD, M. L. Gray’s manual of botany. ed. 8. Ixiv + 1632 pp. New York. 1950.
[Cyperaceae, 236-38 |.]
Gappy, L. L. Twelve new ant-dispersed species from the southern Appalachians. Bull.
Torrey Bot. Club 113: 247-251. 1986. [Carex laxiflora, C. nigromarginata, C. stria-
tula, Scleria triglomerata.]
Gisss, R. D. emotaxonomy of flowering plants. 4 vols. xxii + 2372 pp. Montreal
and London. 1974. [Cyperaceae, 3: 1890-1893.]
Goprrey, R. K., & J. W. Wooten. Aquatic and wetland plants of southeastern United
ne Cyperaceae. Te 34: 617-639. “1985. [A thorough evaluation of the published
names. ]
Soe N. F., et al. Cyperaceae. Jn: V. L. KomMARov & B. K. SCHISCHKIN, eds.
FI. URSS 3: 1 464, 1935 (in Russian); Fl. USSR 3: 1-455. 1964 (English translation
Goop, R. E., D. F. WHIGHAM, & R. L. Simpson. Freshwater wetlands: se ae pro-
cesses and management potential. xvii + 378 pp. New York. 19 se ena
information on ecology and life history of Carex, Eleocharis, and hae
Haines, R. W. Amphicarpy in East African Cyperaceae. Mitt. Bot. Staatssam. a rineien
e . 538. 1971. ear from Bulbostylis, Scirpus.]
HAKA Meiosis and pollen mitosis in x-rayed and untreated spikelets of He-
pee ena Hereditas 40: 325-345. 1954.
ntric chromosomes in Eleocharis. Ibid. 44: 531-540. 1958.
HARBORNE, J.B. Distribution and taxonomic significance of flavonoids in the leaves of
the Cyperaceae. Phytochemistry 10: 1569-1574. 1971. [Eleven genera, 62 species;
luteolin, tricin, and glycoflavones are the characteristic flavonoids in the leaves of
sedges. ]
1987] TUCKER, CYPERACEAE Slow)
,C. A. Witiiams, & K. L. Witson. Flavonoids in leaves and inflorescences of
Australian Cyperaceae. Phytochemistry 24: 751-766. 1985. [Thirty-five genera, 170
Lae
Harris, S. W., & W. H. MARSHALL. Ecology of water level manipulation of a northern
marsh. Ecology 44: 331-343. 1963. [Permanent flooding eliminated Scirpus validus,
Eleocharis palustris, and Carex sp. after four years. ]
HEGNAUER, R. Chemotaxonomie der Pflanzen. Band 2, Monocotyledoneae. 540 pp.
Basel and Stuttgart. 1963. [Cyperaceae, 124-133.]
Heppner, J. B. The sedge moths of North America Segsesrariis Glyphipterigidae).
vii + 254 pp. Leiden. 1985. [Thirty-six species; host plan
Hesta, B. I., L. L. Treszen, & S. K. IMBAmBA. A system ae survey of C, and C,
photesviihesios in the Cyperaceae of Kenya, East Africa. Photosynthetica 16: 196-
205. 1982. [Eight genera, 220 species; 6'? carbon values for each species. ]
Hesse, M. Entwicklungsgeschichte und Ultrastruktur von Pollenkit und "Exine bei nahe
verwandten entomophilen und anemophilen Angiospermensippen der Alismata-
ceae, Liliaceae, Juncaceae, Cyperaceae, Poaceae und Araceae. Pl. Syst. Evol. 134:
229-267. 1980. [Descriptions, illustrations, and discussions of pollen of Carex acu-
tiformis Ehrh., C. vulpina L., and C. baldensis L.
Heusser, C. J. Pollen and spores of Chile. xiv + 167 pp. Tucson. 1971. [Cyperaceae,
16, 17, figs. 96-104.]
Hoi, L., J. V. PANCHO, J. P. HERBERGER, & D. L. PLuckNetr. A geographical atlas
of world weeds. (English, Arabic, Chinese, French, German, Hindi, Indonesian,
Japanese, Russian, and Spanish introductions.) xliv + 391 pp. New York. 1979.
Ho.ttum, R. E. The spikelet in Cyperaceae. Bot. Rev. 14: 525-541. 1948. [Review.]
Hotcnkiss, N. Common marsh, underwater, and floating-leaved plants of the United
States and Canada. vi + 124 pp. New York. 1972. (Reprint of U. S. Dep. Int. Fish
Wildlife Serv. Bur. Sport Fish. Wildlife Resource Publ. 44. 1967; Ibid. 93. 1970.)
[Cyperaceae, 5-8, 12-17; illustrations.]
Huana, T.C. Pollen flora of Taiwan. vi + 297 pp. + 177 pls. Taipei. 1972. [Cyperaceae,
250-260, pis. 169, 170.]
Bake G.E. A treatise on limnology. Vol. 3, Limnological botany. x + 660
w York. 1982. [Much useful information on the ecology of aquatic Cyperaceae:
hori index.
Hutcuinson, J. The families of flowering plants. 2 vols. 792 pp. Oxford. 1959. [Cy-
oar 2: 704-710; includes worldwide key to genera.
Kap.ec, J. A., & W. A. Wentz. State-of-the-art survey and evaluation of marsh plant
establishment techniques: induced and natural. 230 pp. + 3 appendices. School Nat.
Resources, Univ. Michigan, Ann Arbor. 1974.* [Review of edaphic parameters of
many aquatic vascular plants.]
Kern, F. D. North American rusts on Cyperus and acacia Mycologia 11: 134-147.
1919. [Nine species of Uredo, Puccinia, and Uromyces, with list of hosts; in several
cases the monokaryotic (sexual) as infects speci of the Compositae.]
KERN, J. H. Cyperaceae. Fl. Males. 7: 350-670.
KESSLER, J. W., & T. STARBUCK. Goperascze new c Texas and Louisiana. Sida 10: 190,
191
Koyama, T. Classification of the family Cyperaceae (1). Jour. Fac. Sci. Univ. Tokyo
Bot. 8: 37-148. 1961. [Important reference; broad generic concepts.] (2). Ibid. e
149-278. 1962a. [Caricoideae of eastern Asia.] (3). Quart. Jour. Tarwan Mus. 14
159-194. 1962b. [Cyperus Op casted. cin
KRAL, R. Further additions t n the flora of the southern states, particularly
Alabama and middle Tennessee. eee 83: 301-315. 1981. [Fifteen state records
and range extensions for species of the Cyperaceae.]
KUKKONEN, I. Gedanken und Probleme zur Systemalik der Familie Cyperaceae. Eine
Zussamenfassung. Aquilo 6: 18-42. 1967.
368 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
. Flavonoid chemistry of the ac a preliminary survey. Mitt. Bot. Staats-
sam. Miinchen 10: 622-638. 1969
. Special features of the inflorescence structure in the family Cyperaceae. Ann.
Bot. Fenn. 23: 107-120. 1986. [Arrangement of bracts, branches, and flowers; many
illustrations. ]
KuntTH, C. S. Enumeratio plantarum. Vol. 2, Cyperographia synoptica.... iii + 591
pp. Stuttgart and Tubingen. 1837. [Worldwide monograph.
Le MaAout, E., & J. Decaisne. Traité général de botanique descriptive et analytique.
vill + 746 pp. Paris. 1868. [Cyperaceae, 600-604; synopsis by tribes (Cypereae,
Scirpeae, Hypolytreae, seman (Rhynchosporeae), Scleriae, Cariceae); represen-
tative illustrations; summary of materia medica
LERMAN, J. C., & J. RAYNAL.. La teneur en isotopes stables du carbone chez les Cyper-
acées: sa valeur taxonomique. Compt. Rend. Acad. Sci. Paris, D. 275: 1391-1394.
1972. [List of C, and C, genera and subgenera; determined by 6'4 carbon values.]
Lioyp, N. P. H., & H. W. WooLHouse. Comparative aspects of photosynthesis, pho-
torespiration, and transpiration in four species of aia from the relict flora
of Teesdale, northern England. New Phytol. 83: 1979. [Carex capillaris, C.
ericetorum, Eriophorum latifolium, and Kobresia ae
Lovet, J. H. The flower and the bee. xvii + 286 pp. New York. 1918. [Flies and beetles
feed on sedge pollen.
Martin, A. C. The comparative internal morphology of seeds. Am. Midl. Nat. 36: 513-
660. 1946. [Cyperaceae, 534.] .
MattreLp, J. Zur Morphologie und Systematik der Cyperaceae. Proc. 6th Int. Bot.
Congr. 1: 330-332. 1935.
McAteg, W. L. Wildfowl food plants: their value, propagation, and management. ix +
141 pp. Ames, Iowa. 1939. [Cyperaceae, 35-45.
MeeuseE, A. D. J. Interpretive floral morphology of the Cyperaceae on the basis of the
anthoid concept. Acta Bot. Neerl. 24: 291-304. 1975. [Defends theory that simple
“bisexual” flowers of Cyperaceae are synanthial in origin.
METCALFE, C. R. Anatomy of the monocotyledons. Vol. 5, Cyperaceae. 597 pp. London.
1977.
uC
Mora, L. E. Beitraége zur Entwicklungsgeschichte und vergleichenden Morphologie der
Cyperaceen. Beitr. Biol. Pflanzen 35: 253-341. 1960.
Napper, D. M. Cyperaceae of East Africa. I. E. Afr. Nat. Hist. Soc. Natl. Mus. Jou
24: 1-18. 1964a. [Floristic account; Caricoideae.] II. [bid. 23-45. 1964b. [Scleriae
Schoeneae.] III. /bid. 25: 1-27. 1965. [Scirpeae, Cypereae (Lipocarpha).] IV. Ibid.
26: 1-17. 1966. ge (Cyperus).]
NEES VON ESENBECK, C. G. Uebersicht der ie as Linnaea 9: 273-306.
1834. [Synopsis of tribes, genera, and subgene
Nos te, R. E., & P. K. MurpHy. Short term eae csineed backwater flooding on
understory vegetation. Castanea 40: 228-238. 1975. [Tensas Parish, Louisiana; pro-
longed spring flooding of Mississippi River reduced populations of Cyperus and
Carex species.]
Ocpen, E. C. Anatomical patterns of some aquatic vascular plants of New York. New
York State Mus. Bull. 424. v + 133 pp. 1974. [Many southeastern species; Rhyn-
chospora capitellata (Michx.) Vahl misidentified as R. glomerata (L.) Vahl.
O’NeEILL, H. T. The sedges of the Yucatan Peninsula. Carnegie Inst. Wash. Publ. 522:
249-322. 1940. [Keys, detailed descriptions. ]
Patcu, E. M. Food-plant catalogue of the aphids of the ee amet the Phyllox-
eridae. Maine Agr. Exper. Sta. Bull. 393. 431 pp. 1935. [Cyperaceae, 66-69; 23
genera of aphids recorded from species of Carex, se us, Eriophorum, “Firabristylis
and Scirpus. ]
1987] TUCKER, CYPERACEAE 369
PLowman, A. B. The comparative anatomy and phylogeny of the Cyperaceae. Ann.
Bot. 20: ae 1906.
Raprorp, A. E., H. E. Antes, & C. R. BELL. Manual of the vascular flora of the Carolinas.
Ixi + 1183 pp. Chapel Hill, North Carolina. 1968. [Cyperaceae, 168-255.]
RAYNAL, J. Répartition et évolution des modes de photosynthése chez les Cypéracées.
Compt. Rend. Acad. Sci. Paris 275: 2231-2234. 1972.
Notes cypérologiques: 19. oe a la classification de la sous-famille des
Cyperoideae. Adansonia 13: 145-17 3.
Notes cypérologiques: 33. Manges nomenclaturaux (2). Ibid. 17: 273-280.
1978. erat of tribe i
Riku, M. Beitrag tomie der Cyperaceen mit besonderer Beriick-
sichtigung pee inneren Parenchymscheide. Jahrb. Wiss. Bot. 27: 485-580. 1895.
[Kranz anatomy in Cyperus, Fimbristylis, and Bulbostylis.]
SAVILE, D. B. O. A study of the species of Cintractia on Carex, oe and Scirpus
in northern America. Canad. Jour. Bot. 30: 410-435. 1952. [Rusts.]
ScuHutze-MotEL, W. Entwicklungsgeschichte und vergleichend- La eee Unter-
suchungen im Bliitenbereich der Cyperaceae. Bot. Jahrb. 78: 129-170. 1959.
. Cyperales. Pp. 602-607 in H. Metcuior, Engler’s Syllabus der Pflanzenfamilien.
ed. 12. Berlin. 1964.
SmitH, R. J., Jn. W. T. Fuincnum, & D. E. SEAMAN. Weed control in U. S. rice
production. U. §. Dep. Agr. Agr. Handb. 497. iv + 78 pp. 1977. [Photographs,
eee discussions of Rhynchospora corniculata, 53, 54; Scirpus mucronatus,
S. fluviatilis, S. acutus, 54, 55; Fimbristylis autumnalis, F. miliacea, 59, 60; Ele-
ocharis spp., 69, 70; Cyperus spp., 70-72.]
Stace, C. A., ed. Hybridization and the flora of the British Isles. xiii + 626 pp. London
and New York. 1975. [Scirpus, 510-512; Eleocharis, 512, 513; Schoenus, 513;
Carex, 513-540.]
STANDLEY, P. C. The Cyperaceae of Central America. Fieldiana Bot. 8: 239-292. 1931.
[Synoptic account. ]
TeerI, J. A., L. G. Stowe, & D. A. Livincstone. The distribution of C, species of
Cyperaceae in North America in relation to climate. Oecologia 47: 30-310. 1980.
[Floristic study; percentage of C, species decreases with latitude; cf. TucKER, 1986b,
under Cyperus.
TuHorne, R. F. A phylogenetic classification of the Angiospermae. Evol. 9: 35-106.
1976
Tietz, H. M. An index to the described life histories, early stages, and hosts of the
Macrolepidoptera of the continental United States and Canada. 2 vols. vi + 1041
pp. Sarasota, Florida. 1972. [Host references sublished through 1950; Carex, 852;
Cyperus, 875; Eleocharis, 881; Scirpus, 991, 992.]
Torrey, J. Monograph of the North American Cyperaceae. Ann. Lyc. Nat. Hist. New
York 5: 181-448. 1836. ca a thorough study.
VANHECKE, L. Embryography of some genera of the Cladiinae and the Gahniinae (Cy-
peraceae) with additional notes on their fruit anatomy. Bull. Jard. Bot. Natl. Belg.
44: 367-400. 1974. [Embryos and fruits of Cladium and Schoenus; illustrations. ]
VEKEN, P. VAN DER. Contribution a 1 i i
Cyperoideae. Bull. Jard. Bot. Natl. Belg. 35: 285-354. 1965. [Descriptions of most
of our genera (remainder treated by VANHECKE) with taxonomic comments; illus-
trations.
Winrrey, H. J., & G. L. SAMSEL. Preliminary effects on algal succession resulting from
nutrient enrichment of two central Virginia ponds with different trophic states.
Castanea 38: 140-152. 1973. [Bulbostylis capillaris, Carex spp., Cyperus strigosus,
Scirpus americanus, S. cyperinus.]
370 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
KEY TO THE GENERA OF CYPERACEAE IN THE
SOUTHEASTERN UNITED STATES
Ceneral characters: perennial (occasionally annual ), often rhizomatous herbs of diverse,
ually wet t; leaves linear, the sheaths
sean a sie ponies simple or variously branched jaeral or crowded at the apices
of the culms; flowers perfect or imperfect (the plants very rarely dioecious), borne in the
axils of scales or in ei perianth bristles present or absent; fruit an achene; embryo
small; endosperm abundan
A. Flowers a ae or carpellate flowers infrequently formed at base or apex
of spikele
B. ae of the spikelets ae) arranged.
C. Achenes without obconical or pyramidal apical tubercle, but sometimes with
eee: ieee nee oe much less than half as wide as the achene.
D. s subtended by 15- . cinnamon or whitish, silky ieee 5-10
RO as long as the achenes. ....................... 2. Eriophorum.
. Achenes subtended by aie at most 3 times as long as the achenes, or
with bristles lacking.
E. Inner whorl of perianth bristles with expanded spongy petaloid a
3. Fui
v)
E. Perianth bristles absent or lacking expanded blades.
F. Bulbous base of style persistent on mature achenes. ...........
Peuee se ule eee ayaa eu aes 6. Bulbostylis.
F. Base of style not persistent.
Ce Sty 168 MDa. aca 35, 5 wach edd eae oy ae dan 5. Fimbristylis.
G. Styles smooth.
H. Spikelets maturing a single achene; bristles absent. .....
Ae Bes atte Sete ot ate gees Pee, eave eg Cladium.
H. aaa maturing several to many achenes; bristles usu-
ally
I. Spikes and spikelets borne on rays, rarely ee
achenes and scales appressed to rachilla. .. 1. Scir
I. Spikelets sessile; achenes and scales borne at ea an-
Ghee see oe alee Be 10. Lipocarpha.
C. Achenes with pyramidal or obconical apical tubercle '2 to nearly as broad as
the achene.
J. Leaf blades absent; inflorescences unbranched, a single spikelet termi-
nating the culm. ......0..000 0.00.00 c ccc ce eee 4. Eleocharis.
J. Leaf blades presen i acca of several to many oe some usu
ally borne on branches. .....................00.. Rhy besbare:
B. Scales of the spikelets eee arranged.
K. Perianth bristles absen
L. Plants bulbous- thickened basally; style base sclerified, persistent; spikelets
(se tiers ence pe tee ent es Det ate us each tas, ae ag nein ae 7. Abildgaardia.
L. Plants not bulbous-thickened basally; style base soft, deciduous; spikelets
numerou
M. ies tence oe gee 1- to many-flowered; rachilla elon-
gate; scales broadly rounded. ....................... "yperus.
M. Inflorescences nee aenea (spikes sessile); spikelets 1- (infrequently
2- : owered; rachilla not or barely elongate; scales conduplicate, con-
CUOUSIY KECIED: co kl siudes Ga een yin een nrceere 9. Kyllinga
K. Perianth nates present.
Leaves cauline; inflorescences several, axillary. ........ 12. Dulichium.
N. Leaves basal; inflorescence solitary, terminal. .......... 13. Schoenus.
1987] TUCKER, CYPERACEAE cep
A. Flowers strictly imperfect.
O. henes naked, often borne ona discoid hypogynium. .......... 15. Scleria.
©. Achenes enclosed in perigyn
P. Spikes single, white; leaf blades broadly Spon apices broadly round-
ed, the midvein not distinguishable from other veins. .. 16. Cymophyllus.
Spikes | to several, greenish, vellowish green, or mien brown; leaf blades
linear, the apices acute, the midvein much larger and more conspicuous ae
OUNCE VEINS: 25 fpr eset tsieea tes ete ee oases ae ene Beate 17. Car
0
Subfamily CY PEROIDEAE
Tribe ScrrPEAE Kunth ex Dumortier, Fl. Belg. 143. 1827.
1. Scirpus Linnaeus, Sp. Pl. 1: 47. 1753; Gen. Pl. ed. 5. 26. 1754.
Small to medium-sized perennials or annuals of shallow fresh or tidal waters,
disturbed moist soils, moist [mesic to dry-mesic] woodlands, marshes, open
mountaintops, and grassy balds. Roots fibrous; perennial species with rhizomes
short, branched, producing loose to dense tussocks of culms; annual species
iiss ise forming dense clumps of culms. Culms trigonous (with
planar, concave, or slightly convex surfaces) or terete, smooth throughout or
ace distally. Leaves all basal or scattered along the culm; sheaths closed,
smooth or sometimes with conspicuous cross veins, greenish white, reddish
brown, or blackish; blades flat, conduplicate, or subterete, 12 to nearly as long
as the culm, stiff or arching (limp when growing underwater); stomata paracytic;
chlorenchyma not radiate; longitudinal air chambers often present. Involucral
leaves (1 or) 2-10, the blades resembling cauline ones but sheaths generally
much shorter, approximate at the summit of the culm or rather widely spaced
over the upper '4 of it, horizontal to ascendent, or the longest nearly vertical
and simulating a continuation of the culm. Inflorescences composed of primary
and secondary (sometimes tertiary) rays, in many species reduced to glomer-
ulate clusters or heads, in some to a cluster of several more or less sessile
spikelets or a single sessile spikelet; prophylls of the rays tubular, obtuse to
acute apically, smooth but usually conspicuously costate; primary rays smooth,
or scabrellate distally or throughout, terete, stiff or flexuous, secondary (and
sometimes tertiary) rays similar to primary ones, but shorter and usually more
slender. Spikelets ovoid to linear-oblong. Scales (3 to) 20 to about 100, spirally
arranged and closely imbricate, with 2 lowermost sterile and others fertile, all
deciduous at maturity, ovate to oblong, with 1-9 subtle to conspicuous nerves
and sometimes a conspicuous midrib, the apex obtuse to acute, entire or mu-
cronulate to strongly cuspidate, the awn straight to strongly excurved. Flowers
perfect, protogynous. Perianth bristles 3—-6(-8) or lacking, smooth or retrorsely
scabrellate, straight, highly curled, or crinkled at maturity, from 1 to 4 times
as long as the mature achene, deciduous or remaining attached to the mature
achene. Stamens (2 or) 3; filaments slender, about equaling the subtending
scales; anthers broadly ellipsoid to narrowly linear, the apices of the connectives
in some species prolonged as subulate appendages up to '4 the length of the
anther, sometimes tipped with crystalline prickles; pollen uniaperturate, sub-
spheroidal in polar view and triangular to obovoid in equatorial view, psilate,
372 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
1987] TUCKER, CYPERACEAE 373
bi- or trinucleate. Styles capillary; stigmas 2 or 3, about equaling the style in
length. Achenes lenticular to trigonous, equilateral in transverse section, or
slightly to strongly dorsiventrally flattened, the base stipitate or cuneate, the
apex apiculate, beaked, or entire, the surface essentially smooth, finely pitted,
reticulate, or rugulose. Embryos ellipsoid, turbinate, or fungiform, the radicle
lateral or basal. Base chromosome numbers 5, 7. LECTOTYPE SPECIES: Scirpus
sylvaticus L.; see Hitchcock & Green, Prop. Brit. Bot. 118. 1929. (Latin name
for a bulrush, probably Scirpus Tabernaemontani Gmelin.)—BULRUSH, REED,
CLUB-RUSH, WOOL-GRASS, THREE-SQUARE.
Scirpus, the third-largest genus of the Cyperaceae, with about 300 species
worldwide, is best represented in temperate regions. North America (including
Mexico), with about 80 species, is the center of diversity. Only about 15 species
occur in the West Indies and Central America, and about 30 in all of South
America, most of these in Argentina and Chile. Twelve species occur in Europe,
and perhaps 50 in Africa. It is difficult to estimate the number of species 1n all
of Asia; 24 grow in the Soviet Union, and ten in Malesia. A recent synopsis
included 44 in Australia (Wilson)
Studies in Scirpus have been hampered by lack of a worldwide treatment
(such as those prepared for several other large genera of the family, 1.e., Carex,
Cyperus, Eleocharis, and Rhynchospora). Some botanists (e.g., Wilson, Koy-
ama) have recognized each of the sections at the generic level. Most American
authors (Fernald, Schuyler), however, have recognized the genus in a broad
sense; this traditional circumscription is accepted here. Although several re-
searchers have lamented the “diverse”’ nature of the genus, most of the kinds
of variation that are represented in Scirpus are also present in Cyperus, which
Ficure I. Scirpus sect: JuNco- -SCIRPUS. a—h, S. eee (GS. validus): a, un-
derwater late in season, apex at right (note re of shoot of current
season and aecalopine eh shoots of next year’s growth), x ; es apex 7 culm with inflo-
rescence, x 1; c, single spikelet with lower flowers past anthesis (filaments visible), upper
ones with anthers visible and styles exserted, x 12; d, flower and subtending scale
moved from spikelet, view of adaxial surface, stigmas exserted, anthers still included
ee barbed bristles), x 20; e, flower, showing different maturation of stamens, the
with exserted stigmas, involucral bracts ee reduced, scalelike, spikelet Tae x 6;
j, achene with smooth bristles, x 12. k, 1, S. koilolepis: k, solitary spikelet, subtended
by scalelike involucre, scales keeled, x 6; 1, mature, trigonous, bristleless achene, x 12
m-o, S. Erismaniae: m, basal flower in axil of leaf, x 6; n, achene from basa 1 flow
12; 0, achene from cauline spikelet, x 12. p, S. cyperinus: achene with elongate aa
x 12.
374 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
has traditionally been maintained as one genus. Moreover, there has yet to
appear a thorough study of Scirpus that presents compelling arguments for
recognizing Schoenoplectus (Reichenb.) Palla, Trichophorum Pers., Baeothryon
A. Dietr., and other segregate genera. Many useful papers on the taxonomy of
single species or groups of species have been written by several authors, most
notably Schuyler.
The achenes of species of Scirpus are probably dispersed after being eaten
by waterfowl (McAtee) (wild ducks in the case of S. paludosus Nelson). Most
are digested, but those that survive have 94 percent germination, compared
with two or three percent for those treated with acid or alkali, and nine percent
after fermentation treatment (Low). Light is required for germination (Isely).
Achenes of many species, particularly S. cyperinus (L.) Kunth, are probably
dispersed by the wind, although their long, contorted perianth bristles likely
also cause them to cling to fur or feathers.
Some 30 species of Scirpus, representing six sections, occur 1n our area.
Following is a brief account of these.
Species with leafy stems are classified in three sections. In all of these, leaves
are borne along the length of the culm, while in plants of other sections they
are basal. Schuyler (1961, 1962, 1963, 1964, 1966, 1967a, 1967b, 1967c,
1971b) has studied the species with leafy stems and has provided most of the
available information on morphological variation, cytology, hybridization, and
distribution.
Section Scirpus (sect. Taphrogeton (Reichenb.) Ascherson; plants leafy
stemmed; spikelets in dense heads; achenes ellipsoid, with perianth bristles
straight, about as long as the achenes) includes the type species, the Eurasian
Scirpus sylvaticus L., n = 31, 32. The section is represented by seven species
in our area, which fall into three groups. The first includes the North American
relatives of S. sy/vaticus, among which the only representative occurring in the
Southeast is S. expansus Fern., n = 32. This bulrush grows mostly in the
Northeast, but it ranges south in the Appalachians to northern Georgia and
northern Alabama. A second eastern North American species, S. microcarpus
Presl (S. rubrotinctus Fern.), n = 33, occurs southward to the uplands of West
Virginia and also in western North America and eastern Asia.
The second group (leaves tristichous, spikelets in glomerules, plants typically
viviparous, bristles straight) includes what was treated as Scirpus atrovirens
Willd. by Fernald (1950). Schuyler (1967a, 1967b, 1967c) demonstrated that
there are four species in this group that can be distinguished morphologically
and separated geographically and phenologically. Scirpus georgianus Harper,
n = 25, 26, 27, is the most common in our area (specimens examined from
every state). It lacks a perianth and leaf cross veins (these are present in the
more northern S. atrovirens Willd., 1 = 28, which is occasional in our area
from the Ridge and Valley Province westward). Scirpus Hattorianus Makino,
n = 28, 18 a northeastern species known in our area from only six collections
from the uplands of North Carolina, Tennessee, and Alabama. Scirpus flaci-
difolius (Fern.) Schuyler, n = 27, 1s endemic to river bottoms in eastern Virginia
and northeastern North Carolina.
The third group of sect. Scrrpus (leaves distichous, spikelets in glomerules,
1987] TUCKER, CYPERACEAE BY)
plants not viviparous, bristles contorted) is represented by a single species in
North America, sia polyphyllus Vahl, n = 29, which is known from all the
Southeastern Stat
Section een (Nees) Bentham (plants leafy stemmed; perianth bristles
smooth, approximately as long as the subtending scales) is represented in our
area by three species. Scirpus pendulinus Muhl. (S. lineatus auct., non Michx.),
n = 20, has the greatest range of the three, occurring from Maine to Minnesota
south to the Gulf Coast. Scirpus lineatus Michx. (S. fontinalis Harper), n =
18, is found along the Coastal Plain from Virginia to Florida; S. divaricatus
Ell., n = 14, has a similar range but is found westward to Louisiana.
Section TRICHOPHORUM (Pers.) Darl., the wool grasses (plants leafy stemmed;
perianth bristles contorted, several times longer than the achenes), comprises
several species of cold-temperate regions. At maturity the elongate, crinkled
bristles give the spikelets and the inflorescences a woolly appearance. Extensive
hybridization in this group has resulted in a nomenclatural mire of species,
varieties, and forms. Schuyler (1962, 1967a) has carefully documented infra-
specific variation, cytology, and hybridization; he concluded that only a single
species, Scirpus cyperinus (L.) Kunth (including S. rubricosus and S. eriophorum
Michx.), » = 33, should be recognized in the Southeast. Three others that
hybridize with S. cyperinus, S. pedicellatus Fern., n = 34; S. Longii Fern.,3 n =
33; and S. atrocinctus Fern., n = 34, occur in the Northeast.
Section OxycARYUM (Nees) Beetle (plants rhizomatous; heads of spikelets
ovoid, pedunculate; scales acute, excurved) is represented in the Southeast by
a single species, Scirpus cubensis Poeppig & Kunth. In our area the species
occurs from southern Florida to Louisiana in brackish or freshwater marshes.
he affinities of this section are unclear, and no chromosome counts are avail-
Section BOLBOSCHOENUS (Ascherson) Beetle (plants tall; spikelets large, few;
scales awned, pubescent) is represented in our area by two species of freshwater
or tidal wetlands. Scirpus robustus Pursh grows in tidal marshes and estuaries
from eastern Canada to Texas. A second species, S. cy/indricus (Torr.) Britton,
occurs in marshes from Delaware to Georgia. It was confused with S. robustus
and S. etuberculatus until it was restudied by Schuyler (1975). The third species,
S. etuberculatus (Steudel) Kuntze, grows in brackish waters and is known near
the coast from Delaware to Louisiana. It is morphologically transitional to the
next wae (Fernald, 1950).
Section JuNco-scirpus Syme? (sect. Pterolepis Beurl., sect. Schoenoplectus
(Reichenb. : Bentham) (plants tall; culms often leafless; involucral leaves | or
2, more or less erect; achenes sessile, beaked, with bristles persistent) is rep-
resented in the Southeast by seven species. Scirpus pungens Vahl (S. americanus
sReported from North Carolina by Cappel and Radford and colleagues. I was unable to locate any
specimens to substantiate this. According to Schuyler (1962: pers. comm.), records of S. Longii from
‘]
‘Scirpus sect. JUNCO-scirPUS Syme in Sowerby, Engl. Bot. ed. 3. 10: 62. 1870. LecroryPE SPECIES
(here designated): S. lacustris L. Syme included three species in this section, S. /acustris, S. triqueter
nd S. pungens Vahl; S. /acustris is the only one with terete culms suggesting those of plants of
the genus Juncus L., a feature emphasized by the sectional name.
376 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
auct., non Pursh), m = 39, of sunny wetlands, 1s widespread in temperate North
America and occurs in all the southeastern states. It is closely related to S.
americanus Pursh (S. Olneyi Gray), n = 39, a taller, thicker-stemmed species
of tidal, alkaline, or saline marshes from Massachusetts to Florida and west to
southern California. The two species occasionally hybridize in brackish upper
edges of tidal marshes, but in general they are isolated ecologically. A recently
described species, S. de/tarum Schuyler, n = 39, occurs in the Mississippi Delta
region, the Mobile Bay area, and disjunctly in the prairie marshes of eastern
Kansas and Missouri. A fourth species, S. subterminalis Torrey, n = 37, is
widespread in eastern North America but is known in the Southeast from only
a few collections from the Coastal Plain and Piedmont of North and South
Carolina. Two growth forms exist: submersed, in which the leaves are filiform
and flaccid, and terrestrial or stranded, in which they are conduplicate and stiff
(Schuyler, 1972b). The highly reduced inflorescence consists of a single spikelet
subtended by one erect involucral bract. The species has an unusual photo-
synthetic metabolism: the tissues of the stem, leaf, and rhizome accumulate
malic acid at night, providing a reservoir of fixed carbon for photosynthetic
reactions during daylight (Beer & Wetzel). Such physiology is similar to that
of terrestrial plants having crassulacean-acid metabolism.
The remaining three species of sect. JUNCO-SCIRPUS were once segregated as
sect. Pterolepis (Fernald, 1950). These reportedly differ in having plumose
bristles and pedunculate clusters of spikelets. However, on a worldwide basis
several extraregional species are intermediate with respect to these two char-
acters; Koyama (1963) therefore concluded that the two sections should be
merged. Scirpus Tabernaemontani Gmelin (S. validus Vahl), n = 21, grows in
freshwater marshes nearly throughout the United States and southern Canada
and in much of the Old World; it is common throughout the Southeast. Scirpus
acutus Bigelow, n = 19, aspecies of the Midwest and Great Plains, is represented
in our area by a few collections from North Carolina and Tennessee. Dabbs
studied these two species in Saskatchewan and found that they were morpho-
logically distinct. Hybrids were occasionally found, but these were sterile and
spread only by rhizomes. A western species, S. californicus (C. Meyer) Steudel,
n = 34, is known from a few places in Louisiana, Mississippi, and South
Carolina. Other North American species of Scirpus lack its plumose perianth
bristles. Scirpus heterochaetus Chase, n = 19, might be found in the north-
western part of our area; it 1s a species of quiet calcareous waters of the St.
Lawrence and upper Mississippi drainages.
ection BAEOTHRYON Dumort.> (plants caespitose, often forming tussocks;
leaves basal; inflorescences ofa single terminal spikelet; involucral bract greatly
reduced, resembling a fertile scale of the spikelet) is represented by four species
in northeastern North America. Only one of these, the circumboreal Scirpus
cespitosus L., reaches our area, growing in the grassy balds of the high mountains
of North Carolina, Georgia, and Tennessee. The southeastern populations are
disjunct from the nearest occurrences of the species in the northeastern United
‘Scirpus sect. BAEOTHRYON Dumort. Fl. Belg. 143. 1827. Fernald (1947, 1950) and other authors
have attributed the sectional name to Endlicher (Gen. Pl. 118. 1836).
1987] TUCKER, CYPERACEAE Sele
States (in the Adirondack Mountains of New York) by some 1200 km. A
widespread but easily overlooked species of the northeastern and midwestern
United States, S. verecundus Fern., has not yet been collected in our area but
might occur in the uplands of North Carolina, Tennessee, or Arkansas. It is
perhaps the most mesic species of the genus in North America, inhabiting dry
woodlands and basic ledges, in contrast to the aquatic habitats of most species
of Scirpus
Section Tonnes (R. Br.) Griseb. (plants annual; inflorescences unbranched;
spikelets sessile, few) is represented in our area by five species. Scirpus koilolepis
(Steudel) Gleason probably occurs in all the states in our area, as well as in
the Midwest and the Great Plains. The remaining species are much less frequent
and are local in range. Scirpus Erismaniae Schuyler, n = 5, is recorded from
Georgia, western Florida, and Alabama. This species produces basal spikelets
on very short culms (see FiGureE 1), as do several African species of this section
(Haines). Scirpus molestus M. C. Johnston, described from Texas, also occurs
in southern Louisiana. The remaining species have perianth bristles (in most
collections) and have been distinguished by some authors (e.g., Fernald, 1950)
as sect. Actaeogeton (Reichenb.) Beetle. Scirpus Hallii Gray, n = 11, known
from widespread localities in the eastern United States, has been collected in
Georgia; and §. Purshianus Fern., n = 19, a primarily northeastern species, is
known in the Southeast from North and South Carolina, Tennessee, and Geor-
gia.
REFERENCES:
Under family references see BADEN et a/.; BARNARD; BARROS (1935); BENTHAM; BLASER
(1941a, 1941c); BURKHALTER; CLARKE (1908, 1909); ErTEN (1976a); EvLes & ROBERTSON;
& RAYNAL; McATEE; MEEUSE; METCALFE; Mora; O° NEILL, RADFORD et al.; RAYNAL
(1972, 1973); Riku; SAVILE; SCHULZE-MOTEL (1959, 1964) ; TEER! et al.; TIETZ;
Torrey; and WINFREY & SAMSEL.
Beer, S., & R. G. WeTzEL. Photosynthetic carbon metabolism in the submerged aquatic
angiosperm Scirpus subterminalis. Pl. Sci. Lett. 21: 199-207. 1981.
oa Le, A. A. Studies of the ae Scirpus L., V. Notes on the section Actaeogeton.
m. Jour. Bot. 29: 653-656. 1942.
. Akey to the North nee species of the genus Scirpus based on the achene
characters. Am. Midl. Nat. 29: 533-538. 1943.
Studies in the genus Scirpus VII. Conspectus of sections represented in the
Americas. Am. Jour. Bot. 31: 261-265. 1944. [See review by FERNALD (1947).]
. Sedge boats in the Andes. Jour. N. Y. Bot. Gard. 46: 1-4. 1945. [Boats in Lake
Titicaca made from culms of S. Tatora Steudel.]
. Cyperaceae: Scirpeae. Scirpus. N. Am. FI. 18: 479-504. 1947. [Keys, descrip-
tions. ]
. Annotated list of original descriptions of Scirpus. Am. Midl. Nat. 41: 453-493.
1949. [Worldwide.]
. Bulrushes (Scirpus) and their multiple uses. Econ. Bot. 4: 132-138. 1950. [Sum-
mary of economic importance.
CappEL, E. D. The genus Scirpus in North Carolina. Jour. Elisha Mitchell Sci. Soc. 70:
85-91. 1954. [Keys, descriptions, distributions
378 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Dasss, D. L. A study of Scirpus acutus | nae validus in the Saskatchewan River
delta. Canad. Jour. Bot. 49: 143-153.
FERNALD, M. L. Studies of the North fens species of Scirpus. Rhodora 45: 279-
296. es
tified bibliography of Scirpus. Ibid. 49: 49-52. 1947, [Review of BEETLE
1944); pie infrageneric names used by Beetle were not published at the ranks
indicated. }
HANSETER, R. Recovery, productivity, and p r tent of selected marsh plants
after repeated cuttings. 81 pp. ae M. S. Thesis, Univ. Wisconsin, Oshkosh.
975.* |S. acutus and S. validus recovered well even when harvested as frequently
as every two weeks; S. fluviatilis, however, decreased in number and size of shoots
under this treatment; information from Goon et al.]
IseLy, D. A study of the conditions that affect the germination of Scirpus seed. Cornell
Univ. Agr. Exper. Sta. Mem. 257. 28 pp. 1952. [Light enhances germination in many
species. }
Koyama, T. Taxonomic study of the genus Scirpus Linné. Jour. Fac. Sci. Univ. Tokyo
Bot. a 271-366. 1958. [Broad generic concept including Eriophorum, Fuirena.]
he genus Scirpus Linn. Some North American aphylloid species. Canad. Jour.
Bot. a 913. 1962.
. The genus Scirpus Linn. Critical species of the section Pterolepis. [bid, 41: 1108-
1131. 1963.
B. C. Stone. The genus Scirpus in the Hawaiian Islands. Bot. Mag. Tokyo
73: 288- . 1960.
Licutcap, B. W., & A. E. SCHUYLER. Scirpus triqueter established along tidal portions
of the ea River. Bartonia 50: 23, 24. 1984.
Low, J. Germination tests of some aquatic plants important as duck foods. 27 pp.
Unpubl. B.S. Thesis, Utah State Univ., Logan. 1937.* [Summarized by G. E.
HUTCHINSON. ]
Lye, K. A. Moderne oppfatning av slekta Scirpus L. Blyttia 29: 141-147. 1971.*
Raymonpb, M. Additional notes on some Southeast Asian Scirpus. Nat. Canad. 84: 111-
957
AL, J. ) :
sect. Supini. Adansonia, II. 16: 119-155. 1976. [Formation of basal spikelets in
SAVILE, D. B. ome rusts of Scirpus and allied genera. Canad. Jour. Bot. 50: 2579-
micv ee
SCHUYLER, A. E. Evidence for the hybrid origin — Peckii. Rhodora 63: 237-
243. 1961. [Sterile hybrid of S. atrovirens and S. atrocinctus or S. pedicellatus.]
Sporadic culm formation in Scirpus Longii. eee 32: 1-S. J unnumbered
pl. 1962. [Report from North Carolina (FERNALD, 1943) based on misidentification
of S. cyperinus, see also SCHUYLER & STASZ.]
—. Notes on five _ of Scirpus in eastern North depen Tbid. 33: 1-6. 1963.
[Comments on taxonomy of S. ancistrochaetus, S. atrovirens, S. divaricatus, S.
fontinalis, and S. jeer chromosome counts for ea
biosystematic study of the Scirpus CD as oes Proc. Acad. Nat. Sci.
Phila. 115: 283-311. 1964. [Hybridization of S. cyperinus and related species.]
. The taxonomic delineation of Scirpus lineatus and Scirpus pendulus. Not. Nat.
390: 1-3. 1966. [With nomenclatural comments.]
A taxonomic revision of the North American leafy species of Scirpus. Proc.
Acad. Nat. Sci. ee 119; 295-323. 1967a. [Keys, descriptions, chromosome num-
bers for 18 species. ]
ens Hattorianus in North America. Not. Nat. 398: 1-5. 1967b. [Common
d with S. atrovirens and S. georgianus; south-
ern range limit i in North Carolina ermal
1987] TUCKER, CYPERACEAE 519
A new status for an eastern North American Scirpus. Rhodora 69: 198-202.
1967¢. [S. jeauailie distinguished from S. atrovirens.]
hree new species of Scirpus (Cyperaceae) in the southern United States. Not.
Nat. 423: |- 12. 1969. [S. Bergsonii, S. Erismaniae, and S. Wilkensii, all from Gulf
Coastal Plain, related to S. Hallii; chromosome counts and specimen citations.]
A new North American aquatic bulrush (Cyperaceae: Scirpus). Ibid. 427: 1-3.
1970, [S. deltarum from Mississippi, Louisiana, Alabama, and Missouri, related to
S. pungens, illustrations. ]
Some relationships in Scirpeae bearing on the delineation of genera. Mitt. Bot.
Staatssam. Miinchen 10: 577-585. 1971la
. Scanning electron microscopy of achene epidermis in species of Scirpus (Cy-
peraceae). Proc. Acad. Nat. Sci. Phila. 123: 29-52. 1971b. [Survey of epidermal
features of Scirpus and some species of Eriophorum with comments on taxonomy;
clear, eae photographs.]
me numbers of Scirpus Purshianus and S. Smithii. Rhodora 74: 398-
406. 19722. eae of named forms of each species; S. Purshianus, n = 19;
S. Smithii, n =
: pclae and anatomical differences in leaf blades of three North Amer-
ican aquatic bulrushes (Cyperaceae: Scirpus). Bartonia 41: 57-60. 1972b. [S. etu-
berculatus, S. subterminalis, and S. Torreyi; illustrations; these closely related species
differ greatly in anatomy of leaf blades.]
. Scirpus cylindricus: an ecologically restricted eastern ae ao tuberous
bulrush. /bid. 43: 29-37. 1974. [Illustrations, specimen citatio
——. Chromosome numbers of some eastern North aes ates of Scirpus.
Ibid. 44: 27-31. 1975.
J. L. Stasz. Influence of fire on reproduction of eee Longii. Bartonia 51:
105-107. 1985. [Fire ie culm formation and flow
SEIDEL, K. Macrophytes and water purification. Pp. (09-122 in J. TouRBIER & R
Pierson, eds., Biological control of water pollution. New York. 1976. [S. lacustris
and other marsh plants used to treat wastewater in artificial marshes in northern
Europe. ]
& R. Kickutu. Biological treatment of phenol-containing wastewater with bul-
rush (Scirpus lacustris L.). Wasserwirtschaft-Wassertechnik 17: 209, 210. 1967.*
ee zed by Goon et a
Séropes, J. B., J. DESCHENES, & J.-P. TourpE. Temps de submersion des marais a
sos (Sims americanus) de Vestuaire du Saint-Laurent. Nat. Canad. 112: 119-
129,
SMITH, S. a i tural hybridization 1 in the Scirpus lacustris complex in the north central
United States. Pp. 175-200 in J. G. GuncKEL, ed., Current topics in plant science.
New York. 1969.
— ology of the rane lacustris complex in North America. Polsk. Arch. Hy-
drobiol. 20: 215, 216. 1973
ENDER pb, M., & R. G. WETZEL. Photorespiration and internal recycling of CO,
in the Bee angiosperm Scirpus subterminalis. Canad. Jour. Bot. 58: 591-598.
1980.
STEINMANN, F., & R. BRANDLE. Carbohydrate and protein metabolism in the rhizomes
of the bulrush (Schoenoplectus lacustris (L.) ee in eum o i ia development
of the whole plant. Aquatic Bot. 19: 53-64. 1984. [S. lacu
WEsTHOFF, V., & M. F. M6rzer Bruns. De gro ae van eee americanus Pers.
op het Groene Strand bij West-Terschelling. (English summary.) Acta Bot. Neerl.
§: 344-354. 1956. [Optimum habitat for S. pungens at upper edge of tidal marsh,
where the salinity was less than 9 g chlorine/liter; disturbance reduced competition
to the benefit of this species. ]
Witson, K. L. A synopsis of the genus Scirpus sens. lat. (Cyperaceae) in Australia.
380 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Telopea 2: 153-172. 1981. [Keys, descriptions, discussions for 43 species; subgenera
recognized as genera. ]
2. Eriophorum Linnaeus, Sp. Pl. 1: 52. 1753; Gen. Pl. 27. 1754.
Small to medium-sized, single-stemmed or loosely caespitose [densely caes-
pitose or tussock-forming] perennials of bogs, swamps, and pocosins. Roots
fibrous; rhizomes short, horizontal to oblique. Culms terete or nearly so, gla-
brous. Leaves basal and cauline; sheaths glabrous, ligules lacking; blades flat
[conduplicate], the midrib conspicuous, the margins scabrellate, especially dis-
tally; chlorenchyma not radiate; air chambers present. Inflorescences of 1 to
several sessile or pedunculate spikelets; bracts 1-6, closely spaced at the summit
of the culm, oblique or slightly reflexed [ascendent to erect], sheaths very short,
blades leaflike; rays short [elongate and drooping or absent]. Spikelets oblong-
ovoid; empty basal scales 3—5[-15]. Scales 50-150, oblong-ellipsoid, acute to
obtuse, 1- to 5-nerved, deciduous after the achenes mature. Flowers perfect.
Perianth bristles [6 to] 12 to ca. 50, about equaling the scales at anthesis but
elongating greatly as the achenes mature. Stamens 1 [or 2 or 3]; filaments
flattened; anthers linear [ellipsoid], the apices of the connectives not prolonged.
Styles capillary, glabrous; stigmas 3, about as long as the style. Achenes tri-
gonous, slightly compressed dorsiventrally, oblong-ellipsoid (widest in distal
half), the apex obtuse, apiculate, the base sessile, the surface smooth, glossy.
Embryos more or less turbinate [obconical or ellipsoid], the radicle sublateral.
Base chromosome number 29. Type species: FE. vaginatum L.; see Britton &
Brown, Illus. Fl. No. U. S. Canada, ed. 2. 1: 322. 1913. (Name from Greek,
erlos, cotton or wool, and phoros, bearing, in reference to the cottony mature
inflorescence.) — COTTON-GRASS, BOG-COTTON.
A genus of about 12 species of boreal regions. About eight species are cir-
cumpolar, occurring in both northern Eurasia and northern North America.
There is relatively little endemism. Only Eriophorum virginicum L. occurs in
the Southeast; it ranges from Newfoundland to Minnesota southward and is
known in our area from a few scattered collections made in the mountain bogs
of North Carolina and Tennessee and the Coastal Plain swamps of North
Carolina, South Carolina, and Georgia (southern limit in the Okefenokee
Swamp). No species of the genus is reported from Missouri or Kentucky, and
only £. virginicum occurs in Virginia and West Virginia.
A few workers (e.g., Koyama) have treated the cotton grasses as constituting
Scirpus sect. Vaginati (Andersson) Koyama, but most have kept Eriophorum
separate from Scirpus. The two genera are readily distinguished by the number
and the length of the perianth bristles. Eriophorum is divided into two sections
(Goncharov et a/.), each with about six species: sect. ERIOPHORUM (sect. Va-
ginati Andersson) contains those species in which the inflorescence is a single
sessile spike, while sect. PHYLLANTHELA Andersson comprises those (including
FE. virginicum) in which the inflorescence consists of several pedunculate spikes.
The genus is almost uniform cytologically; ten of the 12 species have been
counted as n = 29. Two are n = 27, and in the case of Eriophorum angustifolium
L., n = 29 and n = 35 have been reported.
1987] TUCKER, CYPERACEAE 381
Hybridization is known among both the Eurasian and the North American
species. Although it is generally not difficult to distinguish Eriophorum virgini-
cum from the other members of the genus, there are species pairs that appear
to intergrade—for example, E. angustifolium and E. viridicarinatum (Engelm.)
Fern. It is surprising that the genus has not received more systematic study,
considering its broad distribution.
The circumboreal Eriophorum alpinum L., n = 29, was placed in Scirpus
(as S. hudsonianus) by Fernald. Following a survey of epidermal features of
achenes of Scirpus and Eriophorum, Schuyler concluded that the species be-
longs in Eriophorum. Its chromosome number also supports this placement.
In the Arctic, species of Eriophorum are dominant and sometimes form a
vegetation type known as “‘tussock tundra.” The plants provide an important
forage for deer and caribou in North America and for sheep, ponies, and
reindeer in northern Europe and Asia. In the United States the plants are seldom
dominant (except in alpine grasslands in limited montane areas). However,
they sometimes form a conspicuous element of fen and bog vegetation because
of their showy fruiting heads.
Wein summarized ecological information about Eriophorum vaginatum, a
circumboreal tussock-forming species. Species of Eriophorum occurring in the
eastern United States are rhizomatous or rather loosely caespitose. There is
much information on the autecology and physiological ecology of the genus,
although nearly all is derived from studies of FE. vaginatum.
Despite its abundance in arctic regions, Eriophorum has conspicuously few
insect herbivores. Larvae of the cottongrass moth, Celaena haworthi Curtis,
tunnel in the culms of E. vaginatum in Europe, but no macrolepidopteran
species is reported to feed on Eriophorum species in North America (Tietz).
The aphid Rhopalosiphum eriophori (Walker) is reported on E. angustifolium
and E. vaginatum. The larvae of the beetle Plateumaris discolor (Panzer) live
in anaerobic conditions among the roots of E. vaginatum in Europe, obtaining
needed oxygen by tapping into the intercellular air spaces in the cortex of the
roots.
REFERENCES:
Under family references see BENTHAM; BERGGREN: BLASER (194 1a, 1941c), GONCHAR-
ov et al.; Hotttum; Le Maout & DECAISNE; LERMAN & RAYNAL, LLoyp & WOOLHOUSE;
METCALFE; PATCH; RAYNAL (1972, 1973); TreTz; and TORREY
Under Scirpus see Koyama (1958) and SCHUYLER (1971b).
Faecri, K. Zur Hybridbildung in der Gattung Eriophorum. Verh. Inst. Rubel Ziirich
33: 50-58. 1958. [Hybridization of several European species, many illustrations.]
FERNALD, M. L. The North American species of Eriophorum. Rhodora 7: 8 1-92, 129-
136. 1905. [Eight species.]
Fetcuer, N., & G. R. SHAVER. Growth and peas patterns within tussocks of Erio-
phorum vaginatum. Holarct. Ecol. 5: 180-1
& Life histories of tillers of Eriophor um vaginatum in relation to tundra
disturbance. Jour. Ecol. 71: 131-14 83.
Goopman, G. T., & D. F. PERKINS. role of mineral nutrients in Eriophorum
communities. III. Growth response to added inorganic elements in two E. vaginatum
382 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
communities. Jour. Ecol. 56: 667-683. 1968; IV. Potassium supply as a limiting
factor in an E. vaginatum reaping Ibid. 685- 696. 1968.
Hype, H. A., & K. F. ADAMS. atlas of airborne pollen grains. xvi + 112 pp. London
and New York. [E. intidielum Honck., 28, 29.]
RayMonb, M. Two new Eriophorum hybrids from northeastern North America. Sv.
Bot. Tidskr. 45: 593. 1951. [E. Pylaieanum (E. spissum x E. russeolum), E. Porsildii
(E. Chamissonis x E. spissum).]
What is Eriophorum Chamissonis C. A. Meyer? Ibid. hai 65. 1954.
ReceRON: K. P., & H. W. WooLHouse. Studies of the seasonal course of carbon uptake
of Eriophorum vaginatum in a moorland habitat. II. The seasonal course of pho-
tosynthesis. Jour. Ecol. 73: 685-700. 1985.
WEIN, R. W. Biological flora of the British Isles: Eriophorum vaginatum L. Jour. Ecol.
61: 601-615. 1973.
L. C. Buiss. Changes in arctic cottongrass tussock tundra communities. Arct.
Alp. Res. 6: 261-274. 1974.
3. Fuirena Rottboell, Descr. Icon. 70. 1772.
Rhizomatous perennials or caespitose annuals of sunny, wet, often disturbed
soils. Rhizomes horizontal, covered with persistent lanceolate scales, producing
cormlike axillary offshoots from which new culms arise. Culms erect or slightly
inclined, unbranched, terete, hollow. Leaves with sheaths tubular, costate, pu-
bescent, barely reaching to decidedly separated from the base of the next sheath,
the ligules hyaline, hispid (or glabrous) apically; basal leaves bladeless, cauline
leaves with blades lanceolate to linear, flat or slightly conduplicate [crescen-
tiform], pubescent (blades absent or reduced to an awned apex of the sheath
in | species); stomata paracytic; chlorenchyma not radiate. Inflorescences of |
to several sessile or pedunculate glomerules in the axils of the upper leaves:
rays lacking or |—-4, smooth or hispidulous. Spikelets 1-6, ovoid to oblong.
Scales 30-60(—100 or more), ovate to oblong, widest at or above the middle,
hispid adaxially, less often glabrous or glabrescent, 3- to 9-nerved, the 3 central
nerves prolonged into a cuspidate, straight, or excurved apex '4 as long as to
nearly equaling the length of the body of the scale, the 3 basal scales sterile,
longer, narrower, and more conspicuously awned than the fertile ones. Flowers
perfect, protogynous. Perianth biseriate [uniseriate or absent], outer whorl (se-
pals) of 3 smooth or retrorsely scabrellate bristles, 4 to nearly as long as the
achene; inner whorl (petals) of bristles bearing expanded, entire [fimbriate],
hyaline to somewhat spongy blades with obtuse, acute, aristate, or emarginate
apices. Stamens 3 (infrequently 1, 2, or 6); filaments ribbonlike, about as long
as the subtending scale; anthers linear to ellipsoid; pollen grains uniaperturate,
obovoid to subspheroidal, psilate, trinucleate. Styles linear, frequently hispid:
stigmas 3, linear, about as long as the styles, pubescent. Achenes trigonous with
conspicuous ridged angles, ellipsoid, the apex acute but not apiculate, the base
stipitate (usually conspicuously so), the faces flat to slightly concave, delicately
striate or smooth [cancellate], glossy. Embryo fungiform. Base chromosome
number 23. (Including Vaginaria Persoon.) Lectotype species: F. umbellata
Rottb.; see Britton & Brown, Illus. Fl. No. U. S. Canada, ed. 2. 1: 337. 1913.
(Named for Joergen Fuiren, 1581-1628, Danish physician.)
1987] TUCKER, CYPERACEAE 383
A warm-temperate and tropical genus of about 30 species. Seven occur in
the Southeast; these are well known through Kral’s recent revision. An addi-
tional three occur in the southwestern United States. Fuirena repens Boeck. 1s
endemic to Mexico, while five primarily South American species extend north-
ward into Central America, Mexico, and the West Indies. About 12 species
occur in South America, and about as many in Africa. Only F. umbellata 1s
recorded in Europe, and it is limited to the southern part of the continent. Five
species occur in southern Asia, but none is recorded from the Soviet Union.
Most of our species are distributed from Texas to Florida along the Gulf
Coastal Plain and northward on the Atlantic Coastal Plain. Fuirena scirpoidea
Michx. and F. /Jonga Chapman occur only as far north as southern Georgia,
F. breviseta Cov. as far as eastern Virginia, F. squarrosa Torrey north to Long
Island, and F. pumila to Cape Cod. The last species is disjunct in southern
Michigan and northern Indiana. Two others in our area, F. Bushii Kral and
F. simplex Vahl, are southern Great Plains species that occur eastward to
Louisiana, Arkansas, and Missouri.
All of the southeastern species have haploid chromosome numbers of 23.
The only exception is Fuirena simplex, for which n = 15 has been reported
from Texas populations, in addition to n = 23 from southeastern representa-
tives (Kral).
Plants of Fuirena have no reported economic significance in North America,
although F. glomerata Lam. and F. umbellata have been reported as important
weeds in Borneo, India, Taiwan, and Malaysia (Holm et al.).
REFERENCES:
Under family references see BEAL; BENTHAM; BLaser (1940, 194 1a); CLARKE (1908,
1909); ee aoe ROBERTSON; FASSETT; GODFREY & WooTEN; HESLA et al.; HOLM
et al.. Hotttum; Huan; J. Hutcuinson; J. H. Kern, KUNTH; Le MA Ba, Ae
METCALFE; Tae aG6s), NEES VON oo O'NEILL; SCHULZE-MOTEL (1959, 1964);
STANDLEY; TORREY; and VAN DER VEK
Under Scirpus see KoyAMA (1958).
Busn, B. F. The North American species of Fuirena. Rep. Missouri Bot. Gard. 16: 87-
99. 1905. [Eight species; keys, descriptions, specimen citations.
Covite, F. V. Revision of the United ee species of the genus Fuirena. Bull. Torrey
Bot. Club 27: 1-14. 1890. [Four speci
Fores, P. L. Studies in Cyperaceae of ee Africa: VI. A new combination in
Fuirena with notes on the species. Jour. S. Afr. Bot. 35: 83-98. 1969. [F. hirsuta
(Berg.) see good illustrations of inflorescences, perianth parts, achenes |
Scanning electron microscopy of the leaf blade epidermis of Fuirena Rottb.
(cee Proc. Electron Microscop. Soc. S. Afr. 3: 27, 28. 1973. [Adaxial epi-
dermis and substomatal chambers showing su uress differences. ]
———. Studies in Cyperaceae in southern Africa: 11. A new species of Fuirena Rottb.
S. Afr. Jour. Bot. 3: 359-362. 1984. [F. ane from eastern Cape Province and
Lesotho; illustrations. ]
C. M. LALKuaN. A preliminary study of silicon distribution in the leaf blade
epidermis of eee coer as (Cyperaceae). Proc. Electron Microscop. Soc. S.
Afr. 13: 79, 80. 1983.
GOVINDARAJALU, E. th systematic anatomy of South Indian Cyperaceae: Fuirena Rottb.
384 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Bot. Jour. Linn. Soc. 62: 27-40. 1969. [F. uncinata Kunth, F. Wallichiana Kunth,
F. pubescens Kunth, F. ciliaris (L.) Roxb., and F. umbellata Rottb.; cross sections
of leaves and culms; species distinguishable by surface features of leaf blades; key.]
Hoi, T. Studies in the Cyperaceae. V. Fuirena squarrosa Michx. and F. scirpoidea
Vahl. Am. Jour. Sci. 154: 13-26. 1897. [Morphological and anatomical study of
two species of the Southeast; illustrations. ]
KRAL, R. A synopsis of Fuirena (Cyperaceae) for the Americas north of South America.
Sida 7: 309-354. 1978. [Keys, descriptions, illustrations, chromosome counts.]
4. Eleocharis R. Brown, Prodr. 224. 1810.
Small to medium-sized, loosely to densely caespitose or single-stemmed,
rhizomatous or stoloniferous, submersed, emergent, or littoral perennials (rare-
ly annuals) of marshes, ditches, and pond and river shores. Roots fibrous:
rhizomes (lacking in some species) slender, horizontal, covered with appressed
ovate to lanceolate scales. Culms terete or ellipsoid (less often trigonous, quad-
rangular, or flattened), solid or hollow (sometimes with thin transverse par-
enchymatous septa), smooth, with numerous paracytic stomata (submersed
lower portions of culms with few or no stomata); in submersed species sec-
ondary branches present, very closely spaced and seemingly verticillate. Leaves
|—4; sheaths closely fitting the base of the culm, the summit firm or scarious
(sometimes apiculate); blades lacking. Inflorescences single spikelets terminat-
ing the culms. Spikelets slenderly cylindrical to ovoid, slightly less than to
about 3 times thicker than the summit of the culm. Scales (2-)20-100, oblong,
lanceolate, obovate, or orbiculate, hyaline, firm, or coriaceous, strongly to
weakly nerved or nerveless, deciduous or persistent. Flowers perfect. Perianth
bristles (3—-)6(—12) or absent, extrorsely or retrorsely barbed or smooth, per-
sistent on the base of the mature achene or falling from it. Stamens 3; filaments
hyaline, about equaling to shorter than the subtending scale; anthers ellipsoid
to linear; pollen grains |- [to 4-Japerturate, obovoid to subspheroidal, psilate
(scabrellate), trinucleate. Styles with swollen, bulbous base; stigmas 2 or 3,
capillary. Achenes lenticular or trigonous, ovoid, obovoid, or ellipsoid, the base
broadly rounded, the apex capped by a small to large, pyramidal, conical, or
swollen tubercle, the surface smooth or variously reticulate, dull, frequently
glossy, or iridescent. Embryos turbinate to fungiform. Base chromosome num-
ber 5. Type species: E. palustris (L.) Roemer & Schultes (Scirpus palustris L.);
see Britton & Brown, Illus. Fl. No. U. S. Canada, ed. 2. 1: 310. 1913. (Name
from Greek, helos, marsh, and charis, grace, from the paludal habitat of most
species.) — SPIKE-RUSH, DOG’S-HAIR GRASS.
A genus of about 250 species, worldwide in distribution. Eleocharis is evi-
dently closely related to Scirpus but is distinguished by its leafless culms and
its single, erect, terminal spikelets. Although the apical tubercles of the achenes
of Eleocharis are similar to those of some species of Fimbristylis Vahl, sug-
gesting that Eleocharis is most closely related to that genus (Svenson, 1929),
recent evidence supports a closer relationship between Scirpus and Eleocharis.
Both of these genera have non-kranz anatomy, while Fimbristylis has kranz
anatomy (Metcalfe). The embryos of Eleocharis (turbinate to fungiform, radicle
basal, coleoptile lateral) are similar to those of species in Scirpus sect.
1987] TUCKER, CYPERACEAE 385
BoOLBOSCHOENUS (Van der Veken), rather than to those of Fimbristylis (turbinate
to fungiform, radicle lateral, coleoptile basal).
Some 40 species occur in the Southeast, and many of these have rather wide
ranges. Holarctic, neotropical, and pantropic groups are represented in our
area. Svenson’s (1929, 1957) division of the genus into seven series has received
wide acceptance, and our species are presented here according to his classifi-
cation. The two largest are ser. ELEOCHARIS (ser. Palustriformes Svenson) and
ser. TENUISSIMAE Svenson, having 13 and ten species in our area, respectively.
Plants of ser. ELEOCHARIS are characterized by slender culms and a stolon-
iferous habit; there are both tristigmatic and distigmatic species. Our repre-
sentatives are mostly northeastern species that occur southward only as far as
Virginia, Tennessee, or Arkansas. However, Eleocharis fallax Weatherby, 2 =
42, and E. arenicola Torrey, 2n = 20, both of the Coastal Plain, are found in
most of the Southeastern States. Eleocharis montevidensis Kunth, 2n = 10, 20,
is a neotropical species that has been found north to the Carolinas and Cali-
fornia; it is sometimes treated as conspecific with E. arenicola.
Plants of ser. TENUISSIMAE are loosely caespitose and have slender, wiry
culms. Our species are mostly restricted to the Coastal Plain. In the Southeast
the neotropical Eleocharis nana Kunth has been found only in southern Florida,
while E. nodulosa (Roth) Schultes occurs along the Gulf Coast from Florida
to Louisiana. The most widely distributed of our species, E. tuberculosa (Michx.)
Roemer & Schultes, 2” = 30, is found throughout the Southeast northward to
Nova Scotia. It is distinctive in having perhaps the largest tubercle in any
species of the genus—as large as the body of the mature achene.
Plants of ser. MUTATAE Svenson are the tallest in the genus; three of our
species regularly reach | m. The plants are characterized by spikelets that are
barely wider than the apices of the subtending culm and that have persistent
scales. The plants are unusual ecologically because they grow in ponds or pools
with a stable water level. Most other species of the genus grow where receding
water levels leave the plants exposed in summer. Species of ser. MUTATAE have
very high chromosome numbers. Briggs has made counts for the Australian
Eleocharis equisetina Presl, 2n = 172, and E. sphacelata R. Br., 2n = 94-100,
140, 180, 188. Six species of the series occur in our area: FE. equisetoides (EIl.)
Torrey and FE. quadrangulata (Michx.) Roemer & Schultes are reported
throughout the Southeast and range north to southern New England; E. ce/lulosa
Torrey, E. interstincta (Vahl) Roemer & Schultes, and E. elongata Chapman
are restricted to the Coastal Plain; and &. Robbinsii Torrey, a species mainly
of the Northeast, ranges south to Virginia and northern Florida along the
Coastal Plain. Tubers of E. dulcis (Burman f.) Trin. ex Henschel, n = ca. 100,
provide the familiar water chestnut of Oriental cuisine. The juice of the tubers
is strongly antibiotic (Hegnauer). The species is closely related to the eastern
North American EF. equisetoides, and the pair serve as an example of the eastern
Asian-eastern North American pattern of disjunction (Wood).
Species of ser. PAUCIFLORAE Svenson are tiny plants with few-flowered spike-
lets. Eleocharis parvula (Roemer & Schultes) Link, 2” = 8, 10, E. rostellata
Torrey, and E. melanocarpa Torrey occur in the Southeast, and all have broad
ranges in our area.
386 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 68
io)
sue
RUWORITESISTNU ANI —
2 a
= = ie. ‘
7 MATT WT. h i)
IND ei dt mm
a
FiGure 2. Eleocharis. a-k, E. cellulosa: a, habit of stoloniferous plants, x 14; b, cross
section of culm, showing air spaces (black) with cross partitions (stippled—cellular detail
too small to be shown), x 10; c, detail of culm with apex of bladeless sheath, x 2; d,
spike of flowers in carpellate phase (flowers protogynous), styles protruding (note flowers
with either 2 or 3 stigmas), x 2; e, abaxial side of flower, stigmas receptive, filaments
1987] TUCKER, CYPERACEAE 387
Plants of ser. ACICULARES Svenson are also small. Three species occur in our
area, and their contrasting distribution patterns are notable. The northern
Eleocharis Wolfii Gray is found southward to Tennessee and Louisiana, while
the neotropical F. radicans (Poiret) Kunth ranges northward to Virginia and
Oklahoma. However, E. acicularis (L.) Roemer & Schultes, 2n = 20, 30-38,
50-58, a widespread north-temperate species, is reported from throughout
eastern North America. Both emergent and submersed growth forms of £.
acicularis a been described. Submersed plants have three large lacunae per
culm, while emergent plants have about ten small ones. These forms are ge-
netically ee and fully interconvertible, as is demonstrated by reciprocal
transplants (Rothrock & Wagner). The plants are able to grow in acidic runoff
from Appalachian coal mines and flourish in streams with pH as low as 2.8.
This is odd and suggests some overlooked variability in the species, because
in northern Europe it nearly always occurs in basic waters (Iversen).
The plants of ser. OvATAE Svenson have broadly ellipsoid to ovoid spikelets.
Three species are in our area: Eleocharis obtusa (Willd.) Schultes, 21 = 10, in
every Southeastern State, is one of the commonest spike-rushes in eastern
North America; the closely related £. Enge/mannii Steudel, 2n = 10, occurs
from Georgia and Missouri south to the Gulf Coast; and £. /anceolata Fern.
is a oe species that just extends into our area in Arkansas and
Louisi
mane af ser. MACULOSAE Svenson are characterized by dark purple to black,
biconvex achenes. Some species grow submersed, while others are found in
littoral habitats. There are four species in our area: Eleocharis caribbaea (Rottb.)
Blake is pantropic (northward to South Carolina and Texas); E£. olivacea Torrey,
2n = 20, is endemic to the Coastal Plain from Virginia to Florida; £. atro-
purpurea (Retz.) Kunth is widely but sporadically distributed in the Southeast
(but otherwise is found throughout temperate and tropical regions of both the
Old and New Worlds); and £. flavescens (Poiret) Urban is neotropical, growing
north along the Coastal Plain to Delaware.
Plants of ser. WEBSTERIA (S. H. Wright) G. Tucker® are submersed, flaccid,
6Eleocharis ser. WEBSTERIA, comb. nov., based on Websteria 8. H. Wright, Bull. Torrey Bot. Club
14: 135. 1887.
f stamens not yet elongated, 4 of 6 perianth bristles visible, x 10; f, apex of spikelet,
carpellate phase past, lower flowers with protruding stamens, * 5; g, flower in staminate
phase with apex of subtending scale, x 10; h, abaxial view of flower in staminate phase
(note 4 of 6 perianth bristles, ovary with enlarged stylar base), x 10; 1, adaxial view of
mature mee with Saat stylar base and smooth perianth bristles, achene lenticular,
2 ature achene, abaxial view, x 12; k, longitudinal cross section of achene
iiiace pericarp en coat, and basal emibrye unshaded, endosperm stippled), - x a
I-n, E. obtusa: 1, spikelet with mature achenes (hidden by subtending scales, few
visible at eee left), x 5:m, abaxial side of mature achene crowned by tubercle eal
style base) and with perianth bristles, x 12; n, detail of perianth bristle to show retrorse
barbs, x 25. 0, p, E. tuberculosa: 0, abaxial side of mature ee (trigonous in cross
section) a eae x 12; p, detail of perianth bristle, x 25.q, E. atropurpurea: abaxial
side of mature achene (lenticular in cross section) with tubercle, eet bristles absent,
5
388 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
slender-branched plants of shallow, still waters. There are one or perhaps two
pantropic species (Eiten, 1976b). Eleocharis confervoides’ is an uncommon
plant of cypress swamps and lakes in Florida, southern Alabama, southern
Georgia, and Louisiana. It is also known from widely scattered localities in the
neotropics and in tropical Africa and Sri Lanka. The species has been variously
placed in Rhynchospora Vahl (Bentham: Kiikenthal, 1948), Scirpus, and the
monotypic Websteria. The slender, leafless culms are similar to those of other
species of Eleocharis. Additional submersed one-flowered species of Eleocharis
occur in Brazil and Africa (Nelmes). At anthesis, the one-flowered spikelets of
E.. confervoides are exserted just above the water surface. The achenes lack the
differentiated tubercle of most species of Eleocharis, but the embryos are typical
of the genus (Van der Veken).
Rikli reported that the inner parenchymatous layer was absent from the
bundle sheaths in many species of E/eocharis, a feature on which he based the
segregate genus Chlorocharis. Metcalfe could not confirm this in any species
including those investigated by Rikli but suggested that further study might be
profitable.
Chromosomes of Eleocharis have been extensively studied. Cytologically,
the genus is the best known in the Cyperaceae. Most species have the diffuse
centric condition typical of the family; some have holocentric chromosomes
(Battaglia). Although aneuploidy has been frequent in most other genera of
Cyperaceae, polyploidy has been important in the evolution of this genus.
Several species have tetraploid (and sometimes hexaploid) races or subspecies.
Strandhede (1965, 1966) studied about 1100 European populations of species
of ser. ELEocHaRIS (ser. Pa/ustriformes) and reported that chromosome break-
age and refusion were common. Most species had several cytotypes, and various
kinds of multivalents were frequent at meiosis. Heterovalents formed in mei-
osis, and aberrant but apparently viable gametes were often observed. Similar
reports of chromosomal variability have been made for North American species.
Karyotypic rearrangements have been noted in Eleocharis flavescens (Poiret)
Lam., which had 30 chromosomes in various combinations of univalents,
bivalents, tetravalents, and ring complexes (Schuyler, 1977).
When the sample size is large, chromosome number can be correlated with
morphology within species and between species pairs. For example, the Eu-
ropean Eleocharis uniglumis (Link) Schultes consists of two subspecies that
differ in ecology and in features of the spikelet scales. Subspecies unig/umis
has n = 46, while subsp. Sterneri Strandhede has n = 74-82. Apparently, the
latter taxon was derived from the former by tetraploidy followed by fusion of
some of the chromosomes, but fusion of different chromosomes in different
populations has also resulted in mixoploidy. In some cases affinities between
species can be confirmed cytologically. For example, E. Engel/mannii Steudel
and £. obtusa are both n = 5 and have very similar arrotvees.
Species with different chromosome numbers are known to hybridize in the
wild. Some hybrids (e.g., Eleocharis mamillata x E. palustris subsp. palustris)
"Eleocharis confervoides eee . Tucker, comb. nov., based on Scirpus confervoides Poiret in
Lam. Encycl. Méth. Bot. 6: 75
1987] TUCKER, CYPERACEAE 389
have greatly reduced fertility, while others (e.g., E. palustris subsp. palustris x
subsp. vulgaris) have fertility comparable to that of the parent species.
Several species are important weeds, especially of rice fields.
REFERENCES:
Under family references see BARROS (1928); BATTAGLIA; BEAL; BENTHAM; BERGGREN;
BLASER (1940, 1941a); BREWBAKER; CLARKE (1908, 1909); Erren (1976a, 1976b); EYLEs
& ROBERTSON; FAsseTT; GODFREY & WOOTEN; GONCHAROV ef al.; Goon et al.; HA-
KANSSON (1954, ee HARBORNE et a/.; HARRIS & MARSHALL, Heowsre: HESLA et
al., Heusser; Hoim et al.; seasons HotcuHkiss; Huanc; G. E. HUTCHINSON;
J. Hutrcuinson; F. D. a - H. Kern; KuNTH; LE MAout ‘& DEcAISNE; MCATEE;
METCALFE; NAPPER (1965); NEES VON pees OGDEN; O’NEILL; PATCH; RIKLI;
SCHULZE-MOTEL (1959, 1964); em et al.; STACE; STANDLEY; TIETZ; TORREY; and VAN
DER VEKEN
BERNARDINI, J. V. Studies of the kinetochore of Eleocharis macrostachya Britt. Proc.
Minnesota Acad. Sci. 27: 104-114. 1959.
Boyp, C. E., & D. H. Vickers. Relationships between production, nutrient accumu-
lation, and chlorophyll ie in an Eleocharis quadrangulata population. Canad.
Jour. Bot. 49: 883-888.
Bricas, B. G. Become numbers in some Australian ae of Eleocharis (Cy-
peraceae). Contr. Natl. Herb. New South Wales 4: 130-136. 1970. [Summary of all
reported chromosome numbers arranged in series; E. ee (L.) Roemer &
Schultes, 2” = 20.
BRUNNER, G. Aquarium plants. (English translation by G. Vevers.) [vil] + 94
Princeton and New York. 1966. [E. acicularis, E. vivipara Link, 23, 24, propagated
by division and runners.]
Evans, P. S. Intercalary growth in the aerial shoot of Eleocharis acuta R. Br. I. Structure
of the growing zone. Ann. Bot. 79: 205-217. 1965.
Harms, L. J. esa studies in ye subser. Pal/ustres: central United
States taxa. Am. Jou t. 55: 966-974. 196
. Cytotaxonomy of ie ree ae complex. [bid. 59: 483-487. 1972.
Horn AF RAntzien, H. Certain aquatic plants collected by Dr. J. T. Baldwin Jr. in
Liberia and the Gold Coast. Bot. Not. 1951: 384-398. 1952. [Subg. RHEOCHARIS
Horn described and illustrated. ]
IversEN, J. Studien iiber die pH-Verhaltnisse danische Gewdsser und ihren Einfluss auf
die Hydrophyten-Vegetation. Bot. Tidsskr. 40: 277-326. 1929. [Distribution of EF
acicularis f. submersa, 95 percent of its occurrences in Denmark are in neutral,
alkaline, or variable waters, only 5 percent in acidic waters; cf. ROTHROCK & WAGNER. |
KUKENTHAL, G. Vorarbeiten zu einer Monographie der Rhynchosporoideae. Rhyn-
chospora. Bot. Jahrb. 74: 375-509. 1949; Ibid. 75: 90-195. 1950; Ibid. 75: 273-
314. 1951.
LEwIs, K. R., & B. JOHN. ee ae in a wild population of Eleocharis palustris.
Chromosoma 12: 433-468.
Nemes, E. Submersed species ie ae haris with |-flowered spikelets. Kew Bull. 1952:
289, 290. 1952. [E. Naumanniana Bock. and FE. Caillei Hutchinson & Dalz. of
Pocan, E. Studies in Eleocharis R. Br. I. Chromosome numbers of E. palustris (L.) R.
et S. and E. uniglumis fous Schult. Acta Biol. Cracov. Bot. 15: 69-76. 1972. [E.
palustris subsp. palustri. a = 16; E. palustris subsp. vulgaris Walters, 2n = 38,
39, 40; E. uniglumis, =
REJMANEK, M., & J. VELAZQUEZ. ie mmunities of emerged fishpond shores and bottoms.
Pp. 206-21 1 in D. DyKyjova & J. Kvet, eds., Pond littoral ecosystems. (Ecological
390 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
studies 28.) New York. 1978. [Maximum standing crop of E. acicularis is 469-644
g/m?.]
Roturock, P. E., & R. H. WAGNER. The autecology ofan acid tolerant sedge, Eleocharis
actcularis (L.) R & S. Castanea al; 279-290. 1976
SCHUYLER, A. E
Rien Brittonia 29: 129-133. 1977.
. FerReN, JR. A new intertidal form of Eleocharis olivacea (Cyperaceae).
Bartonia 43: 46-48. 1975
SEISCHAB, F. K., J. M. BERNARD ,& K. Fiata. Above- and belowground standing crop
partitioning of biomass by Eleocharis rostellata in the Byron-Bergen Swamp, Ge-
nesee County, New York. Am. Midl. Nat. 114: 70-76. 1985.
STRANDHEDE, S. O. Chromosome studies in Eleocharis ae Palustres. II. Obser-
vations on western European taxa. Op. Bot. 9(2): l- 1965.
Eleocharis subser. Palustres in North een el comments and
chromosome numbers. Bot. Not. 120: 355-368. 1967.
SveNson, H. K. Monographic studies in the genus Eleocharis. Rhodora 31: 121-135,
152-163, 167-191, 199-219, 224-242. 1929. [The basic monograph, worldwide:
keys, a distribution maps, synonymices. ]
onographic studies in the genus Eleocharis—IIL. Ibid. 34: 193-203, 215-227.
pl. 221. 1932; IN. Ibid. 35: 377-389. pls. 320, 321. 1935; IV. Ibid. 39: 210-273.
pls. 460-465. 1937; V. Ibid. 41: 1-77, 93-110. pls. 537-547.
. The group of Eleocharis palustris in North America. Rhodora ne 61-67. 1947.
. Eleocharis. N. Am. FI. 18: 509-540. 1957. [Keys, descriptions.]
Wa ters, 8S. M. Eleocharis. In: Biological flora of the British Isles. Jour. Ecol. 37: 192-
206. 1949.
tern North American E/eocharis
——.. On the vegetative morphology of Eleocharis R. Br. New Phytol. 49: 1-7. 1950.
Warp, D. B., & E. M. H. LeiGu. aes tions to the flora of Florida—8, Eleocharis
(Cyperaceae). Castanea 40: 16-36.
Woop, C. E., JR. Morphology and aa. the classical approach to the study
of disjunctions. Ann. Missouri Bot. Gard. 59: 107-124. 1972. [Eastern North Amer-
ican—Eastern Asian disjunctions; Dulichium arundinaceum, extant and fossil distri-
bution, 118.]
WriGut, S. H. A new genus in Cyperaceae. Bull. Torrey Bot. Club 14: 1887. [Websteria.]
5. Fimbristylis Vahl, Enum. Pl. 2: 285. 1805.
Small to medium-sized annuals or perennials of disturbed, open, wet habitats.
Roots fibrous; rhizomes regularly present in some species. Culms slender, terete
or nearly so, glabrous. Leaves all basal; sheaths smooth or pubescent, with
ligule present or not, glabrous or ciliate; blades linear to filiform, flat, condu-
plicate, or involute, glabrous or pubescent, the margins glabrous or scabrellate:
chlorenchyma radiate; bundle sheaths 3-layered (““Fimbristylis type”). Inflo-
rescences terminal, branched (rarely sessile, capitate); bracts 1-6, erect to oblique,
the sheaths greatly reduced to essentially absent, the blades leaflike; primary
rays absent or 1-10, glabrous or scabrellate, secondary rays regularly produced
in some species. Spikelets single or in clusters of 2-5, ovoid to lanceolate. Scales
5-100, ovate to oblong, obtuse or acute, blunt or mucronate [aristate], glabrous
or puberulent abaxially, 1- to 5-nerved medially, nerveless laterally, deciduous
at maturity. Flowers perfect. Perianth lacking. Stamens (1, 2, or) 3; filaments
about as long as the subtending scales, flattened; anthers oblong, the apices of
the connectives sometimes prolonged; pollen grains uniaperturate, obovoid,
1987] TUCKER, CYPERACEAE 391
subspheroidal, or spheroidal, scabrate, trinucleate. Styles slender, terete
throughout or trigonous basally, usually fimbriate distally, deciduous from the
mature achene; stigmas 2 (or 3), about as long as the style, glabrous. Achenes
lenticular or trigonous, ovoid, oblong, or obovoid, the apex broadly rounded
to subacute, apiculate or not, the base cuneate or stipitate, the surface smooth,
warty, or reticulate with isodiametric or horizontally arranged rectangular cells,
these cells concave or with a central papilla. Embryos turbinate, radicle lateral,
coleoptile basal. Base chromosome number 5. Type species: F. dichotoma (L.)
Vahl, typ. cons. (Name from Latin fimbria, fringe, and stylus, style, referring
to the fringed style of most species.)
A genus of about 200 species, mainly pantropic but also well represented in
warm-temperate regions. Most of the species grow in disturbed wet habitats,
especially roadsides and croplands. The center of diversity is southeastern Asia
(Goetghebeur & Coudijzer). Thirteen species are recorded from the United
States. Twelve of these occur in the Southeast, while Fimbristylis thermalis S.
Watson is endemic to California, Arizona, and Nevada (Kral). Kral’s thorough
monograph includes illustrations and chromosome counts for all species in
North America.
Fimbristylis is closely related to Bulbostylis and Abildgaardia. Chromosome
numbers in the three genera are based on five (Gordon-Gray, Kral), and their
kranz anatomy is similar (three-layered bundle sheaths). Such anatomy is not
reported in any other genera of the Cyperaceae (Metcalfe; Raynal, 1972). The
three genera have been distinguished from the remainder of the Scirpeae as
tribe Abildgaardiae Lye (Fimbristylideae Raynal).
Koyama (1961) treated Bulbostylis as a subgenus of Fimbristylis, while Kral
recognized three genera, Bulbostylis, Abildgaardia, and Fimbristylis. Additional
information supports Kral’s belief. Gordon-Gray made a careful study of the
southern African representatives of the three genera. Abi/dgaardia can be dis-
tinguished from Bulbostylis and Fimbristylis by its distichous spikelet scales.
Bulbostylis and Fimbristylis are separated by a suite of characters. The embryos
are consistently different (in Fimbristylis the radicle is lateral, the coleoptile
basal; in Bulbostylis, the radicle is basal and the coleoptile lateral), although
there is no single morphological character that separates the two genera. The
styles of Fimbristylis are usually fimbriate (occasionally entire) and are decid-
uous, while those of Bu/hostylis are always entire and have a persistent base.
The spikelet scales of Fimbristylis are generally glabrous, while those of Bul-
bostylis are generally puberulent. The ligules of Fimbristylis are glabrous, while
those of Bulbostylis are hispid. Species of Fimbristylis always lack intrapro-
phyllar buds at the base of the inflorescence rays, while such buds are frequently
present in Bulbostylis (Guaglianone). Species of the two genera differ in surface
ornamentation of the achenes. Goetghebeur & Coudijzer examined about 100
species from throughout the world and found that the epidermal cells of Fim-
bristylis are horizontally elongate (infrequently isodiametric) and in vertical
bands, but those of Bulbostylis are vertically elongate in horizontal bands. The
two genera also differ in habit and habitat: Fimbristylis species are mostly
perennials of moist soils, while Bu/bostylis species are generally annuals of dry
sandy soils.
392 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Svenson recognized two sections in Fimbristylis, Kral did not comment on
the infrageneric classification. Plants of sect. FiMBRISTYLIS (sect. Dichelostylis
Bentham) have two stigmas, lenticular achenes, and styles commonly fringed
apically. This section includes eleven of the fourteen species of the southeastern
United States. Most of our species are somewhat weedy plants of disturbed
wet habitats: Fimbristylis tomentosa Vahl, n = 5; F. dichotoma (L.) Vahl, n =
10, 15; F. decipiens Kral, n = 10; F. annua (All.) Roemer & Schultes, » = 15;
F. Vahlii (Lam.) Link, n = 10; F. puberula (Michx.) Vahl, n = 10, 20; and F.
perpusilla Harper, n = 5. In general these are widely distributed in the Southeast.
Fimbristylis perpusilla, endemic to southeastern North America, is a notable
exception. Kral knew of only two localities in southwestern Georgia for this
tiny annual. Recently, the species has been reported in Horry County, South
Carolina (Leonard) and in eastern Maryland (Schuyler, pers. comm.). The four
remaining southeastern species of sect. FimBristyLis, F. caroliniana (Lam.)
Fern. (” = 10, 20, 30), F. schoenoides (Retz.) Vahl (n = 5), F. spathacea Roth
(n = 24), and F. castanea (Michx.) Vahl (7 = 10), are tall plants of tidal marshes.
Plants of sect. TRICHELOSTYLIS Bentham have three stigmas, lenticular achenes,
and entire styles. In our area this section is represented by Fimbristylis autum-
nalis (L.) Roemer & Schultes, F. comp/anata (Retz.) Link, and F. miliacea (L.)
Vahl, all n =
Fimbristylis autumnalis and F. miliacea are detrimental weeds in rice fields
in the Southeast and California (Smith ef a/.), as well as in Asia and Africa
(Holm et al.). Fimbristylis tomentosa 1s rapidly becoming a common weed in
rice fields from South Carolina to Texas (Kral).
REFERENCES:
Under family references see BADEN ef al.; BARROS (1945); BEAL; BENTHAM; BREWBAKER;
Brown; CAROLIN et al/.; CLARKE (1908, 1909): eres GODFREY & WOOTEN; GONCHA-
ROV et al; HARBORNE; HARBORNE et al.; HOLM et al.; HoLttUM; HUANG: J. HUTCHINSON:
J. H. KERN; KOYAMA (1961); KUKKONEN (1969): ce Le Maout & DECAISNE; LER-
MAN & RAYNAL; METCALFE; NApPER (1965); NEES VON ESENBECK; O’ NEILL; PATCH; RAYNAL
(1972, 1973, 1978); Riku, SCHULZE-MoTEL (1959, 1964); SmitH et al.; STANDLEY; TEERI
et al., Torrey; and VAN DER VEKEN
GoOETGHEBEUR, P., & J. Coupuzer. Studies in Cyperaceae 3. Fimbristylis and Abild-
gaardia in Central — Bull. Jard. Bot. Natl. Belg. 54: 65-89. 1984. [SEM pho-
tographs of achene
GORDON-GRAY, K. D. i Tmbristylis ae oo hers sae as seen by a student
of southern African species. Mitt. Bot. Staatssam. Miinc 10: 549-574. 1971.
GUAGLIANONE, E. R. Un nuevo ee er en la distincion generic entre Fimbristylis
Vahl y Bulbostylis Kunth (Cyperaceae). Darwiniana 16: 4 70.
Hom, T. Studies in the Cyperaceae. X. Fimbristylis Vahl; an anatomical treatise of
North American species. Am. Jour. Sci. 157: 435-4 899.
KrAL, R. A treatment of . ee Bulbostylis, and Fimbristylis (Cyperaceae) for
North America. Sida 4: 57-227. 1971
LEONARD, a - Fimbristylis a Paper in South Carolina. Castanea 46: 235,
236.
SVENSON, 7 2 eee ee and Abildgaardia (Cyperaceae: Scirpeae). N.
Am. Fl. 18: 540-556.
1987] TUCKER, CYPERACEAE 393
Warp, D. B. Contributions to a flora of Florida, 4. Fimbristylis (Cyperaceae). Castanea
33: 123-134. 1968a.
—. Supplemental note to Fimbristylis of Florida. [bid. 350. 1968b.
6. Bulbostylis Kunth ex C. B. Clarke in Hooker f. Fl. Brit. India 6: 651. 1983,
nom. cons.
Small to medium-sized, tufted (solitary-stemmed) perennials or annuals of
open or disturbed, dry or wet habitats. Roots fibrous; rhizomes lacking [present].
Culms slender, terete, glabrous. Leaves all basal; sheaths expanded basally or
not, with ligule fimbriate or ciliate apically; blades filiform or narrowly linear,
shorter than to slightly exceeding the culm, conduplicate or involute, often
pubescent on one or both surfaces, the margins and midvein scabrellate or
smooth; chlorenchyma radiate; bundle sheaths 3-layered (““Fimbristylis type”’).
Inflorescences terminal, capitate or branched; bracts 1-4, erect to oblique,
shorter than to exceeding the length of the rays; primary rays lacking or 1-6,
erect or spreading, subterete, glabrous or scabrellate, secondary rays absent.
Spikelets solitary or in small clusters, ovoid to oblong or lanceolate. Scales 2-
50, ovate to oblong, mucronulate, mucronate, or aristate, glabrous or scabrel-
late, or puberulent abaxially, 3- to 7-nerved, deciduous at maturity, the 1-4
lowest ones sterile. Flowers perfect. Perianth lacking. Stamens (1, 2, or) 3;
filaments slender, hyaline, about as long as the subtending scales; anthers ob-
long, the apices of the connectives prolonged as tiny subulate tips; pollen grains
uniaperturate, subspheroidal or obovoid, psilate or scabrate, trinucleate. Styles
papillate, the bulbous basal portion persistent on the mature achene; stigmas
3, slender, glabrous, equaling to exceeding the style in length. Achenes trigonous
(rarely biconvex), ovoid to oblong or ellipsoid, the apex obtuse to acute, crowned
by the persistent bulbous style base, the base cuneate to stipitate, the surface
smooth or reticulate with vertically elongate, rectangular (rarely isodiametric)
cells, these cells smooth or sometimes with a single central papilla. Embryos
turbinate, radicle basal, coleoptile lateral. Base chromosome number 5. TyPE
species: B. capillaris (L.) C. B. Clarke, typ. cons. (Name from Latin bulbus,
bulbous, and stylus, style, referring to the characteristic bulbous style base.)
A genus of about 120 species, mostly pantropic but with some in the warm-
temperate regions. The genus is related to Abildgaardia and Fimbristylis. (A
discussion of the distinguishing features of these genera appears under Fim-
bristylis.) Bulbostylis was first distinguished from Fimbristylis as the genus
Stenophyllus Raf. (Neogenyton, 4. 1828). Although the generic name Bulbo-
stylis Kunth was published in synonymy (Kunth) and validated by Clarke (q.v.),
it has been conserved over Stenophyllus. Kral’s illustrated monograph (in-
cluding chromosome numbers) is the basic reference for the North American
species
Bulbostylis is represented in the United States by eight species, me of which
occur in the Southeast. Bulbostylis barbata (Rottb.) C. B. Clarke, n = 5,
ae (L.) C. B. Clarke, n = 36, and B. ciliatifolia (Ell) Fern., n = in have
ach been reported from all or nearly all the southeastern states. Bulbostylis
eis (Ell.) C. B. Clarke and B. Warei (Torrey) C. B. Clarke, both n =
394 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
15, are more restricted in range than the three preceding species. Both occur
along the Coastal Plain from Florida to North Carolina. Three more species,
B. Funckii (Steudel) C. B. Clarke, n = 10, B. juncoides (Vahl) Kiikenthal, n =
60, and B. Schaffneri (Boeck.) C. B. Clarke, occur in the Southwest. About 15
species occur in Mexico, Central America, and the West Indies, with perhaps
20 in all of South America. The center of diversity for the genus is tropical
Africa, where 30-40 species are reported.
The southeastern species of Bulbostylis are generally found in open, dry,
sandy places, such as pine flatwoods, sand hills, palmetto scrub, roadsides, and
shores. They are annuals or short-lived perennials. The neotropical B. paradoxa
(Sprengel) Lindm., a long-lived perennial that flowers in response to fires (Kral),
occurs in pinelands and savannas in Cuba and from Mexico to northern South
America.
Plants with basal clusters of spikelets are occasionally encountered in several
species of Bulbostylis (e.g., B. capillaris and B. Funckii). Formation of such
spikelets may be the result of drought, but no studies have been made to
document this supposition. In some species achenes produced by the basal
spikelets are 12-2 times larger than those produced by typical elongate culms.
Such amphicarpy has also been reported in certain African species (Haines).
Bulbostylis barbata is a weed of old fields and sandy croplands in the south-
eastern Coastal Plain. Three species (including B. barbata) are reported as
significant weeds in tropical Africa and Asia (Holm et al.).
REFERENCES:
er family references see eae (1945); BEAL; BENTHAM: BROWN; CAROLIN et al.:
epee (1908, 1909); Goprrey & WoorTeNn; GONCHAROV ef al.: HAINES; HARBoRNe:
HARBORNE et al/.; HOLM et ale oreu Huana; J. HuTCHINSON; J. H. KERN; KUKKONEN
(1969); KUNTH; LE MAout & DecAIsSNE; LERMAN & RAYNAL: METCALFE: NAPPER (1965);
NEEs VON EseNBECK; O'NEILL; RAYNAL (1972, 1973, 1978); Riki; SCHULZE-MOTEL (1959,
1964); STANDLEY; TEER! et a/.; TORREY; VAN DER VEKEN: and WINFREY & SAMSEL.
Under Fimbristylis see GORDON-GRAY, KRAL, and SVENSON.
GOETGHEBEUR, P. Studies in Cyperaceae 4. New species and a new combination in
Cen tral African Bulbostylis. Bull. Jard. Bot. Natl. Belg. 54: 91-104. 1984.
U, e systematic anatomy ne a Indian Cyperaceae: Bulbostylis
Kunth. Jour. Linn. Soc. Bot. 59: 289-304.
Lye, K. A. The generic concept of Bulbostylis en ex C. B. Cl. Mitt. Bot. Staatssam.
Miinchen 10: 539-547. 1971,
7. Abildgaardia Vahl, Enum. Pl. 2: 296. 1805.
Small, single-stemmed or tufted, bulbous-based, glabrous perennials of trop-
ical and subtropical grasslands. Roots fibrous: rhizomes lacking. Culms sub-
terete, smooth. Leaves about ' as long as the culms; sheaths expanded, their
overlapping bases forming the bulblike base of the plant, ligules lacking; blades
linear-filiform, slightly involute, thickened at margins, scabrellate distally;
chlorenchyma radiate; bundle sheaths 3-layered (‘“Fimbristylis type’). Inflo-
rescences simple cymes of 1-3[-6] sessile or pedunculate spikelets: bracts sol-
1987] TUCKER, CYPERACEAE 395
itary, filiform. Spikelets broadly lanceolate, slightly compressed, the scales
distichous or essentially so. Scales 3-15, ovate, acute, mucronate, 3- to
5-nerved medially, nerveless laterally, deciduous as the achenes mature. Flow-
ers perfect (although frequently the distal flowers of a spikelet staminate only).
Perianth lacking. Stamens (1, 2, or) 3; filaments flattened; anthers linear, the
apices of the connectives not prolonged; pollen grains uniaperturate, obovoid
to subspheroidal, scabrate, trinucleate. Style trigonous basally, slender and
capillary distally, deciduous from the mature achene; stigmas 3, linear, about
as long as the style, glabrous. Achenes rounded-trigonous, ovoid, the apex
broadly rounded, apiculate, the base abruptly contracted to a stipe, the surface
pebbled. Embryo turbinate, radicle basal. Base chromosome number 10. (Named
for P. S. Abildgaard, an eighteenth-century Danish botanist.) TyPeE SPECIES: A.
ovata (Burman f.) Kral (Carex ovata Burman f.; A. monostachya (L.) Vahl);
see Britton & Millspaugh, Bahama FI. 52. 1920.
A pantropic genus of about 15 species, distinguished from Bulbostylis and
Fimbristylis, with which it has been united, by its distichous spikelet scales
and its deciduous style bases. Chemical data support the recognition of Abild-
gaardia. The four Australian species produce the flavones luteolin and tricin,
whereas the 15 species of Fimbristylis and Bulbostylis examined had only tricin
(Harborne ef al.).
Abildgaardia is represented in the New World by two species. Abildgaardia
mexicana (Palla) Kral, n = 10, is endemic to grasslands of the Mexican High
Plateau. The southeastern representative, 4. ovata, n = 10, occurs in Florida,
the West Indies, and the lowlands of Central and South America. Abildgaardia
ovata is found in grasslands over limestone in southern Florida (Dade and
Monroe counties) and in the vicinity of Tampa (Citrus County; Kral).
Species of Abildgaardia have no reported economic significance. None has
been noted as a weed.
REFERENCES:
Under family references see BARROS (1945); BENTHAM; CLARKE (1908); GODFREY &
WooTEN; HARBORNE; HARBORNE et al.; HUANG; KUNTH; LERMAN & RAY NAL; METCALFE;
NAPPER aay NEES VON ESENBECK; O’ NEILL; SCHULZE-MOTEL (1959, 1964); and VAN
DER VEKE
Under Fimbristylis see KRAL and SVENSON.
Lye, K. A. Studies in African Cyperaceae VIII. The taxonomic position of Abildgaardia
Vahl and Nemum Hamilton. Bot. Not. 126: 325-329. 1973.
Tribe CyPEREAE
8. Cyperus Linnaeus, Sp. Pl. 1: 44. 1753; Gen. Pl. ed. 5. 27. 1754.
Tufted or rhizomatous, perennial or less often annual herbs of disturbed wet
to dry soils, marshes, ditches, shallow swamps, and shores in full sun or light
shade. Roots fibrous; rhizomes or stolons sometimes present, horizontal to
oblique. Culms trigonous (sometimes with winged angles) or terete, smooth or
scabrellate. Leaves all basal; sheaths glabrous, sometimes with conspicuous
396 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
cross veins, especially in emergent plants, ligule present or lacking; blades linear
to lanceolate, flat, conduplicate, plicate, filiform, crescentiform, or involute,
the margins and midvein usually scabrellate; stomata paracytic, sometimes
surrounded by 1-4 papillae; chlorenchyma radiate or not (if radiate the bundle
sheaths 2-layered—“*Cyperus type’’). Inflorescences terminal, diffusely branched,
spicate, or capitate; bracts (1-)3-6(-22), the sheaths very short, the blades
leaflike, closely spaced and appearing verticillate at the apex of the culm, usually
ascendent but in some species erect (the inflorescence thus appearing lateral),
horizontal, or reflexed, forming a conspicuous involucre; rays glabrous (rarely
scabrellate or hispidulous), unequal in length, produced singly from the axils
of the inflorescence bracts; spikes digitate, glomerulate, or spicate; rachis smooth,
rarely scabrellate. Spikelets (1—)5—30(-150), cylindrical to compressed, ovate,
lanceolate, or linear, the scales distichous; rachilla deciduous or persistent,
internodes winged or wingless, spongy and thickened in a few species. Scales
(1 or) 2—20(-80), oblong, elliptic, or ovate, obtuse, acute, mucronulate, or
cuspidate, 3- to | 1-nerved, deciduous or persistent, the 2 lowermost (bract and
prophyll) sterile. Flowers perfect [imperfect, the plants dioecious]. Perianth
lacking. Stamens (1, 2, or) 3; filaments ribbonlike, usually as long as the sub-
tending scales; anthers ovoid, ellipsoid, or linear, the apices of the connectives
sometimes prolonged as small, reddish, entire or scabrellate appendages; pollen
grains obovoid, subspheroidal, rectangular, or triangular, (1- or) 4-aperturate,
psilate, trinucleate. Styles slender, the base sometimes persistent as an apiculus
or beak on the mature achene; stigmas capillary, shorter than, equaling, or
exceeding the style in length, glabrous [glandular]. Achenes trigonous or len-
ticular, ovoid, ellipsoid, or narrowly oblong, obtuse or acute, apiculate or not,
stipitate, substipitate, or sessile, smooth, puncticulate, or reticulate. Embryos
broadly to narrowly ellipsoid. Base chromosome number 8. (Incl. Pycreus
Beauv., Mariscus Vahl, Juncellus (Griseb.) C. B. Clarke, Acorellus Palla, Re-
mirea Aublet, Torulinium C. B. Clarke.) Lecrotyre species: C. esculentus L.:
see Britton & Brown, Illus. Fl. No. U.S. Canada, ed. 2. 1: 297. 1913. (Name
from Greek kupeiros, ancient name for C. /ongus L.)—FLAT-SEDGE, UMBREL-
LA-SEDGE, SEDGE-GRASS, GALINGALE (Britain).
A very large genus of about 650 species widely distributed throughout the
tropical and warm- and cool-temperate regions of the world. It is the second
largest genus of the Cyperaceae; only Carex L. is larger. Cyperus is morpho-
logically coherent and is readily recognized by the distichous arrangement of
scales on the spikelets. Six subgenera have been recognized: subg. Cyperus,
subg. Pycnostacuys C. B. Clarke,* subg. Pycreus (Beauv.) Gray,’ subg.
JUNCELLUS (Griseb.) Kiikenthal, subg. ToruLiniuM (Desv.) Kiikenthal, and
subg. Fimpricyperus K. A. Lye. These are circumscribed by features of the
achenes, spikelets, and vegetative anatomy. Most recent workers have followed
*Cyperus subg. Pycnostacuys C. B. Clarke in Hooker f. Fl. Brit. India 6: 597, 1893. Lectotype
SPECIES (here designated): C. diffusus Vahl. Synonym: Cyperus subg. Protocyperus K. A. Lye, Nordic
Jour. Bot. 1: 54. 1981. Type species: C. difformis L.
°Cyperus subg. Pycreus (Beauy.) Gray, Man. Bot. ed. 1.517. 1848. This combination is consistently,
but erroneously, attributed to C. B. Clarke, Jour. Linn. Soc. Bot. 21: 33. 1884.
1987] TUCKER, CYPERACEAE 397)
Kiikenthal and Fernald, who treated the genus in the broad sense. Others
(Koyama, 1962b; Vorster; Raynal, 1972, 1973) have followed Clarke (1908)
and recognized the subgenera as genera. Subgenera Pycreus and JUNCELLUS
differ from the others in having the derived conditions of lenticular (vs. trig-
onous) achenes and bifid (vs. trifid) styles (Blaser, 1941a; Raynal, 1972). In
subg. Pycreus the achenes are laterally compressed, while in subg. JUNCELLUS
the compression is dorsiventral, suggesting that the bicarpellate condition
evolved twice. Several other genera of the family (e.g., Carex and Bulbostylis)
are divided into subgenera on the basis of carpel number.
Subgenus ToRULINIUM differs from all other subgenera in having the rachilla
articulate at the base of each scale (i.e., an abscission layer forms) (vs. contin-
uous or articulate only at the base of the spikelet). Thus, the mature spikelet
of plants of subg. TORULINIUM breaks up into one-fruited segments, each con-
sisting of an internode of the rachilla, a scale, and an achene.
Subgenera JUNCELLUS, Pycreus, and ToRULINIUM are readily distinguished
from each other and from the remaining subgenera. However, the subgeneric
classification of the remaining species of the genus has been a matter of long
debate. Traditionally, the species here recognized as constituting subgenera
PycnostAcHys and Cyperus (Lye, 1981) have been circumscribed differently
as subgenera Mariscus and Cyperus. Clarke (1908) and Kikenthal (1935-
1936) defined subg. Cyperus as differing from subg. Mariscus in having the
spikelet rachilla firmly attached to the rachis, while the scales are deciduous,
falling from the rachilla as the achenes mature. In species of subg. Mariscus,
the scales remain firmly attached to the rachilla even after the spikelet has
fallen from the rachis. O’Neill (1942) listed some twenty species (e.g., Cyperus
strigosus L., a common species throughout the United States) having charac-
teristics of both subgenera—both the rachillas and the scales are more or less
deciduous. Kiikenthal placed such intermediate species in his concept of subg.
Mariscus, but they are clearly transitional between subg. Cyperus and subg.
Mariscus. Also, as O’Neill (1942) observed, C. rotundus L. and C. esculentus
L., both of which have always been placed in subg. Cyperus, have persistent
scales, a feature attributed solely to subg. Mariscus by Kiikenthal. Federowicz
surveyed the epidermal features of leaves and achenes of both subgenera and
found no consistent differences between the two. There is no single character
that consistently separates them. O’Neill (1942, p. 47) stated: “It is ill-advised
to maintain Mariscus as a genus when it is very ill-defined even as a subgenus.”
More recently, Koyama (1962b) and Vorster have recognized Mariscus at the
generic level.
Rikli surveyed the anatomy of the leaves and culms of many genera of the
Cyperaceae. He divided Cyperus into two genera, Eucyperus (= Cyperus) and
Chlorocyperus. The latter was characterized by having radiate chlorenchyma
(i.c., kranz anatomy), while the former had nonradiate. Lerman & Raynal
examined the distribution of the C, photosynthetic pathway in the family and
found that Cyperus contained both C, and C, species. These physiological
differences were correllated with the division that Rikli had based on anatom-
ical information. Subgenus PycNosTAcuys corresponds to “Pars Pycnostachys”
(not a valid taxonomic rank) in Kiikenthal’s monograph of the genus. Lye
398 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
concurred with O’Neill that Mariscus could not be maintained even at the
subgeneric rank but ought to be included in subg. Cyperus. The recognition
of subgenera PycNosTAcHys and Cyperus (rather than subgenera Mariscus and
CYPERUS sensu Kiikenthal) is a natural classification that reflects current knowl-
edge of the phylogeny of the genus, as outlined by Raynal (1973).
Van der Veken surveyed variation in embryo shape within the subfamily,
including 162 species of Cyperus. Throughout this genus the embryos were
broadly ellipsoid. There were interspecific differences in size, but these did not
follow taxonomic lines. Van der Veken’s data supported a broad concept of
the genus.
Harborne and colleagues surveyed the distribution of flavonoids in South
American, African, and Australian species of Cyperus. They examined about
150 species and reported that each subgenus had a distinct profile of com-
pounds. Subgenus PycNnostTacuys is characterized by flavonols, which are ab-
sent in the other subgenera (these have flavones instead). Aurones, which give
a yellowish hue to the inflorescences, are present in subgenera Cyperus (in-
cluding subg. Mariscus) and ToRuLINIuM but lacking in subgenera Pycreus
and Pycnostacnys. These investigators believed the differences they reported
confirmed the recognition of PycNosTAcuys as a subgenus distinct from subg.
Cyperus. They also concluded that the flavonoid data indicated that no sub-
genus was sufficiently unlike the others to merit generic status. Thus, these
authors also favored a broad concept of the genus.
Chromosome numbers have been reported for about 40 species of Cyperus.
However, even this limited number of counts gives some information about
evolution in the genus. One significant trend is that subg. PycNosTACHyYs has
haploid numbers from 8 to 28 (mostly 15-20), while subg. Cyperus has n =
8-86 (mostly 45-60). The generally lower chromosome numbers of subg.
PYCNOSTACHYS suggest that it is the most primitive subgenus; this is also
indicated by its being the only subgenus with the C, pathway. Different chro-
mosome numbers have been reported for several species. In some species (e.g.,
C. rotundus, n = 16, 48, 54, 76) polyploid races are indicated; in others (e.g.,
C. Houghtonii Torrey, n = 84, 85, 86), mixoploid.
Cyperus in the southeastern United States comprises 63 species in four sub-
genera: five species are adventives from the Old World, seven are endemic, 17
are shared with the northeastern states, 15 are shared with the neotropics, and
the remaining ones have either pantropic or cosmopolitan distributions.
Subgenus Pycnostacnys (C, photosynthesis, spikelets in glomerules or dig-
itate clusters, achenes trigonous), with 150 species worldwide (Lye), includes
14 in our area. Eight of these belong to the New World sect. LuzEOLOIDE!
(Kunth) Clarke (spikelets in glomerulate clusters, scales with proximal abaxial
groove, stamen one per flower). The group has been revised by Denton (1978,
1983), who has also investigated the morphology of the achenes and leaf blades.
She showed that epidermal features of the achenes could be used to distinguish
species. Only one chromosome count is available for this section: Cyperus
Eragrostis Lam., 2n = 42. This species has been collected as a waif in South
Carolina; it is native to the Pacific coast of the United States and temperate
South America and is naturalized in southern Europe and southeastern Texas.
1987] TUCKER, CYPERACEAE 399
The remaining six southeastern species of the subgenus are scattered among
four sections. Section HAsPANI (Kunth) Clarke'® (wetland plants; spikelets dig-
itate; achenes ovoid, papillose), is represented in our area by three species.
Cyperus Haspan L. occurs in Coastal Plain wetlands from Virginia southward.
It is one of the few truly pantropic species and is believed to be native to
southeastern Asia, tropical Africa, and the New World tropics. Cyperus dentatus
Torrey, 2n = 34, isa northeastern species of pond shores that extends southward
to South Carolina and Tennessee. This is the only species of the subgenus with
tuberiferous stolons. It is closely related to the southeastern endemic C. Leconte
Torrey ex Steudel,'! a Coastal Plain species ranging from North Carolina to
Louisiana.
Section Fusci (Kunth) Clarke!’ (plants annual; scales ovate; styles and stigmas
very short; achenes ovoid, glossy) is represented in the Southeast by one in-
troduced species. Cyperus difformis L., 2n = 34, a weedy Asian species, was
first collected in the eastern United States in Norfolk Co., Virginia, in 1935 by
Fernald (Tyndale). Lipscomb (1980b) has provided an interesting account of
the spread of this species in North America. The species was first collected in
the New World in New Mexico in 1850. It is a significant weed of rice fields
in California but has not yet become a problem in the southern rice-producing
states (Bryson). In contrast to the other weedy species of the genus (e.g., C.
esculentus), C. difformis is an annual that is capable of completing its life cycle
in only one month; a single plant can produce thousands of achenes. The species
is adapted to ground that is frequently flooded, such as rice fields. The seeds
germinate best under shallow water (McIntire). The type species of the section,
C. fuscus L., 2n = 72, is Eurasian; it is sparingly adventive from Massachusetts
to Nebraska and Virginia but has not yet been reported from the Southeast.
Subgenus Pycreus is characterized by having lenticular, laterally compressed
achenes and C, photosynthesis. There are about 120 species worldwide, of
which eight occur in our area. All our species are fibrous-rooted annuals, mostly
less than 30 cm tall, of disturbed wet soils. One, Cyperus louisianensis Thieret,
is endemic to southeastern Louisiana. Five pantropic species occur in our area:
C. flavescens L., 2n = 50, C. pumilus L., 2n = 94, C. flavicomus Michx. (C.
albomarginatus “Nees,” see Tucker, 1985a), C. polystachyos Rottb., and C.
lanceolatus Poiret. Cyperus bipartitus Torrey (C. rivularis Kunth, see Tucker,
1983a), n = 27, is a widespread North American species that also occurs in
the mountains of Mexico, Central America, and southern South America
(Tucker, 1983a). Cyperus filicinus Vahl is endemic to eastern North America
(tidal marshes from Maine to Louisiana).
Subgenus JuNcELLUs has only about six species worldwide. The pantropic
Cyperus laevigatus L., 2n = 80-84, was collected as a ballast plant in Wil-
mington, North Carolina (G. McCarthy s.n. in 1888, Gu!). It apparently never
Cyperus sect. HaspAni (Kunth) Clarke, Jour. Linn. Soc. Bot. 21: 119. 1884. Type species: C.
‘The name has been attributed to Torrey, but he published it provisionally under C. dentatus var.
multiradiatus Torrey (Ann. Lyc. Nat. Hist. Ne w York 3: 273. 1836). The name C. Leconte? was first
validly published by Steudel (Syn. Pl. Glum. 2: 17. 1854).
Cyperus sect. Fusci (Kunth) Clarke, Jour. eon Soc. Bot. 21: 131. 1884. Type spectes: C. fuscus L.
400 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
became established in the eastern United States. This species, which grows in
alkaline or brackish soils, is native to the area from western Texas to southern
California and southern Mexico, to the Lesser Antilles, and to South America.
Subgenus Cyperus contains about 400 species worldwide and about 35 in
the Southeast. Among these are pantropic, neotropical, and cosmopolitan rep-
resentatives. About half of the 35 are endemic to the United States, and many
of these are endemic to the Southeast; four are introduced from the Old World.
Plants of sect. UMBELLATI C. B. Clarke are characterized by their caespitose
habit, deciduous rachillas, and appressed, mostly persistent scales. This pan-
tropic group has twelve species in the southeastern United States: Cyperus
croceus Vahl (C. globulosus auct., non Aublet), C. echinatus (L.) Wood (C.
ovularis (Michx.) Torrey), C. Plukenetii Fern., C. ovatus Baldwin (C. Pollardii
Britton), C. Aystricinus Fern., C. refractus Torrey, C. retrofractus (L.) Torrey
(C. dipsaciformis Fern., see Carter & Jarvis), C. lancastriensis Porter, C. re-
trorsus Chapman (C. Nashii Britton), n = ca. 90 (Marcks, 1972a), C. thyrsi-
florus Jungh., C. retroflexus Buckley (C. uniflorus Torrey & Hooker, non Thunb.),
and C. /entiginosus Millsp. & Chase. Carter (1984) revised the North American
representatives, some of which were also studied by Marcks (1972b) and Tucker
(1983a, 1985b).
Plants of sect. LAxiGLumi'? are characterized by their rhizomatous, single-
stemmed habit, deciduous rachillas, and spreading, more or less deciduous
scales. Species of this section infrequently hybridize with those of the preceding
one (Marcks, 1972a, 1972b). Eight species occur in the eastern United States,
of which four are in our area; there are ten in the mountains of the southwestern
United States, Mexico, and Central and South America. The plants typically
grow in open, dry, sandy or gravelly habitats. The American species were
studied biosystematically by Marcks (1972a, 1972b), and the Mexican and
Central American ones by Tucker (1983a, 1984, 1985a). The species are cy-
tologically similar: all are n = 82 except Cyperus Schweinitzii Torrey, n = 84,
85 (Marcks, 1972b). Cyperus filiculmis Vahl (C. Martindalei Britton), C. lu-
pulinus (Sprengel) Marcks (C. filicu/mis auct., non Vahl), C. Grayi Torrey, and
C. Grayoides Mohlenbrock occur in our area.
The remaining southeastern species are scattered among six mainly pantropic
sections. Section Cyperus (sects. Esculenti Kiikenthal and Rotundi C. B. Clarke)
s most diverse in Australasia (Blake, J. H. Kern). In members of this section
oi the scales and the spikelets are persistent (a combination of characters
unknown elsewhere in the genus), and the stolons are tuberiferous. Cyperus
rotundus L., purple nut-sedge, is generally acknowledged to be the world’s worst
weed. It occurs throughout the Southeast, except in the mountains, but extends
only as far north as southern Missouri and southeastern Virginia. It does not
grow north of the mean 1°C January isotherm (Stoller). Cyperus esculentus L.,
yellow nut-sedge, is able to tolerate winter air temperatures as low as —
and is a serious weed in much of the world, especially in cooler regions where
Cyperus sect. LaxicLumi (C. B. Clarke) Kiikenthal, Pflanzenr. IV. 20(Heft 101); 220. 1936: based
on Mariscus suSeet Pexigiem B. Clarke, eeaae ae i Ser. 8: 103. 1908, “Laxighimae.”
I Mariscu K.) C. B. Clarke (= C. Manimae HBK.).
1987] TUCKER, CYPERACEAE 401
the more tropical C. rotundus does not grow. These two species also differ in
their thermal optima for growth. In Mexico C. esculentus is found from sea
level to about 2600 m, while C. rotundus occurs from sea level to about 1500
m (Tucker, 1985b). It is unclear whether these species are native to the New
orld. Cyperus esculentus now occurs in all 50 states and in southern Canada.
The stoloniferous nature of these two species underlies their success as weeds.
A single tuber can produce a population covering 2-4 m? in two months
(Horowitz). The sharp-pointed stolons can cause puncture wounds in the hands
of farm workers and curious agronomists and penetrate root crops such as
potatoes and yams. In 1821 Elliott noted that Cyperus rotundus was a great
problem for farmers in Georgia and South Carolina. He outlined a method for
removing an infestation by cultivating a fallow field weekly for a year (including
winter), thus allowing the tubers to be killed by exposure to drying and cold
air. Mulligan & Junkins provided a thorough summary of its biology, empha-
sizing weed control and management. Horak & Holt analyzed isozymes in ten
widely separated populations of C. escu/entus in California. Genetic variation
served to determine the relative importance of sexual and asexual reproduction.
Results indicated that reproduction by seeds is unimportant in maintenance
of populations in croplands. Stolons and tubers are the primary means of
reproduction. Germinability of seeds from northeastern populations ranged
from 7 to 95 percent; such variation was believed to be genetic (Mulligan &
Junkins). Seeds from a 50-year-old herbarium specimen had 5 percent ger-
mination (Mulligan & Junkins). Cyperus esculentus is self-incompatible (Horak
Holt).
Members of sect. Compress! Nees!’ are caespitose annuals with cuspidate
scales and emarginate achenes. Most of the species are native to the Old World
tropics. The pantropic Cyperus compressus L., n = 64, is the only representative
in the United States. It is found throughout the Coastal Plain and Piedmont,
as far north as Pennsylvania and Missouri. The only other New World species,
C. Wilburii G. Tucker, is endemic to the lowlands of southern Mexico. Its
larger size suggests that it may be a tetraploid derived from C. compressus.
Section IR1orpE1 Nees'> comprises several tropical and temperate eastern
Asian species. The plants are annual and have cae -appressed spikelets
and three-nerved, orbiculate scales. Cyperus Tria L. an oes in all
tropical and temperate regions of the New World an ad is a comm e
throughout the southeastern Coastal Plain and Piedmont. persis 1m plants
are cleistogamous. The staminal filaments elongate only enough to bring the
minute anthers into contact with the very short stigmas, which remain inside
the scales at anthesis. Often the anthers are later found agglutinated to the
stigmas.
Section Viscosi C. B. Clarke! is endemic to the New World and 1s represented
'4Cyperus sect. Compress! Nees, Linnaea 9: 234. 1834. Type species: C. compressus L.
'5Cyperus sect. IRio1pe1 Nees, Linnaea 9: 235. 1834. Type species: C. Iria L. Synonym: sect. /riae
(Kunth) C. B. Clarke, Kew Bull. a Ser. 8: 99. 1908.
'6Cynerus sect. Viscost C. B. Clarke, Jour. Linn. Soc. Bot. 21: 114. 1884. Type species: C. viscosus
eee - C. elegans L.). aN sect. Glutinosi (Béck.) Kiikenthal, Pflanzenr. IV. 20(Heft 101):
163.
402 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
by two species in the Southeast. Plants of this section have spicate inflores-
cences; the spikes are short and dense and appear glomerulate, which apparently
caused Kiikenthal to believe them to be closely related to plants of sect.
LUZEOLOIDEI (subg. PycNosTAcHys). The plants have kranz anatomy, further
supporting their placement in subg. Cyperus (Tucker, 1985b). They secrete a
viscid fluid and are sticky when living, hence the appropriate sectional name.
Two species occur in the Southeast. Cyperus elegans L. grows from southern
Florida and Texas south to Ecuador. Cyperus oxylepis Nees ex Steudel is a
South American species that has recently become an adventive in the United
States, where it was first noted in Texas (O’Neill). More recently it has been
reported in Louisiana (Thieret, 1964) and in Charleston County, South Carolina
(MacDougal 1501, 5 Aug. 1981, DUKE, NCU, NYS).
Subgenus TORULINIUM has a single representative in our area, the pantropic
and warm-temperate Cyperus odoratus L. It is a common species of disturbed,
wet soils, especially pond shores and stream banks. Five segregate species (e.g.,
C. Engelmannii Steudel, C. ferruginescens Bock.) have been recognized at
various subspecific ranks. Evidence for treating these segregates as conspecific
with C. odoratus has been published (Tucker, 1984). Three other species of
this subgenus occur in the New World tropics: C. Corre/lii (Koyama) G. Tucker
in the Bahamas, C. rhizophorae (C. B. Clarke) Standley along the Pacific Coast
of Central America, and C. filiformis Sw. in the Greater and Lesser Antilles.
Section RemireEA (Aublet) Kern contains a single pantropic species, Cyperus
pedunculatus (R. Br.) Kern (Remirea maritima Aublet), beach-stars. In our
area it occurs only 1n Peninsular Florida. The rhizomatous plants form mats
that bind sand dunes. This species has been treated as constituting a monotypic
genus, Remirea, which Kikenthal placed in the Rhynchosporoideae. Metcalfe
and Oteng- Yeboah showed convincingly that the anatomy of C. pedunculatus
is similar to that of the kranz species of Cyperus. Within Cyperus, the thickened
upper internode (“‘corky organ’’) of the one-flowered spikelets suggests a re-
lationship with subg. TORULINIUM (C. odoratus typically has spongy, thickened
rachilla internodes). Such internodes may serve to make the achenes buoyant,
thus contributing to dispersal by water, but experimental evidence for this
supposition is lacking.
REFERENCES:
Under family references see ALLAN et al.; shee BARROS (1938); BEAL; BENTHAM:
BLASER (1940, 1941a, 1941¢); BRASELTON; BREW KER; BROWN; BURKHALTER; CAROLIN
et al., CLARKE (1908, 1909); Ce oer ErreNn (1976a); EyLes & ROBERTSON;
FASSETT; FERNALD; GODFREY & WOOTEN; GONCHAROV ef al.; Goon et al.; HARBORNE;
HARBORNE ef al.; Hom ef al.; HotttumM; HuANG:; J. Hutrcuinson; F. D. Cer J. H.
A
RAYNAL; LOUROGNON; MCATEE; MEEUSE; METCALFE; NAPPER (1965); NEES von Es-
ENBECK; Nos_e & Murpuy; O’ NEILL; PATCH; RAYNAL (1972, 1973); RIKLI; SCHULZE-MOTEL
(1959, 1964); Smitu et al.; STANDLEY; TEERI et a/l.; TieTz; TORREY; VAN DER VEKEN; and
WINFREY & SAMSEL.
Ayers, B. The genus Cyperus in Mexico. Cathol. Univ. Am. Biol. Stud. 1: 1-103. 1946.
[Eighty species. ]
1987] TUCKER, CYPERACEAE 403
BAIJNATH, E. A study of Cyperus alternifolius L. sens. lat. (Cyperaceae). Kew Bull. 30:
521-526. 1975. [Includes C. involucratus Rottb. (C. alternifolius of authors, not L.),
a species introduced in the Southeast.
BASKIN, J. ne & C. C. Baskin. Germination of Cyperus inflexus Muhl. Bot. Gaz. 132:
3-9, a. [C. squarrosus L., temperate and pantropic species occurring in the
EAs Aven dormant seeds. Dormancy broken by stratification, scarification,
or nitrogenous compounds; light needed for germination
e possible ecological significance of the light requirement for ger-
mination in Cyperus inflexus. Bull. Torrey Bot. Club 98: 25-33. 1971b.
Effect of photoperiod on germination of Cyperus inflexus seeds. Bot.
oe 137: 269- 273. 1976.
Seasonal changes in the germination response of Cyperus inflexus
a to temperature and their ecological significance. [bid. 139: 231-235. 1978.
& Effects of wetting and drying cycles on the germination of seeds of
Cyperus inflexus. Ecology 63: 248-252. 1982. peloes and drying cycles decrease
time needed for germination once other conditions a
BENDIXEN, L. E. Anatomy and sprouting of yellow ee tubers. Weed Sci. 21: 501-
Betria, A. [., & E. R. MonTALp. Light effects on. bulb differentiation and leaf growth
in Cyperus rotundus L Phyton 32: 1-8. 1974
BLAKE, S. T. Cyperus rotundus (nut grass) and its allies in Australia. 14 pp. + 7 pls.
Brisbane, Australia. | 2. [Six species; keys, descriptions, illustrations. ]
Bryson, C. T. Weed a ene umbrella sedge (Cyperus difformis L.). So. Weed
Sci. Soc. Newsl. 7(1): 6. 1984.
CARTER, J. R., JR. A systematic study of the New World species of section Umbellati
of Cyperus. 279 pp. Unpubl. Ph.D. Thesis, Vanderbilt Univ. 1984. [Descriptions,
keys, ecology.]
& C. E. Jarvis. Re-evaluation and lectotypification of Scirpus retrofractus L.
Rhodora 88: 451-456. 1986. [C. retrofractus (L.) Torrey the correct name for C
dipsaciformis Fern.
CHERMEZON, H. Sur la position syst¢matique du genre Remirea. Bull. Soc. Bot. France
69: 809-814. 1922.
CHETRAM, R. S., & L. E. BENpIxEN. Gibberellic acid plus cytokinins induces basal bulbs
of purple nutsedge above ground. Weed Sci. 22: 55-58. 1974.
CLARKE, C. B. On the Indian species of Cyperus; with remarks on some that specially
illustrate the subdivisions of the genus. Jour. Linn. Soc. Bot. 21: 1-202. 1884.
[Important for sectional nomenclature. ]
Cray, K., T. N. HARDY, & A. M. HAMMonD. Fungal endophytes of Cyperus and their
effect on an insect herbivore. Am. Jour. Bot. 72: 1284-1289. 1985. [C. rotundus
and C. virens.]
Co tins, R. P., & M. B. Jones. The seasonal pattern of growth and production of a
temperate C, species, Cyperus longus. Jour. Exper. Bot. 37: 1823-1835. oe [Eur-
asian species; photosynthetic rates similar to those of temperate C, grass
CorcorRAN, M. L. revision of the ey aie in North and South ee.
Cathol. Univ. Am. Biol. Ser. 37: 1-68.
Costa, J., & A. P. AppLesy. Response of a ibe nutsedge varieties to three her-
bicides. Weed Sci. 24: 56-58. 1976.
Cour, P. Cyperus esculentus L., C. rotundus L., et C. rotundus var. brevibracteatus Legr.;
caractéres discriminatifs e distribution géographique. Biarritz Centre Etudes Rech.
Sci. B. 3: 181-192. :
Cusick, A. W. An a eat of halophytes in northern Ohio. Rhodora 72: 285. 1970.
[C. esculentus in runoff from salt well.
Denton, M. F. taxonomic treatment of the Luzulae group of Cyperus. Contr. Univ.
Mich. Herb. 11: 197-271. 1
404 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Anatomical studies of the Luzulae group of Cyperus (Cyperaceae). Syst. Bot. 8:
250- 262. 1983.
Druyts-Voets, E. Types van stengel—in bladstrukturen in het genus Cyperus L. (French
abstract.) Natuurwet. Tijdschr. 52: 28-49. 1980. [List of C, and C, species and
sections, illustrations. ]
Eames, A. J. Comparative effects of spray treatments with growth regulating substances
on the nut grass, Cyperus rotundus L., and anatomical modifications following treat-
ment with butyl 2,4-dichlorophenoxyacetate. Am. Jour. Bot. 36: 571-584. 1949.
E.uiotr, S. A sketch of the botany of South Carolina and Georgia. Vol. 1. iv + 606
pp. Charleston, South Carolina. 1821.
Feperowicz, M. F. The significance of the achene and stoma in the status of Eucyperus
and Mariscus (Cyperaceae) He on the studies of plastic replicas. Cathol. Univ.
Am. Biol. Studies 75: 1-50.
FisHer, J. B. Development of - intencalany meristem in Cyperus alternifolius. Am.
Jour. Bot. 57: 691-703. 1970a. [= C. involucratus Rottb., a ata cultivated in
pools and greenhouses, sparingly naturalized in the Southea
Xylem derived ae intercalary meristem of Cyperus ie nifolius. Bull. Torrey
Bot. Club 97: 58-66. 1970b.
Control of the eee intercalary meristem of Cyperus alternifolius. Am.
Jour. Bot. 57: 1017-1026. 1970c.
ForeERO Pinto, L. E. Etnobotanico de las comunidades indigenas Cuna y Waunana,
Chocd (Colombia). (English abstract.) Cespedesia 9(33-34): 115-301. 1980. [De-
coction of roots of C. chalaranthus Presl used for stomachaches, inflorescences of
C. luzulae used for decoration.]
FRIEDMAN, T., & M. Horowitz. Biologically active substances in subterranean parts of
purple nutsedge. Weed Sci. 19: 398-401. 1971
Gara, D. . E. BENpIxEN, & S. R. ANDERSON. Rhizome differentiation in yellow
nutsedge. Weeds 15: 124-128. 1967. [C. esculentus.]
GarRONI, L. W., & W.H. Murpy. Systematic relationship of the granite outcrop endemic
Cyperus granitophilus McVaugh to Cyperus inflexus Muhl. (Abstract.) ASB Bull. 11:
1964. [C. granitophilus an autotetraploid derivative of C. squarrosus L. (C.
inflexus). ]
GUAGLIANONE, E. R. Caracteres diferenciales entre Cyperus rotundus L. y C. esculentus
.. presencia de un pliegue ligular en el primero. Rev. Assoc. Argent. Malezas 6:
21-35. 1978.
GUIGNARD, J. L. Cypéracées. Développement de ’embryon chez le Cyperus vegetus
Willd. ae Rend. Acad. Sci. Paris 252: 2125-2127. 1961.
Gupta, S. K., R. C. SHARMA, O. P. AGGARWAL, & R. B. Arora. Anti-inflammatory
rea of * isolated from Cyperus scariosus R. Br. Indian Jour. Exper. Biol. 10:
41, 42. 1972.*
ee J. B., C. A. Wittiams, & K. L. Witson. Flavonoids in leaves and inflores-
cences of Reel Cyperus species. Phytochemistry 21: 2491-2507. 1982.
Hauser, E. W. Development of purple nutsedge under field conditions. Weeds 10: 315-
321. 1962. [C. rotundus.]
Haynes, R. R., & A. LASSEIGNE. Cyperus giganteus (Cyperaceae) in Florida. Sida 3:
345. 1969, ene state record. ]
Hikino, H., K. Aorta, D. UWA ANO, & T MoTO. Structure and absolute configu-
ration of alpha- eae and He rotunol sesquiterpenoids of Cyperus rotundus.
Tetrahedron 27: 4831-4836. 197
Hockina, P. J. Effects of sodium se potassium chlorides on the growth and accu-
mulation of mineral ions by Cyperus involucratus Rottb. Aquatic Bot. 21: 201-217.
1985. [Potential use in treating wastewater.
Hoim, T. Studies in the Cyperaceae. XXIII. The inflorescence of Cyperus in North
America. Am. Jour. Sci. TV. 18: 301-307. 1904.
1987] TUCKER, CYPERACEAE 405
Horak, M. J., & J. S. Horr. Isozyme variability and breeding systems in populations
of yellow ee (Cyperus esculentus). Weed Sci. 34: 538-543. 1986.
Horowitz, M. Growth, tuber formation, and spread of Cyperus rotundus L. from single
79.
Horvat, M. L. A revision of the subgenus Mariscus found in the United States. Cathol.
Univ. Am. Biol. Ser. 33: 1-147. 1941. [Keys, descriptions, specimen citations. ]
JAN, P. Essais de lutte contre le Cyperus rotundus, étude bibliographique. Agron. Trop.
27: 255-262. 1972.
Jones, M. B. Papyrus: a new fuel for the Third World. New Sci. 99: 418-421. 1983.
& T.R. MicsurN. Photosynthesis in papyrus (Cyperus Papyrus). Photosynthetica
12: 197-199. 1978.
F. M. Mutuvurti. The canopy structure and microclimate of papyrus (Cyperus
Papyrus) swamps. Jour. Ecol. 73: 481-491. 1985.
Justice, O. L. Germination ea . seeds of nutgrass (Cyperus rotundus L.). Assoc.
Off. Seed Anal. Proc. 46: 67-7 56.
. Germination, dormancy, se viability in seeds of certain weedy species of
Cyperus. Ibid. 47: 167-175. 1957.
KAMAKHINA, G. L. The nae a of purple nutsedge, Cyperus rotundus L. Weed
Abstr. 21: no. 894.
ae J. W. Cyperus Se ) Torr. var. cylindricus (Ell.) Torr. (Cyperaceae)
w to New Mexico. Sida 10: 258. 1984.
Kian NNA, P. A contribution to the ea, of Cyperus rotundus L. Proc. 43rd Indian
Sci. Congr. Sed 236, 237.
Koyama, T. ew species of ee ulinium (Cyperaceae) from the Sere Islands.
Brittonia 28: 252- 254. 1976. [= Cyperus Correllii (Koyama) G. T
KUKENTHAL, G. Cyperus. In: L. Diets, ed., Pflanzenr. IV. 20(Heft 101): 1.620. 1935-
1936.
LEMAIRE, R. J. Recent se records for Nebraska. Rhodora 72: 283, 284. 1970. [C.
difformis in “marsh” on roof of office building in Lincoln.
Lipscoms, B. L. ae ae te (Cyperaceae): new to Arkansas, Kansas, and
ae Sida 8: 300-327. 1980a.
Cyperus difformis L. eee in North America. /bid. 320-327. 1980b.
fied Gon, spread, and present range of this Old World species in North and
Central America.]
Lye, K. A. Two new subgenera of Cyperus. Nordic Jour. Bot. 1: 57-61. 1981. [Subgenera
ee (synonym of subg. PycNosTACHYs) and FIMBRICYPERUS described; il-
lustratio
Man, A. P. ene vascular bundles in Cyperus. Sci. ou 25: 437, 438. 1960.*
. Air-space tissue in Cyperus. [bid. 28: 39, 40 62.
Marcks, B. G. Population studies in North American Cyperus section seal (Cy-
peraceae). v + 405 pp. Unpubl. Ph.D. Thesis, Univ. Wisconsin, Madiso .1972a
. Preliminary reports on the flora of Wisconsin, no. 66. Cy eae ode
Family II. The genus Cyperus—the umbrella sedges. Proc. Wisc. Acad. Sci. Arts
Lett. 62: 261-284. 1972b. [Keys, descriptions, illustrations, chromosome numbers,
es maps.]
H. H. Ixtis. Post- ee hybridization of Cyperus Schweinitzii and C. maci-
lentus. (Abstract.) Am. Jour. Bot. 54: 659, 660. 1967.
McGivney, M. V. A revision , the subgenus Eucyperus found in the United States.
Cathol. Univ. Am. Biol. Ser. 26: 1-74. 1938. [Keys, descriptions, specimen citations;
illustrations of scales and achenes.]
McIntire, S. Seed reserves in temperate Australian rice fields following pasture rotation
and continuous cropping. Jour. Appl. Ecol. 22: 875-884. 1985. [C. difformis is one
of the two most abundant weeds; its seeds germinate best under flooded conditions. ]
406 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
McLauGuuin, A. D. The genus Cyperus in the West Indies. Cathol. Univ. Am. Biol.
Stud. 5: 1-108. 1944
Mounan Raw, H. Y.., ATRA. Stimulation of flower formation by cytokinins in
the excised immature inflorescences of Cyperus rotundus. Phytomorphology 20: 22-
29. 1970.
mara ae R.H. Anew species of Cyperus from the Illinois sand prairies. Brittonia
: 255, 256. 1959. [C. Grayoides, illustrated; also in Louisiana and eastern Texas;
see MARCKS (1972a).
Cyperaceae of Illinois. I. Cyperus. Am. Midl. Nat. 63: 270-306. 1960.
[Keys, descriptions, gg tae
MUuLuGan, G. A., & B. E. Junkins. The biology of Canadian weeds. 17. Cyperus
esculentus L. Canad. a Pl. Sci. 56: 339-350. 1976.
O’NEILL, H. T.
The status and distribution of some Cyperaceae in North and South
America. Rhodora 44: 43-64. 1942.
OTENG-YEBOAH, A. A, ology, anatomy, and taxonomy of the genus Remirea
Aublet (Cyperaceae). Boissiera 24: 197-205. 1975.
Overton, J. B. Studies on the relation oe living cells to transpiration and sap-flow
in Cyperus. Bot. Gaz. 51: 28-63. 191
Papuye, M. D. Studies in Cyperaceae. I. eee of Cyperus Iria L. Nat. Inst. Sci.
India Proc. i 1-10. 1971.
PuaTAK, S. C., D. R. Sumner, H. D. We ts, D. K. BELL, & N. C. Giaze. Biologica
control of ee sce with ne indigenous rust fungus Puccinia pn
Science 219: 1446, 1447. 3.
Piccoul, F., & R. GERDOL. les field weed communities in Ferrara Province, nort
her
Italy. Aquatic Bot. 10: 317-328. 1981. [Cyperus difformis L. and C. serotinus Rottb.
are important weeds. ]
Porcuer, F. P. Resources of the southern fields and forests. ed. 2. [i +] xv + 733 pp.
Charleston, South Carolina. 1869. [Cyperus esculentus, 684, as C. repens: C. rotun-
dus, 685, as C. hydra.]
PRAKASH, N. A survey of the leaf structure and its relationship to photosynthetic path-
ways in certain Malaysian plants. Malaysian Jour. Sci. 4(A): 67-73. 1976. [C. diffusus
Vahl is C,
RAYMOND, M. A note on x Cyperus Weatherbianus. Rhodora 64: 349, 350. 1962.
Raynat, J. Notes cypérologiques: V. Sur un groupe de Cyperus montagnards
américains. Adansonia 6: 385-392. 1967. [Illustrations; distribution map of C.
lixus HBK. omits occurrences in Louisiana and Central America; see TUCKER (1983a),]
. Notes cypérologiques: 31. Mélanges nomenclaturaux (Cyperoideae). Adansonia
17: 43-47. 1977. [Notes on typification of C. giganteus Vahl and C. odoratus, both
aie t]
te in the Southeas
Reep, M.S. The genus Cyperus in North Carolina. Jour. Elisha Mitchell Sci. Soc. 52:
295- 306. 1936.
SCANLON, G. M. A study of the genus Cyperus in the Hawaiian a Cathol. Univ.
Am. Biol. Ser. 41: 1-62. 1942. eee descriptions, specimen citations. ]
SHARMA, O. P., & R. SHIAM. of cuticular papillae in in ease pa
Bangalore 50: 236. 1981. [C. pilosus Vabl has one to four papillae s ach
abaxial stoma; such papillae are rarely present in C. digitatus Roxb., C. ta.
Retz., and C. rotundus.]
Son, S. L., P. B. KAUFMAN, & W. C. BiGELow. Electron microprobe analysis of silica
cells in leaf eoitenaal cells of Cyperus alternifolius. Plant Soil 36: 121-128. 1972a.
ctron microprobe analysis of silicon and other ie ae
in developing silica cells of leaf and ae of Cyperus alternifolius. Ann.
(London) 36: 611-619 b.
STOLLER, E. W. Effect of soil minimum temperature on differential distribution of
1987] TUCKER, CYPERACEAE 407
Cyperus rotundus and Cyperus esculentus in the United States. Weed Res. 13: 209-
Qe
MA, & V. M. BAHN. Yellow nutsedge tuber germination and seedling
ee Weed Sci. 20: 93-97. 1972.
Taytor, J. R., & D. K. Evans. A taxonomic study of the genus Cyperus (Cyperaceae)
in West Virginia. ASB Bull. 25: 64. 1978. [Fourteen species; C. croceus (C. globu-
losus), new state record.]
Turret, J. W. More additions to the Louisiana flora. Sida 1: 294, 295. 1964. [Cyperus
difformis, C. oxylepis, and C. retroflexus (as C. uniflorus).]
Cyperus louisianensis (Cyperaceae), a new species from southern Louisiana.
Proc. Louisiana Acad. Sci. 40: 23-26. 1977. [Subgenus Pycreus; related to C. bi-
partitus Torrey; illustrations. ]
THompson, K., P. R. SHewry, & H. W. WootHouse. Papyrus swamp development in
the Upemba Basin, Zaire: studies of population structure in Cyperus papyrus stands.
Bot. Jour. Linn. Soc. 78: 299-316. 197
Tucker, G.C. Taxonomy of the genus Cyperus (Cyperaceae) in Costa Rica and Panama.
Syst. Bot. Monogr. 2: 1-85. 1983a. [Fifty species, keys, descriptions, distribution
maps; subgeneric Sea atioue ]
Two new species of Cyperus (subgenus Protocyperus) from Mexico and Central
riences Bull. ae Bot. Club 110: 343-347. 1983b. [C. microbrunneus, C. na-
yaritensis of subg. Pycnostachys; illustrations. ]
Taxonomic notes on two common neotropical species of Cyperus. Sida 10: 298-
307. 1984. [C. edoratus.]
——. Cyperus flavicomus, the correct name for Cyperus ee Rhodora 87:
539-541. 1985a. [A pantropic species occurring 1n the Southe
. Arevision of the Mexican species of Cyperus L. (Cyeracee) 285 pp. Unpubl.
Ph.D. Dissertation, Duke Univ. 1985b. [Eighty-five speci
The correct name for Cyperus cayennensis (C. flavus), Se Southw. Nat
30: 607, 608. 1985c. [C. aggregatus (Willd.) Endl. the correct name for a neotropical
species that occurs northward to Texas and Louisiana.]
. The species of Cyperus described by Liebmann in “Mexicos halvgraes.”’ Syst.
Bot. 11: 14-19. 1986a.
e distribution of C, and C, species of Cyperus (Cyperaceae) in North and
Central America. (Abstract.) Am. Jour. Bot. 73: 792. 1986b. [Abundance calculated
from number of herbarium collections; relative abundance of C, species increases
with latitude. ]
. New records of Cyperus (Cyperaceae) from West Virginia. Castanea 52: 145,
146. 1987. [C. Houghtonii, C. iria, C. polystachyos.]
TumBLESOoN, M. E., & T. KOMMEDAHL. Reproductive potential of Cyperus esculentus L.
by tubers. Weeds 9: 646-653. 1961.
TYNDALE, R. W. Distribution of Cyperus difformis L. (Cyperaceae) in the southeastern
United States. Castanea 48: 277-280. 1983.
VerMA, 8S. C., A. PAL, & B. L. — Anatomical studies on some species of
eee L. Pl. Sci. 5: 52-59. 1973.
VorstTeER, P. J. Revision of the ee of Mariscus Benth. and related genera in
southern Africa. 348 pp. Unpubl. D.Sc. Dissertation, Univ. Pretoria. 1978.
WILuiaMs, R. D. Intraspecific competition of yellow nutsedge. Proc. So. Weed Sci. Soc.
34: 231-238. 1981.
WILts, G. D., & G. A. Briscoe. Anatomy of purple nutsedge. Weed Sci. 18: 631-635.
1970.
WEEDON, R. R., & H. A. STEVENS. Cyperus fuscus in Nebraska and South Dakota.
Rhodora or, 433. 1969.
wee Pals: Sevier and photoperiodic responses of yellow nutsedge. Weed Sci.
210-219. 197
408 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
9. Kyllinga Rottboell, Descr. Icon. Rar. Nov. Pl. 12. 1773, nom. cons.
Small, rhizomatous or tufted perennials (1 species annual). Culms trigonous
or roundly trigonous, smooth. Leaves 1-5, basal; sheaths short, closely fitting
the culms, ligule lacking; blades flat or V-shaped in cross section [lacking], the
margins and keels scabrellate, especially distally; chlorenchyma radiate; bundle
sheaths 2-layered (“Cyperus type’). Involucral bracts 2-4, leaflike, horizontal
to slightly reflexed or erect. Spikes 1-4, sessile, densely ovoid to cylindrical.
Spikelets 15-150 per spike, not readily distinguishable without magnification,
ovate to lanceolate, decidedly flattened. Scales 4, the 2 basal minute, the 2
distal much longer, making up the bulk of the spikelet, the lower of these (the
third scale of the spikelet) subtending a perfect flower, the upper (fourth scale)
slightly smaller, sterile or infrequently bearing | or 2 (often abortive) stamens.
The fertile scale of the spikelet ovate, conduplicate, with a conspicuous smooth
or spinulose-scabrellate [fimbriate or erose] keel terminating in a mucronate
or mucronulate [aristate] apex, laterally 2- to 4-nerved. Flowers perfect. Peri-
anth lacking. Stamens |-3; filaments ribbonlike, about as long as the subtending
scales; anthers oblong-elliptic to linear, the apices of the connectives not pro-
longed; pollen grains 4-aperturate [uniaperturate], obovoid, psilate, trinucleate.
Styles capillary, smooth; stigmas 2, about as long as the styles. Achenes len-
ticular, laterally compressed, narrowly ovoid to oblong or ellipsoid, about '2
the length of the subtending scale, the apex obtuse, apiculate, the base cuneate
to rounded, barely to decidedly stipitate, the surface puncticulate. Embryos
narrowly ellipsoid. Base chromosome number 60. (Cyperus subg. Kyllinga
(Rottb.) Valck.-Suringar.) Type species: K. monocephala Rottb., nom. illeg.
(= K. nemoralis (J. R. & G. Forster) Dandy ex Hutchinson & Dalz., typ. cons.).
(Named for Peter Kylling, Danish botanist, d. 1696.)
A genus of about 40-45 species, nearly all of which are tropical. The greatest
diversity is in tropical East Africa and Madagascar, where there are 30-35
species. Eight occur in southern Asia, three or four in eastern Asia, and two
in Australasia. Two (neither endemic) grow in the Hawaiian Islands, but none
occurs in Europe. There are eight species in the New World; three of these,
Kyllinga pumila Michx., K. odorata Vahl, and K. brevifolia, 2n = 120, which
occur in the Southeast, are pantropic. Kyllinga vaginata Lam. and K. tibialis
Ledeb. are species of littoral habitats in the Caribbean, South America, and
tropical West Africa. Kyllinga nudiceps C. B. Clarke is endemic to Isla del
Coco, in the Pacific some 300 km southwest of Costa Rica. Ky/linga squamulata
Thonn. ex Vahl (Cyperus Metzii Mattf. & Kiikenthal), from tropical Asia, is
introduced in Florida and the West Indies; K. brevifolioides (Delahoussaye &
Thieret) G. Tucker,'’ from temperate eastern Asia, has become sparingly es-
tablished in the eastern United States in the area from Connecticut to western
North Carolina and Tennessee. The four southeastern species are mostly weedy
plants of disturbed, usually moist, sunny places. Kyllinga pumila is a common
“"Kyllinga brevifolioides (Delahoussaye & Ae G. Tucker, comb. nov., based on Cyperus brevi-
folioides Delahoussaye & Thieret, Sida 3: 131.
1987] TUCKER, CYPERACEAE 409
weed of lawns and croplands in the eastern United States from Pennsylvania
and Missouri south to the Gulf Coast.
Kyllinga differs from Cyperus, with which it has been combined by some
workers, in its very short rachilla and in the two lowest sterile scales of its
spikelets being greatly reduced. Taxonomically useful characters have been
reviewed by Tucker. The most important of these are habit (rhizomatous
perennials or caespitose annuals), length and orientation of the involucral bracts,
and length of the anthers. Such characters as number of stamens and presence
of spinulose prickles on the keels of the scales have previously been used
(Delahoussaye & Thieret) but frequently vary within individuals of the same
species and sometimes within spikes of a single plant.
The plants are probably at least partly wind pollinated. However, because
of the close spacing of the spikelets within an inflorescence, some anthers
probably shed their pollen directly onto stigmas of adjacent spikelets. Insect
pollination may be important in some species with conspicuous, whitish or
cream-colored spikes (e.g., Kv/linga odorata), as it is in many species of Rhyn-
chospora sect. DICHROMENA. Syrphid flies have been observed visiting indi-
viduals of K. tibialis in Costa Rica (MacDougal 1190, Duke) and K. odorata
in Mexico (Tucker 2222, DUKE).
REFERENCES:
Under family references see BARROos (1935); BEAL; BENTHAM; BLASER (1940, 1941a);
CAROLIN et al.; CLARKE (1908, 1909); Erren (1976a); FASsETT, ae af eee
GONCHAROV et al.; seme eye ay ee et al., Hom et al.; HottrumM; HUAN
J. Hutcuinson; Kuntu; LE MAout & DecatsNe; LERMAN & RAYNAL; Rieck (1966).
NEES VON ESENBECK; ONENLL: ween (1972, 1973); Riki; ScHULZE-MoTEL (1959,
1964); Torrey; and VAN DER VEKEN
DELAHOUSSAYE, A. J., & J. W. THIERET. Cyperus subgenus Kyllinga (Cyperaceae) in the
ntinental United States. Sida 3: 128-136. 1967. [Synopsis; illustrations of spikelets
oe achenes, distribution maps.]
GOVINDA Lu, E. The systematic en nese Indian Cyperaceae: Cyperus sub-
genus Kyllinga (Rottb.) Suringar. Jour. Linn. Soc. Bot. 62: 41-58. 1969.
LYE, *K. A. New taxa and combinations in Ky llinga. Nordic Jour. Bot. 1: 741-747.
1981
McNauscuton, S. J. Ecology of a grazing ecosystem: the Serengeti. Ecol. Monogr. 55:
259-294. 1985. [During the wet season, leaves of Kyllinga nervosa provide forage
for the Thomson’s gazelle.]
Papuye, M. D. Studies in the Cyperaceae. III. Life history of Kyllinga brevifolia Rottb.
with ; brief discussion on the taxonomic position of Kyllinga. Bot. Gaz. 132: 172-
179.
TUCKER, a C. A revision of the genus Kyllinga Rottb. (Cyperaceae) in Mexico and
Central America. Rhodora 86: 507-538. 1984. [Six species; keys, descriptions, dis-
tribution maps, extensive specimen citations.
10. Lipocarpha R. Brown in Tuckey, Narr. Exped. Congo 5: 459. 1818, nom.
cons
Small, caespitose annuals of wet sandy or peaty soils. Roots fibrous, rhizomes
absent. Culms 1—20(-100), usually densely clustered, erect, spreading, or curved,
410 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
filiform, terete, glabrous. Leaves | or 2, basal, filiform, about as wide as the
culms, the lower reduced to a bladeless sheath or a sheath bearing merely an
involute appendage, the upper with blade up to 4 as long as the culm, or
reduced like the lower one; stomata paracytic; chlorenchyma radiate; the bundle
sheaths 2-layered (“Cyperus type’). Inflorescences unbranched, a sessile cluster
of 1-4 dense spikes; bracts 1-4, filiform, 1-4 times as long as the spikes, leaflike,
the longest erect, appearing as a continuation of the culm, the other(s) shorter
than or equaling the spikes, borne approximately perpendicular to the culm:
rays none. Spikes (“spikelets”) 1-4, sessile, ovoid [globose]; denuded rachis
persistent, with rhombic scars where the spikelets were attached. Spikelets
(“flowers”) [20-]50-150, densely spirally arranged, borne approximately per-
pendicular to the rachis, deciduous. Scales (1, 2, or) 3; outer scale lanceolate
to ovate-lanceolate, planar or nearly so, with 2 conspicuous medial veins and
a less conspicuous central one, laterally weakly 1- or 2-nerved or essentially
nerveless, mucronulate [aristate]; inner scale hyaline, equaling or shorter than
the outer, or reduced to a scalelike appendage much shorter than the outer,
with 3—5 inconspicuous veins or veinless, or absent; third scale present between
the outer scale and the achene in some species, similar to or smaller than the
second. Flowers perfect. Perianth lacking. Stamens | or 2; filaments capillary,
about *% as long as the outer scale; anthers ovoid, the apices of the connectives
not prolonged; pollen grains 4-aperturate, obovoid to subspheroidal, psilate or
scabrate. Styles filiform; stigmas 2, about '2 as long as the styles, minutely
swollen apically, glabrous, deciduous before the achenes mature. Achenes tri-
gonous to terete, obovoid to cylindrical, slightly shorter than the outer scale,
the base sessile to stipitate, the apex obtuse to subtruncate, apiculate, the surface
papillose. Embryos ellipsoid. Base chromosome number 6. (Incl. Ascolepis Nees
ex Steudel, Hemicarpha Nees ex Arnott.'*) Type species: L. senegalensis (Lam.)
T. & H. Durand (L. argenteum (Vahl) R. Br., nom. illeg.; see Haines & Lye).
(Name from Greek, /ipo, to fall, and carpha, chaff, referring to the deciduous
hyaline inner scale of the spikelet.)
A genus of about eight species occurring in tropical and warm-temperate
regions. Five grow in North America: Lipocarpha maculata (Michx.) Torrey,
on the Coastal Plain from Virginia to Texas, southward into the tropics; L.
occidentalis, restricted to the Pacific coast; L. Drummondii, from Oklahoma
and Texas west to New Mexico; L. aristu/ata, across the United States from
South Carolina and Florida west to Washington and California; and L. mi-
'8The inclusion of Hemicarpha in Lipocarpha necessitates the following new combinations for
species occurring in the New World:
Lipocarpha artstulata (Cov.) G. Tucker, based on Hemicarpha micrantha var. aristulata Cov. Bull.
Torrey Club 21: 36. 1894.
z plea canlests (Nees) G. Tucker, based on Hemicarpha Drummondii Nees in Martius, Fl. Brasil.
2(1): 62
a a Be G. Tucker, based on Scirpus micranthus Vahl, Enum. 2: 254.
.. occidentalis (Gray) G. Tucker, based on Hemicarpha occidentalis Gray, Proc. Rede: 7: 391.
868.
~
MON
L. Schomburgkit (Friedl.) G. Tucker, based on Hemicarpha Schomburgkii Friedl. Am. Jour. Bot. 28:
860. 1941
1987] TUCKER, CYPERACEAE 41]
crantha, throughout the United States and southeastern Canada, southward to
tropical South America. Lipocarpha Schomburgkii is known only from the
Guyana region of northern South America.
All species are small, inconspicuous plants of disturbed wet soils, especially
shores of ponds and pools. Because of their small size (less than 30 cm tall,
and often less than 1 cm!), they are easily overlooked and are probably more
frequent and widely distributed than available collections indicate.
aynal’s view that Lipocarpha is a highly reduced derivative of Cyperus
seems well founded and is accepted here. The fact that both genera have
“Cynerus-type” kranz anatomy (Metcalfe) further strengthens this conclusion.
The achene and subtending scales of Hemicarpha are probably homologous to
a single spikelet of Kyilinga or Cyperus. Friedland suggested that the inner
hyaline scale represented five perianth members that correspond to the bristles
subtending the achenes in some species of Scirpus. Raynal’s interpretation of
the inner scales of Lipocarpha (and Hemicarpha) as reduced scales of a spikelet
appears more plausible than Friedland’s view.
Haines & Lye studied the African species previously assigned to Hemicarpha
and Lipocarpha and concluded that the two genera should perhaps be merged.
Goetghebeur (pers. comm.) has recently studied all the Old World species of
these genera, as well as those of the closely related genus Ascolepis. He con-
cluded, as I had from my independent investigations, that the three genera
should be combined.
Chromosome numbers have been reported for Lipocarpha argentea R. Br.
(2n = 26) and L. microcephala Kunth (2n = 46). This suggests a base chro-
mosome number o
No species is gathered as food or for medicinal purposes. Lipocarpha argentea
and L. microcephala (R. Br.) Kunth are recorded as weeds in eastern Asia
(Holm et al.).
REFERENCES:
Under family references see BARROS (1938); BEAL; BENTHAM; BLASER ae 1941a);
Brown; CAROLIN ef al.; CLARKE (1908); ErTEN (1976a); FasseTT; GODFREY & WOOTEN;
Ho. et al.; HUANG; J. Hutcuinson; J. H. KERN; Koyama (1962b); een Le Maout
& DECAISNE, LERMAN & RAYNAL; METCALFE; NAPPER (1965); NEES VON ESENBECK;
O’NEILL; SCHULZE-MOoTEL (1959, 1964); STANDLEY; TEERI et a/.; TORREY; and VAN DER
VEKEN
FRIEDLAND, S. The American species of Hemicarpha. Am, Jour. Bot. 28: 855-861. 1941.
[Revision of the North and South American species; keys, distribution map, de-
scriptions; discussion of morphology of the spikelets; no specimen citations. ]
Haines, R. W., & K. A. Lye. Studies in African Cyperaceae IV, Lipocarpha R. Br.,
Hemicarpha Nees, and Jsolepis R. Br. Bot. Not. 124: 473-4 1971.
Koyama, T. The genus Lipocarpha R. Br., its morphology and systemat ic position in
the family Cyperaceae. (In Japanese, English abstract.) Acta Phytotax. Geobot. 33:
218-226. 1982.
Pata, E. Uber den morphologischen Wert der Bliite der Gattungen Lipocarpha und
Platylepis. Ber. Deutsch. Bot. Ges. 23: 316-323. pl. XIV. 1905. [Floral diagrams. ]
RAYNAL, J. Notes cypérologiques: VII. Sur quelques Lipocarpha africains. Adansonia,
IJ. 7: 81-87. 1967. [Two new species; illustrations. ]
412 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
FiGure 3. Rhynchospora sect. DICHROMENA. a-c, R. colorata: a, habit (note rhizomes
to right), x 2; b, mature achene, tubercle scarcely decurrent on body of achene, x 20;
c, same, in longitudinal section, the 2 layers of the achene wall separated by dotted line,
seed coat unshaded, endosperm stippled, embryo unshaded, x 20. d-k, R. floridensis:
d, head of spikelets subtended by involucral bracts, x 3; e, | large and | small spikelet
enclosed by 2 scales, = 6; f, abaxial surface of spikelet, 2 scales removed, flowers pro-
tandrous, x 10; g, same spikelet, adaxial surface, | stamen and 3 scales removed, x 6;
1987] TUCKER, CYPERACEAE 413
Tribe SCHOENEAE Dumortier, FI. Belg. 144. 1827. (Tribe Rhynchosporeae Fenzl
in Endlicher, Gen. Pl. 2: 115. 1836.)
11. Rhynchospora Vahl, Enum. Pl. 2: 229. 1806, nom. cons.
Small to large, caespitose or single-stemmed, perennial [annual] herbs of
moist open woods, bogs, pocosins, ditches, and pond shores. Roots fibrous;
rhizomes or stolons present in a few species. Culms trigonous, subtrigonous,
or terete, smooth throughout or ribbed just below the inflorescence, glabrous,
leafy [leafless]. Leaves numerous, basal, cauline, or both; basal leaves with
blades flat to conduplicate or involute-filiform, the margins and midveins
generally scabrellate with unicellular [multicellular] prickles, the surfaces gla-
brous or with prickles like those on the margins, or pubescent with long, flexible,
unicellular hairs, or papillose (in R. alba); cauline leaves shorter than but
otherwise similar to the basal ones; stomata paracytic, generally confined to
the abaxial surface; chlorenchyma not radiate [radiate in some tropical species].
Inflorescences terminal (sometimes also lateral, the lateral ones smaller and
less branched than the terminal), fasciculate or cymose; bracts 1-6, leaflike
(sometimes basally whitened); rays slender, terete, smooth or scabrellate; heads
loosely to densely ovoid or capitate. Spikelets solitary, globose, ellipsoid, or
slenderly lanceolate, the 1-5 basal scales sterile. Scales spirally arranged, closely
imbricate, ovate to lanceolate, entire or mucronulate at apex, nerveless to rather
prominently nerved, the midvein most conspicuous. Flowers perfect (the ter-
minal | or 2 scales sterile or subtending rudimentary ovaries and functional
stamens). Perianth bristles lacking or !-6(—20), smooth, barbed, or plumose,
persistent. Stamens (1—)3(-1 2); filaments capillary or ribbonlike; anthers elliptic
to oblong, the apices of the connectives not prolonged; pollen grains uniaper-
turate, obovoid, psilate or scabrate, binucleate. Styles glabrous; the stigmas
longer than, equaling, or much shorter than the style. Achenes lenticular (dor-
siventrally flattened), ovoid to slenderly ellipsoid, crowned with a pyramidal
to subulate tubercle shorter than to 3 times longer than the body of the achene,
the base sessile to conspicuously stipitate, the lateral edges often raised to form
a conspicuous ridged margin, the surface alveolate to cancellate (rarely smooth
or nearly so), transversely rugulose or not. Base chromosome number 5. (Incl.
Psilocarya Torrey, Dichromena Pers., C alyptrostylis Nees.) TYPE SPECIES: R.
alba (L.) Vahl (Schoenus albus L.), typ. cons. (Name from Greek, rhynchos,
snout, and spora, seed, in reference to the prominently beaked achenes.)
A genus of about 225 species, worldwide in distribution, with greatest di-
versity in the New World tropics; about 60 occur in the southeastern United
States. Temperate North America, especially the southeastern Coastal Plain,
is rich in species, and there are many others in the Old World tropics. Only a
h, flower with subtending scale, anthers fallen, 5 scales and rachilla of spikelet removed,
x 10; i, flower removed from spikelet, anthers dehiscing, styles not yet elongated, stigmas
not receptive, x 12; j, nearly mature achene with persistent style and stigmas, x 20; k,
0.
mature achene, tubercle decurrent on body of achene, x
414 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
few species are indigenous to temperate Eurasia—three species in Europe and
four in the Soviet Union east of the Urals.
Kukenthal’s worldwide monograph (1949, 1950, 1951) provided a basis for
identification and further study of the genus RAynchospora. Gale, in her careful,
well-illustrated monograph, did much to clarify the taxonomy of the North
American species. Thomas (1984) has recently investigated the tropical section
DicHRomEna (Pers.) Pfeiffer and confirmed its inclusion in RAynchospora.
The genus is little known cytologically; chromosome numbers have been
published for only ten species (summarized by Thomas, 1984). These suggest
a base chromosome number of 5, in keeping with the base number for other
genera of the family.
There are three subgenera in RAynchospora (Kiikenthal, 1949, 1950, 1951).
The largest of these, including about 54 of the 60 species in our area, is subg.
RuHyYNcHospora (Eurhynchosporae Gray), species of which have papery spikelet
scales and stigmas equaling or longer than the styles. Complete descriptions of
the southeastern species were provided by Gale.
Species of sect. DICHROMENA have sessile capitate inflorescences and whitish
spikelets often subtended by whitish bracts and lack perianth bristles. The
section is primarily neotropical in distribution and contains 23 species, of which
four are present in the United States. Three occur in the Southeast. Insect
pollination has evolved in plants of this section, as was first noted in 1893 by
De Lagerheim and later studied by Uphof and Leppik.
Thomas (1984) reviewed previous investigations of entomophily in species
of sect. DiCcHROMENA and made thorough field and laboratory studies. Fifteen
species of bees (Hymenoptera) visit flowers of plants included in this section.
The bees exhibit constancy, visiting four to ten inflorescences in a population
before leaving. The flowers have no fragrance and no nectar: the white color
of the bracts and spikelets attracts the bees, and pollen is the only reward. The
pollen grains have a sticky “‘pollenkit’’; thus, they aggregate and stick to the
bee’s body and legs. There is probably some transfer of pollen by the wind.
All species of sect. DICHROMENA are self-compatible. Thomas (1984) postulated
that the evolution of entomophily may have permitted the species to radiate
into shaded tropical forests, where a lack of air movement necessary for wind
pollination is compensated for by insect and self-pollination.
No species of Rhynchospora is gathered for food or medicinal uses. Several
species are detrimental weeds in rice fields, both in the Old World and in the
southeastern United States.
REFERENCES:
Phe tay references see BADEN ef al.; BEAL; BENTHAM; BERGGREN; BLASER es
194 CLARKE (1908, 1909): Erren (1976a); EyLes & ROBER TSON; FASSETT
tials & Woores: GONCHAROV et al.; Goon ef al.: eee HARBORNE et al:
HESLA et al.; aes Ho_m et al.; HoL_trum; Hotc ; G. E. Hurcuinson;
J. Hutcuinson; J. H. Kern; Koyama (1961); ee ex. (1969. 1986); KUNTH; LE
MaoutT & a ere & RAYNAL; MEEUSE; METCALFE; NAPPER (1964b); NEEs
VON ESENBECK; OGDEN; SCHULZE-MOTEL ( ae pe) SMITH et al.; STANDLEY; TEERI ef
al.; TORREY; VANHECKE; and VAN DER VEKE
1987] TUCKER, CYPERACEAE 415
Under Eleocharis see KUKENTHAL.
Gate, 8. Rhynchospora section Eurhynchospora in Canada, the United States, and the
West Indies. Rhodora 46: 90-134, 159-197, 207-249, 255-278. 1944. [The basic
monograph; distribution maps, descriptions, keys, and illustrations. ]
Gorpon-Gray, K. D., & L. L. BANbu. Silica deposits in Rhynchospora species. Proc.
Electron Microscop. Soc. S. Afr. 8: 83, 84. 1978.
AL Observations on new kinds of silica deposits in Rhynchospora
microcarpa, SEM photographs of achenes.
Alphabetisches Verzeichnis fiir Rhynchospora Vahl. 19 pp. San Isidro, Argen-
tina. 1981. [Index to generic and specific names, including synonyms, for KUKENTHAL
(1949, 1950, 1951).]
_ Contribucidn al estudio del género Rhynchospora Vahl (Cyperaceae) IV: R.
iberae, nueva especie de América Austral. Darwiniana 24: 469-473. 1982. [New
species related to R. californica Gale; cross sections of leaves and SEMs of achenes.]
Hitt, E. J. The perianth of Rynchospora capillacea var. leviseta. Rhodora 8: 186, 187.
19
Hoim, T. Studies in the Cyperaceae. VI. Dichromena leucocephala Vahl, and D. latifolia
Baldw. Am. Jour. Sci. 154: 298-305. 1897. [Taxonomic history; morphology and
anatomy. ]
Krat, R. A new species of Rhynchospora (C yperaceae) from southwestern Georgia.
Sida 7: 42-50. 1977. [R. Thornei, from Baker Co.; illustrations, key to new species
and relatives: R. divergens, R. pusilla, R. rariflora, and R. stenophylla. |
LAGERHEIM, M. G. pe. Note sur un Cypéracée entomophile. Jour. Bot. (Morot) 7: 181-
3. 1893
Leppik, E. E. Dichromena ciliata, a noteworthy entomophilous plant among the Cyp-
eraceae. Am. Jour. Bot. 42: 455-458. 1955.
Macsripe, J. F. Some Peruvian sedges. The status of Rhynchospora. Fieldiana Bot. 4:
RaAGonesE, A. M., E. R. GUAGLIANONE, & C. DIZEO DE STRITTMATTER. Desarollo del
pericarpio con cuerpos de silice de dos especies de RAynchospora Vahl (Cyperaceae).
(English abstract.) Darwiniana 25: 27-41. 1984. [Developmental study of the peri-
carp in R. corymbosa (L.) Britton and R. scutellata Griseb., emphasizing the origin
and differentiation of the silica bodies in the outer cell walls; line drawings and
EM
S.
TAKEDA, T., O. OeNu, & W. AGATA. The occurrence of C, species in the genus Rhyn-
chospora and its significance in kranz anatomy of the Cyperaceae. Bot. Mag. Tokyo
93: 55-65. 1980.
Upuor, J. C. T. Die Entomophilie der Cyperaceengattung Dichromena Michx. Ber.
Deutsch. Bot. Ges. 50: 208-214. 1932.
12. Dulichium Persoon, Syn. Pl. 1: 65. 1805.
Perennial herbs of swamps, fens, and shores. Roots fibrous; rhizomes hor-
izontal. Culms 1-3, terete, hollow, glabrous. Basal leaves bladeless; sheaths
appressed; cauline leaves several, the blades lanceolate, about 1-2 times longer
416 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
than the sheaths, auriculate, planar, with margins and midveins densely sca-
brellate abaxially; stomata confined to the adaxial surfaces (sometimes a few
present near the margins on the abaxial surface); chlorenchyma not radiate:
air cavities present. Inflorescences solitary in the axils of the upper leaves; rays
and rachises slender, compressed, scabrellate on the edges; spikes loosely ovoid,
appearing flattened from the distichous arrangement of the spikelets. Spikelets
3-20, linear-lanceolate, flattened; rachilla persistent, the internodes with hya-
line margins, the lowermost scale sterile (except in the terminal spikelet). Scales
3-9, deciduous as the achenes mature, lanceolate, conduplicate, acute, 5- to
9-nerved, the midveins scabrellate. Flowers perfect. Perianth bristles 6-9, I-
2 times as long as the mature achene, retrorsely barbed. Stamens 3: filaments
ribbonlike, nearly as long as the scales; anthers linear, the apices of the con-
nectives minute. Style capillary, glabrous; stigmas 2, about as long as the style,
glandular-pubescent. Achenes planoconvex, narrowly ellipsoid, the apex acute,
the base stipitate, the surface puncticulate. Embryos turbinate. Base chromo-
some number 16. Type species: D. arundinaceum (L.) Britton. (Name from
the Greek duo, two, and /eichon, scale, referring to the two-ranked scales of
the spikelets.)
A monotypic genus of wetland plants endemic to temperate North America.
Dulichium is easily distinguished from other Cyperaceae by its characteristic
distichous spikelet scales and its three-ranked cauline leaves. An interesting,
apparently uninvestigated feature of the plants is that in adjacent culms arising
from the same rhizome, the leaves are spiraled clockwise in one and counter-
clockwise in the next. The single species, D. arundinaceum, is distributed from
Newfoundland to southeastern Manitoba, south to southern Florida and eastern
Texas, and disjunctively in the area from northwestern Montana and south-
western British Columbia south, mostly west of the Cascades and the Sierra
Nevada, to central California (Wood, 1972, map). The genus had a wider
distribution during the Pleistocene when it occurred in Europe (Wood, 1971,
map). Fossils of this species are known from the Pliocene in the Soviet Union
(Daghlian). Infraspecific variation in fossil achenes from Europe has been stud-
ied by Truchanowiczowna.
Dulichium has usually been placed in the tribe Cypereae, near Cyperus.
Linnaeus (Sp. Pl. 1: 45. 1753) included the species in Cyperus, presumably
because of its distichous spikelet scales. The two genera differ, however, in
several important features: Dulichium has widely spaced axillary inflorescences
subtended by leaflike bracts with conspicuous sheaths, while Cyperus has api-
cally clustered inflorescence branches subtended by sheathless bracts; Duli-
chium has one sterile scale at the base of each spikelet, and Cyperus has two;
Dulichium has perianth bristles, but Cyperus does not.
The embryos of Dulichium resemble those found in Rhynchospora, rather
than those of any genus of the Cypereae (Van der Veken). A new monotypic
tribe, the Dulichieae, has recently been proposed for this genus by Schulze-
Motel (1959),
Plants of this genus have been neither reported to have economic use nor
noted as weeds.
1987] TUCKER, CYPERACEAE 417
REFERENCES:
Under family sine see BEAL; BENTHAM; BLASER (1940, 1941a, 1941b); CLARKE
(1908); Cook; DAGHLIAN; EyLes & ROBERTSON; FASSETT; GODFREY & WOOTEN; GOOD
et al.; HOTCHKISS; Kunst Le Maout & DECAISNE; LERMAN & RAYNAL; MATTFELD;
METCALFE: NEES VON ESENBECK; OGDEN; RADFORD ef al.; SCHULZE-MOTEL (1959, 1964);
TorReEY; and VAN DER VEKEN
Under Eleocharis see Woop.
BELL, F. G. Fossil ofan American sedge, Dulichium arundinaceum (L.) Britt., in Britain.
Nature 227: 629, 630. 1970. [[llustrations.]
ScHuLzE-MoTEL, W. Dulichieae, eine neue Tribus der Cyperaceae-Scirpoideae. Will-
denowia 2: 170-175. 1959.
TrALAu, H. Extinct aquatic plants of Europe. Bot. Not. 112: 385-406. 1959.
TRUCHANOWICZOWNA, J. Variability of the recent and fossil fruits of the genus Duli-
chium. (Polish and English summaries.) Acta Palaeobot. 14: 119-143. 1973.
Woon, C. E., Jk. Some floristic relationships between the southern Appa lachians and
western North America. Pp. 331-404 in P. C. Hott, ed., The distributional history
of the biota of the southern Appalachians. Part IJ. Flora. Behe Virginia. 1971.
[fig. 1, extant and known former distribution of D. arundinaceum
13. Schoenus Linnaeus, Sp. Pl. 42. 1753; Gen. Pl. ed. 5. 26. 1754.
Caespitose perennials of open sunny wetlands. Rhizomes short, oblique.
Culms terete, hollow, glabrous. Leaves all basal; sheaths tough, glossy, glabrous,
ligule lacking; blades linear, subcylindrical, upper surface flat or broadly convex;
stomata paracytic, on both surfaces [mostly adaxial]; chlorenchyma not radiate.
Inflorescences terminal, sessile, capitate [diffusely branched]; bracts 1 or 2,
oblique to erect, sheathless or essentially so, basally expanded and partly clasp-
ing the spikelets, distally linear; rays lacking. Spikelets (1-)10-25, oblong-
ellipsoid, flattened, the 2 or 3 basal scales sterile; rachilla wingless, more or
less deciduous at maturity. Scales distichous, 3-8, oblong, acute but not mu-
cronate, distally scabrellate, laterally nerveless, medially 1-nerved. Flowers
perfect. Perianth bristles lacking to 6, smooth or scabrellate. Stamens 3; fila-
ments ribbonlike; anthers linear, the apices of the connectives subulate, con-
spicuous; pollen grains 4-porate, obovoid, finely scabrate (pore areas frustillate).
Styles trigonous to subtrigonous, glandular; stigmas 3, capillary, shorter than
the styles, glandular. Achenes roundly trigonous to subterete, ovoid to ellipsoid,
the apex broadly rounded, the base gradually tapered to a stipe, the surface
smooth or barely reticulate, glossy. Base chromosome number 20(?). TyPe
species: S. nigricans L.; see Britton & Millspaugh, Bahama FI. 56. 1920. (Name
from Greek schoinos, for a rushlike plant.)— BLACK-HEADED SEDGE.
A genus of about 80 species, mostly restricted to Australasia but with a few
occurring in Africa, Eurasia, and the New World. Schoenus nigricans L., 2n =
54, 55, is present in North America. It is common in southern Florida but rare
in the Florida Panhandle, where it grows in wet grasslands over limestone
outcrops; it also occurs in the southwestern United States in the mountains
and valleys of western Texas, southern California, and southwestern Nevada,
where it grows in marshes and thermal springs. It is also reported from the
West Indies, Europe, and Asia.
418 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Kikenthal published a worldwide revision of Schoenus, and the genus has
received little subsequent systematic attention. The European species have been
investigated ecologically. Schoenus nigricans requires aluminum ions for growth,
and its range in the blanket bogs of the British Isles is thus limited to the coastal
region of western Ireland.
Plants of the genus have little economic significance. Wet meadows domi-
nated by Schoenus ferrugineus L. are mowed for fodder in northern and central
Europe. The species is adapted to low nutrient levels and is quickly displaced
by grasses when fertilizers are regularly applied.
REFERENCES:
Under family references see BENTHAM; BERGGREN; CLARKE (1908, 1909); GopFREY
& WooTEN; GONCHAROV ef a/.; HARBORNE; HARBORNE ef al.; J. HUTCHINSON; KUKKONEN
(1986); KuntH; LE Maour & DecaisNe; LERMAN & RAYNAL; METCALFE; NEES VON
ESENBECK; SCHULZE-MOorEL (1959, 1964); Torrey; and VANHECKE.
GANZERT, C., & J. PFADENHAUER. Seasonal dynamics of shoot nutrients in Schoenus
Prrscines (Cyperaceae). Holarctic Ecol. 9: 137-142. 1986. [Seasonal patterns of
1omass in an annually mowed calcareous fen in southern Germany; peak standing
crop in mid-July.]
KUKENTHAL, G. Vorarbeiten zu einer Monographie der Rhynchosporoideae. Schoenus.
Repert. Sp. Nov. 44: 1-32, 162-195. 1938. [Worldwide monograph; keys, descrip-
uons; 83 species.]
SPARLING, J. H. The occurrence of Schoenus nigricans L. in blanket bogs. I. Environ-
mental conditions affecting the growth of S. nigricans in blanket bogs. Jour. Ecol.
55: 1-13. 1967a. II. ae on the growth of S. nigricans under controlled
conditions. /bid. 14-31.
WHEELER, B. D. An Sa nee of Schoenus ferrugineus L. in Scotland. Watsonia
14: 249-256. 1983. [Autecology of a rare species
14. Cladium P. Browne, Civ. Nat. Hist. Jamaica, 114. 1756.
Stoloniferous, single-stemmed or loosely clustered, medium to large peren-
nials of sunny wetlands. Culms terete, roundly trigonous, or thickly crescen-
tform, hollow, glabrous. Leaves all cauline; sheaths glabrous, much shorter
than the blades; blades flat or slightly conduplicate to subinvolute, the margins
and midveins sparsely scabrellate to harshly scabrous; chlorenchyma not ra-
diate; alternate bundles inverted. Inflorescences pedunculate, terminal or both
lateral and terminal, diffusely branched; bracts leaflike but with shorter blades:
primary rays terete, wirelike and slightly drooping, glabrous; secondary rays
similar to primary but shorter and more slender; tertiary and quaternary rays
regularly produced in some species, these subtended by lanceolate scalelike
bracts and sheathing prophylls (involucels). Spikelets in glomerules of 1-S,
narrowly ellipsoid to lanceolate; rachilla wingless. Scales 3-5, the basal 1-3
sterile, ovate to oblong-lanceolate. Flowers perfect or imperfect (the distal
flower ofa spikelet perfect, the subdistal staminate). Perianth lacking. Stamens
2 or 3; filaments about as long as the subtending scale, flattened; anthers linear,
the apices of the connectives subulate; pollen grains 4-porate, narrowly obovoid
(sometimes with a peculiar apical appendage containing the degenerate nuclei),
scabrate. Styles subtrigonous, glabrous; stigmas 3, longer than the styles, glan-
1987] TUCKER, CYPERACEAE 419
dular. Achenes terete, ovoid, the apex broadly round (the withered style base
sometimes persistent), the base truncate and impressed, sometimes stipitate,
the surface smooth or nearly so. Embryos small, broadly obovoid, scarcely
differentiated (the first leaf not developed). Base chromosome number 20. TyPE
species: C. Mariscus (L.) Pohl (Schoenus Mariscus L.; see Britton & Brown,
Illus. Fl. No. U. S. Canada, ed. 2. 1: 347. 1913). (Name from Greek c/ados,
branch, referring to the highly branched inflorescences.)— TwiG-RuSH,
SAW-GRASS
Cladium is here accepted in the strict sense —i.e., consisting of three species:
C. Mariscus, C. mariscoides (Muhl.) Torrey, and C. jamaicense Crantz. Ku-
kenthal treated the genus more broadly, including Machaerina Vahl. Recent
studies by Vanhecke and Metcalfe argue against such a broad circumscription.
Species of Cladium consistently differ from those of Machaerina in their small-
er, less differentiated embryos and their isobilateral leaves with inverted bun-
dles (illustrated by Metcalfe).
wo species occur in our area. Cladium jamaicense, the saw-grass of the
Florida Everglades, grows in tidal marshes and coastal wetlands from eastern
Virginia to Mexico and the West Indies. Some authors (Kiikenthal, Raynal)
included C. jamaicense in the European C. Mariscus; Kern also included the
Australasian C. procerus S. T. Blake. The second species in our area, C. ma-
riscoides, occurs in brackish wetlands and inland fens and marshes from New-
foundland to Saskatchewan to Florida and Missouri; it is rare in the Southeast.
Raynal, without discussion, treated C. mariscoides and C. jamaicense as syn-
onyms of C. Mariscus, an extreme view not followed by anyone else.
Cladium jamaicense is important as the dominant species of much of the
Florida Everglades. The culms and leaves of C. Mariscus are gathered and
used in the manufacture of paper products in the Danube Delta, Romania.
REFERENCES:
Under family references see BEAL; BENTHAM; CLARKE (1908); aaeaioey & HARBORNE;
ERDAS pie eee ERTSON; FASSETT; GODFREY & WOOTEN; Goon et al.; HARBORNE;
a tTTUM; HoTcHkiss; G. E. HUTCHINSON; J. eee J. H. KERN;
KUKKONEN ee: rene & RAYNAL; MEEUSE; METCALFE; SCHULZE-MOTEL (1959,
1964); Torrey; and VANHECKE.
Conway, V. M. Biological flora of the British Isles: Cladium Mariscus (L.) R. Br. Jour.
Ecol. 30: 211-216. 1942.
DeviLiez, F., & J. R. DeSLtoover. Influence de prétraitements chauds et froids sur
germination des graines de Cladium Mariscus. (English summary.) Bull. Soc. Bot.
Belg. 113: 45-58. 1980. [Warm followed by cold pretreatment gives best results.]
GuICHARD, A. Sur l’existence de faisceaux libéro-ligneux 4 l’orientation inverse dans la
feuille végétative de oem MGS ba ri Compt. Rend. Acad. Sci. Paris 187:
509-511. 1928a. [Ilust pti finverted vascular bundles in leaf blades.]
Or rigine, parcours et torsion des faisceaux libéro-ligneux inverse du Cladium
Mariscus P. Br. Ibid. 567-569. 1928b. [Illustrations; basipetal differentiation of
vasculature in leaf blades.]
KUKENTHAL, G. Vorarbeiten zu einer Monographie der Rhynchosporoideae. XI. 10.
Cladium Crantz [sic]. Repert. Spec. Nov. 50: 1-17, 139-193. 1942. [Worldwide
revision of the genus in the broad sense; 47 species; C. jamaicense treated as sub-
species of Eurasian C. Mariscus.]
420 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
PFADENHAUER, J., & U. Eska. Untersuchungen zum Nahrstoffhaushalt eines Scheidried-
Bestandes (Cladietum marisci). Veréff. Geobot. Inst. Riibel — 309-327. 1986.
[Ecological study; maximum aboveground biomass in September. ]
RAYNAL, J. Notes cypérologiques 17. Révision des Cladium P. ee s. lat. (Cyper-
aceac) de Madagascar et des Mascareignes. Adansonia, II. 12: 103-112. : 72,
Rupescu, L. The use of sawgrass for paper product manufacture: an examinat f
properties. Pp. 191-195 in J. TourBIER & R. W. PIERSON, JR., eds., Acie
of water pollution. Philadelphia. 1976.
STEWARD, K. K. Physiological, edaphic, and environmental characteristics of typical
stands of sawgrass. Aquat. Ecol. Newsl. 9: 22, 23. 1976. [Tested for use in filtering
waste water; only 12 percent of phosphorus incorporated by plants; system saturated
after just eight weeks. ]
& W.H. Ornes. The autecology of sawgrass in the Florida Everglades. Ecology
56: 162-171. 1975.
Subfam. CARICOIDEAE Pax, Bot. Jahrb. 7: 307. 1886.
Tribe SCLERIAE Kunth ex Fenzl in Endlicher, Gen. Pl. 2: 114. 1836.
15. Scleria Bergius, Sv. Vet.-akad. Handl. 26: 142. 1765.
Small to medium, erect [scandent], perennial or annual herbs of grasslands,
open woods, fens, and shores. Roots fibrous; rhizomes regularly present in
many species, indurate, sometimes tuberlike, simple or branched. Culms trigo-
nous, glabrous, pubescent, or scabrellate [retrorsely scabrous], sometimes bul-
bous basally. Basal leaves bladeless or nearly so. Cauline leaves several; sheaths
3-angled, glabrous or more often scabrellate or pubescent; blades lanceolate to
linear or filiform, flat to slightly conduplicate [involute or thickened], glabrous,
scabrellate, or pubescent; chlorenchyma not radiate. Inflorescences paniculate,
1 to several, terminal or lateral and terminal; bracts leaflike but shorter than
or equaling the cauline leaves; rays trigonous, scabrellate on the angles or
smooth, secondary rays regularly produced in some species. Spikelets 1-6,
lanceolate to linear or oblong. Scales 1-6, ovate-deltoid, acute, mucronulate
to cuspidate, conspicuously medially |-nerved, laterally nerveless, glabrous or
pubescent. Flowers imperfect; carpellate flower(s) | (or 2), borne at the base
of the spikelets or in separate spikelets. Perianth bristles lacking. Stamens 1-
3; filaments capillary; anthers narrowly ellipsoid to linear, the apices of the
connectives frequently prolonged as slender, subulate, reddish appendages;
pollen grains uniaperturate, obovoid to subspheroid, psilate. Hypogynium, if
present, pebbled or warty, entire or with 3 acute to obtuse [truncate or acu-
minate], ciliate or glabrous lobes clasping the base of the achene. Styles slender,
glandular; stigmas 3, capillary, shorter than the styles. Achenes roundly tri-
gonous to terete, globose to ellipsoid, the apex broadly rounded (sometimes
apiculate), the base sessile to broadly stipitate, the surface smooth, reticulate,
trabeculate, rugose, glabrous, or pubescent. Base chromosome number 7(?).
Type species: S. flagellum-nigrorum Berg.; see Britton & Brown, Illus. Fl. No.
U.S. Canada, ed. 2. 1: 348. 1913. (Name from Greek sk/eros, harsh, the culms
of the type species being bound together into whips for beating slaves in Sur-
inam; often incorrectly said to be derived from Greek sk/eria, tough, in reference
to the achene walls; see Holm, 1898).—Nurt-RUusH.
1987] TUCKER, CYPERACEAE 421
A predominantly tropical genus of some 200 to 225 species. Centers of
diversity are tropical South America, tropical Africa, and southeastern Asia.
Twelve species occur in the United States, all east of the Great Plains. All are
present in the Southeast. Several range northward into northeastern North
America, reaching Massachusetts, southern Ontario, and southern Minnesota.
Two of our representatives occur southward into the West Indies. Many of our
species are endemic, as are most other taxa of Scleria. Many African species,
for example, occur only in Africa, and several are restricted to a single country
or are known from only one collection. Such endemism contrasts with the
distribution of the other large, mostly tropical genera of the family, such as
Cyperus, in which about one-fifth of the species are pantropic. Only two species
of Scleria, C. lithosperma (L.) Sw. and S. hirtella Sw., are reported from both
the Old World and the New.
The morphology of the achenes and the hypogynia has traditionally provided
the chief criteria for the circumscription of species. Core noted that some
species—for example, the South American Scleria leptostachya Kunth—pro-
duced both smooth and verrucose achenes, sometimes within a single collection
and sometimes within the same inflorescence. Nelmes (1955, 1956) reported
similar problems with certain African species, and he relied on features of the
rhizomes, ligules, and inflorescence (in addition to achene morphology) in his
classification of the African species. The hypogynium is apparently derived
from receptacular tissue, as is shown by its vascularization (Blaser, 1940, 1941b).
Robinson (1966) indicated that many of the southern African species of
Scleria are strong calcicoles. This autecology contrasts with that of the Amer-
ican species, most of which grow in acidic coastal plain habitats. Apparently
only one American species, S. nitida Willd. (which Fairey treated as a synonym
of S. verticillata) is a calciphile (Fernald).
Core recognized five sections in the genus, of which two, sects. SCLERIA (sect.
Euscleria Endl.) and HypoporuM (Nees) Endl., are represented in the Southeast.
In sect. Hypoporum the species have androgynecandrous spikelets (carpellate
flowers below the staminate) and lack hypogynia. There are five species in our
area: S. verticillata Willd., S. hirtella Sw., S. Baldwinii (Torrey) Steudel, S.
georgiana Core, and S. lithosperma (L.) Sw. Species of sect. SCLERIA have
unisexual spikelets and three-lobed, entire hypogynia. In our area this section
includes seven species: S. triglomerata Michx., S. minor Stone, S. oligantha
Michx., S. ciliata Michx., S. pauciflora Willd., S. Curtissii Britton, and S.
reticularis Michx.
genus is scarcely known cytologically. Reports are available only for
Scleria tesselata, 2n = 28, of southeastern Asia. This suggests the base number
x = 7 for the genus.
Species of Scleria have unusual embryological features (Nijalingappa). In S.
foliosa A. Rich. the embryos have both chalazal and micropylar haustoria.
Wall formation in the endosperm is complete in the Cyperaceae, except in
Scleria, where it is incomplete. The surface of the cotyledon is papillose in
Scleria but smooth in other genera of the family.
Robinson (1966) stated that several southern African species had “‘citrus-
scented” foliage; in fact, he used this as a lead characteristic in his key. Thus,
further investigation of the chemistry of these plants might be fruitful.
422 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
The fruits of Scleria triglomerata are dispersed by ants. The hypogynium
functions as an elaiosome (Gaddy). However, Robinson (1962) suggested that
the hypogynium provided buoyancy for the achenes of several southern African
species and was thus an adaptation for dispersal along water channels that
might later provide appropriate conditions for germination and growth of
seedlings.
No species of Sc/eria is gathered for food. Rhizomes of S. hirtella have been
employed medicinally in Colombia (Core). The tough, scabrous foliage of Sc/e-
ria 18 unsuitable for cattle forage. About ten species are noted as significant
weeds in Central and South America, tropical Africa, and southeastern Asia.
Scleria sumatrensis Retz. is a detrimental weed in Borneo (Holm et al.).
REFERENCES:
Under family references see BENTHAM; BLASER (1940, 1941b); CLARKE (1908, 1909);
CLIFFORD; EITEN (1976a); FERNALD; GADDY; HARBORNE; HARBORNE ef a/.; HOLM et al.:
Ho.tttum; HuAna; J. HuTcHINSson; J. H. KERN; KoYAMA; KUKKONEN (1969); LE MAoutT
& DECAISNE; LERMAN & RAYNAL; MEEUSE; METCALFE; Ses PER (1964b); O’NEILL;
SCHULZE-MOTEL (1959, 1964); STANDLEY; TEERI ef al.; and TorREY.
Core, E.L. The American species of Scleria. Brittonia 2: 1-105. 1936. [Basic monograph
for the New World species; keys, descriptions, representative specimens; illustrations
of achenes.]
Fairey, J. E., Ill. The genus Sc/eria in the southeastern United States. Catanea 32: 37-
7
of Seleria Ibid. 157: 5- 12. 1899. [Discussion of several southeastern species; illus-
tratio
NELMES, E. “Notes on Cyperaceae: XX XVIII. Scleria Berg. sect. Hypoporum (Nees) Endl.
in Africa. Kew Bull. 10: 415-453. 1955; XX XIX. African species of Scleria excluding
sect. Hypoporum. Ibid. 11: 73-111. 1956. [Keys, descriptions, discussions, specimen
citations; illustrations of many species.]
NJALINGApPA, B. H. M. Embryology of Scleria foliosa (Cyperaceae). Pl. Syst. Evol. 152:
219-230. 1986. [Illustrations.]
Rosinson, E. A. Notes on Sc/eria: I. The African species of sect. Tesselatae. Kirkia 2:
172-192. 1961; III. Scleria hirtella and some allied species. [bid. 4: 175-184. 1964.
. Scleria in Central Africa. Descriptions and notes: II. Ibid. 3: 8-14 ener
rovisional account of the genus Scleria Berg. (Cyperaceae) in t
Zambesiaca” area. Kew Bull. 18: 487-551. 1966. [Keys, descriptions, ee
Tribe CARICEAE Kunth ex Dumortier, Fl. Belg. 144. 1827.
16. Cymophyllus Mackenzie in Britton & Brown, Illus. Fl. No. U.S. Can. ed.
2. 1: 441. 1913
Loosely caespitose perennials of mesic montane forests. Rhizomes oblique.
Culms subterete, smooth, aphyllopodic. Leaves several; lowest with papery
sheath only, bladeless; ce laecelnes sheathless, the blade broadly lanceolate,
broadly rounded at apex, undulate at margins (especially so when dried), con-
spicuously multinerved but lacking a differentiated midvein and ligule. Inflo-
rescences single densely ellipsoid spikes, | per culm, terminal, with the pistillate
flowers below the staminate; bracts single broadly deltoid entire scales, | per
1987] TUCKER, CYPERACEAE 423
Figure 4. Cymophyllus. a-n, C. Fraseri: a, habit (portion of plant, leaf of preceding
season, plus new shoot terminated by inflorescence just past anthesis), x 1; b, detail of
undulate leaf margin, < 6; c, longitudinal section of inflorescence, staminate flowers
above, carpellate below, x 2; d, staminate flower with subtending scale, x 3; e, anther
(basifixed), x 12; f, 3 carpellate flowers enclosed in perigynia, re in axil of a scale
x 3; g, longitudinal section of perigynium to show carpellate flower ee ae
rachilla), x 5; h, stigma (note lack of papillae—species is insect pollinated), x 12;
longitudinal section of gynoecium to show single basal anatropous ovule, x 12; j, ai
stage of developing ee growth of gynoecium producing kink in style, x 5; k, perigynium
enclosing mature achene, x 6; 1, immature achene (note rachilla at base), x 6; m, achene,
x 6; n, embryo, eae from base of achene, 2 views, x 25
spike, broader than but otherwise like the pistillate scales immediately above
it. Flowers imperfect. Perianth lacking. Scales oblong-ovate, entire, co
conspicuous midvein or nerves. Stamens 3; filaments slender, 1-3 times as
long as the subtending scales; anthers slenderly ellipsoid, the eee not
424 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
prolonged. Perigynia 10-30, broadly ellipsoid, roundly trigonous, abruptly con-
tracted to a short, entire beak, weakly 20- to 30-nerved, glabrous; rachilla
filiform, “3-2 as long as the perigynium. Styles slender; stigmas 3, slightly
longer than the style, exserted from the beak of the perigynium. Achenes trigo-
nous, broadly ellipsoid, the apex broadly rounded, the base abruptly stipitate,
the surface smooth, glossy. Chromosome number unknown. TyPE SPECIEs: C.
Fraseri (Andrews) Mackenzie (Carex Fraseri Andrews; see Britton & Brown,
Illus. Fl. No. U. S. Canada, ed. 2. 1: 441. 1913.) (Name from Greek kuma,
wave, and phyllon, leaf, in reference to the undulate margins of the leaves.)
— FRASER’S SEDGE.
A monotypic genus endemic to the southern Appalachians. The sole species,
Cymophyllus Fraseri, 1s well known for its attractive white spikes that are
conspicuous when the plants flower in the spring. The plants grow in mesic to
somewhat damp soils in mixed hardwood forests, particularly on northern and
western slopes at middle elevations. The species is known from eastern Ten-
nessee and northwestern South Carolina, north through the Ridge and Valley
and Blue Ridge provinces to extreme south-central Pennsylvania (Somerset
County). Clarkson listed known collections arranged by state and county.
The systematic position of the genus has been disputed. Kiikenthal treated
the species as Carex Fraseri (sect. Leucocephali Holm of subg. Primocarex
Kiikenthal). Mackenzie, Fernald, Metcalfe, and Reznicek (pers. comm.) rec-
ognized Cymophyllus as a distinct genus. The conspicuous white inflorescences
of C. Fraseri, while unique among North American species of the tribe Cariceae,
are also known in at least one Old World species of Carex (C. baldensis L.).
White inflorescences are associated with insect pollination (discussed below)
and have evolved in Cyperus and Rhynchospora. In C. Fraseri there is a rachilla
within the perigynium. While a rachilla is not present in any temperate North
American species of Carex, it does occur in several other species (e.g., C.
microglochin Wahlenb. (boreal North America, cold-temperate Eurasia, south-
ern South America, fide Fernald)). Anatomical evidence (summarized by Met-
calfe) gives the strongest support for the generic status of Cymophyllus. In
Cymophyllus Fraseri the culms are terete (trigonous (rarely hexagonal) in Car-
ex); the leaves lack ligules (which are always present in Carex); the uppermost
leaf lacks a sheath and consists of blade only (sheaths are always present in the
cauline leaves of Carex); the large leaf blade is broadly rounded apically (acute
in Carex) and lacks the differentiated midrib and the adaxial layer of bulliform
cells typical of Carex (Holm; Metcalfe). In Cymophyllus Fraseri the median
vascular bundle has an incomplete adaxial sclerenchyma cap, and there is an
abaxial sclerenchyma girder (Metcalfe). The presence of perigynia in Carex
and C'ymophyllus clearly indicates that they are closely related, although it is
unclear how. The presence ofa rachilla in Cymophyllus suggests that this genus
might be closer to the Southern Hemisphere Uncinia Pers. than to Carex.
Cymophyllus Fraseri has long been suspected of being entomophilous (Clark-
son), although there has been only a single field study documenting entomophily
(Thomas). Four bee and one fly species were observed to visit spikes of this
species, which flowers from late April to mid-June. The insects collect pollen
1987] TUCKER, CYPERACEAE 425
for food and transfer it from plant to plant. They land on the lower, relatively
broad carpellate portion of the spikes, where they deposit pollen on the stigmas.
hey then crawl up to the anthers, collect pollen, and fly to another inflores-
cence. The pattern of stigmas first, then anthers, probably enhances outcrossing
(Thomas).
REFERENCES:
Under family references see BENTHAM, CLARKE (1908), FERNALD, METCALFE,
SCHULZE-MOTEL (1964), and TorREy.
CLARKSON, R. B. Fraser’s sedge, Cymophyllus Fraseri (Andrews) Mackenzie. Castanea
26: 129-136. 1961. [Ecology; summary of literature and known distribution. ]
Hom, T. Studies in the Cyperaceae, III. Carex Fraseri Andrews, a morpho logical and
anatomical study. Am. Jour. Sci. IV. 4: 121-128. pl. 1V. 1897. [Detailed description
with taxonomic Seat ae
Horn, G. S. vAN, & L. G. Wittiams. New county records for endangered and threatened
species in Tennessee. ne 46: 343-345. 1981. [C. Fraseri in Polk Co.
Jounson, R. H., & J. W. WaALLAcE, Jr. The flavonoid profile of Cymophyllus Fraseri
ieee? (Abstract.) Am. Jour. Bot. 73: 727, 728. 1986. [Contains methylated
apigen
ene G. Cyperaceae—Caricoideae. Jn: A. ENGLER, ed., Pflanzenr. IV. 20(Heft
38): 1-824. 1909.
MACKENZIE, K. K. Cyperaceae: Caricoideae. N. Am. Fl. 18(2, pts. 1-7): ‘1-478. 1931-
Rayner, D., ef al. Native vascular plants: endangered, threatened, or otherwise in
jeopardy in South Carolina. So. Carolina Mus. Bull. 4. 22 pp. 1979. [C. Fraseri
extirpated in South Carolina.]
Sims, J. Carex Fraseriana. Fraser’s carex. ie Mag. 33: no. 1391. 1811. [C. Fraseriana
Sims, a synonym of C. Fraseri Andrew
Tuomas, W. W. Insect pollination of C. ee Fraseri (Andrews) Mackenzie. Cas-
tanea ve 94, 95. 1984.
17. Carex Linnaeus, Sp. Pl. 2: 972. 1753; Gen. Pl. 280. 1754.
Caespitose or single-stemmed, small to medium-sized perennials of wet to
dry woods, grasslands, rock outcrops, pocosins, fens, bogs, marshes, and swamps.
Roots fibrous, smooth or pubescent; rhizomes (infrequently lacking) short and
oblique or long and horizontal, with closely appressed, lanceolate scales. Culms
loosely to densely clustered or solitary, fertile or both vegetative and fertile,
trigonous [hexagonal], the angles smooth or scabrellate. Basal leaves several to
many; sheaths smooth; ligule hyaline, glabrous; blades flat, conduplicate, pli-
cate, or involute, scabrellate (or smooth) on margins and midveins, sometimes
microscopically papillate on 1 or both surfaces, infrequently glaucous; stomata
paracytic, present on one or both surfaces; chlorenchyma not radiate; air cham-
bers frequently present; cauline leaves similar to basal ones but shorter and
fewer, sometimes lacking. Inflorescences simple or compound, monoecious
(rarely dioecious); bracts lacking or 1-6; spikes | to several, loosely to densely
ovoid to slenderly cylindrical, sessile or borne on simple [branched] erect to
pendent peduncles; each spike subtended by a leaflike or filiform basal bract;
spikes wholly carpellate or wholly staminate or gynecandrous or androgynous.
426 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
FiGure 5. Carex subg. VIGNEA: 8 species shown, each representing a different section
(A, Carex retroflexa (sect. PHAESTOGLOCHIN), B, C. vulpinoidea (sect. MULTIFLORAE); C,
C. decomposita (sect. HELEOGLOCHIN); D, C. laevivaginata (sect. VULPINAE); E, C. brun-
nescens subsp. sphaerostachya (sect. GLAREOSAE); F, C. bromoides (sect. DEWEYANAE);
1987] TUCKER, CYPERACEAE 427
Staminate scales lanceolate (the margins rarely fused basally), hyaline to char-
taceous, 1- (to 3-)nerved; carpellate scales lanceolate to broadly ovate, char-
taceous, 1- (to 3-)nerved. Flowers imperfect, protogynous or protandrous. Peri-
anth lacking. Stamens 3; filaments capillary or ribbonlike, longer than the
subtending scales; anthers broadly to slenderly ellipsoid; pollen grains 1- or
4-aperturate, obovoid or subspheroidal, psilate, trinucleate. Perigynia solitary
in the axils of carpellate scales, lenticular, subterete, trigonous, or slightly to
strongly compressed (beak, when present, less than to equaling or sometimes
longer than the body), coriaceous to chartaceous, the faces nerveless or with
1-15 nerves, minutely papillose or not, scabrellate or essentially smooth, dull
or glossy. Styles capillary, straight or curved; stigmas 2 or 3 [or 4], equaling or
exceeding the styles in length, smooth, papillose, or glandular, at anthesis
exserted through the orifice of the perigynia. Achenes lenticular or trigonous
[4-sided], ovoid to ellipsoid, 4 as long as to nearly as long as the body of the
perigynium, sessile or stipitate, apiculate or entire, the faces flat, convex, or
concave, the edges obtuse or acute (invaginate in a few species), the epidermal
cells translucent, opaque, or glossy. Embryos obconical, the radicle basal. Base
chromosome number 5. Type species: C. Airta L., not C. pulicaris L.; see
Hitchcock & Green, Prop. Brit. Bot. 187. 1929, and comments by Voss, Mich.
Bot. 11: 31, 32. 1972. (The classical Latin name, perhaps derived from the
Greek keirein, to cut, due to the sharp margins and keels of the leaf blades.)
A very large, cosmopolitan genus, reported to contain from 1000 to 2000
or even 2500 species (Standley, 1985a), including 165 that occur in the South-
east. Four subgenera have been recognized, = which two are Tepresented in
the United States. Subgenus INDocARExX Baillo branched,
branches subtended by tubular prophylls) eee about 50 species of the
Old World tropics. Subgenus VIGNEA (Lestib.) Kiikenthal (spikes all either
gynecandrous or androgynous, sessile, stigmas two, perigynia and achenes len-
ticular) includes about 500 species; it is worldwide in distribution but is most
diverse in the northern temperate and boreal regions. Subgenus Carex (subg.
Eucarex, spikes sessile or pedunculate, some exclusively staminate or pistillate,
stigmas 3 (rarely 2), perigynia and achenes trigonous) is the largest subgenus,
with about 800 species. Subgenus Primocarex Kiikenthal (spikes solitary,
terminal, stigmas 2 or 3, achenes lenticular or trigonous) is not represented in
our area.
The evolution of the tribe Cariceae is largely unclear. Due to shared features
of the inflorescences, Smith & Faulkner suggested that it arose from ancestors
akin to the Scleriae or the Hypolytreae. Kukkonen (1963), because of similar
G, C. Howei (sect. STELLULATAE); H, C. scoparia (sect. OVALES)). Four or 5 items illus-
=?)
face, x 10; 4, mature achene, abaxial surface, x 10; ‘5, longitudinal section of mature
perigynium and achene (C and D only), x 10.
428 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
FiGure 6. Carex subg. Carex (subg. Eucarex). a-i, C. gigantea: a, inflorescence,
uppermost 3 vies staminate, x 1/2; b, staminate flower and subtending scale, adaxial
view, x 5; c, perigynium in axil of subtending scale, stigmas of carpellate flower pro-
truding, x 5; d, carpellate flower (gynoecium), perigynium removed, x 5:e, ovule, ee
view, micropyle not visible, x 25; f, mature perigynium enclosing achene, x 5: g,a
with persistent style, x 5; h, longitudinal section of achene, seed coat not shown, pe
basal, endosperm above, x 5; i, seed removed from achene, x 5. j-r, C. glaucescens: J,
inflorescence, staminate spike uppermost, x '; k, staminate flower and subtending scale,
most of 2 stamens removed, x 5;1, perigynium and llat
flower protruding, x 5: m, oo flower, x 5; n, ovule, ena opye visible, “raphe
behind, x 25; 0, mature perigynium enclosing achene, x 5; p, perigynium, detail of
surface, Seats slbulat to ellipsoid cells that produce neeae effect, x 25; q, achene,
b 5.
1987] TUCKER, CYPERACEAE 429
infestations of smut fungi, indicated a probable close relationship with subfam.
Rhynchosporoideae; Koyama concurred with this opinion. A clearer under-
standing of generic relationships of the genus must await a better picture of
evolution within the genus. Its very large size and worldwide distribution
continue to hamper such studies.
Kikenthal believed that subg. PRIMOCAREX Kiikenthal was a most prim-
itive within the genus. A succession of more recent cyp
Nelmes; Koyama, 1962a; Le Cohu, 1968; Haines & Lye; ae & Faulkner;
Reznicek, 1986b) have taken the opposite view. In their opinion the unispicate
condition of subg. PRIMOCAREX was derived (perhaps polyphyletically) from
ancestors with richly branched inflorescences like those of subg. INDOCAREX.
However, the presence of a rachilla within the perigynium of some species of
subg. PRIMOCAREX suggests that it is the most primitive subgenus. Smith &
Faulkner believed that subgenera CAREX and VIGNEA might have evolved from
subg. INDOCAREX by reduction in inflorescence structure (a pattern also sug-
gested for several other genera of the family, e.g., Cyperus and Scirpus). This
would have involved loss of cladoprophylls (tubular prophylls subtending
branches) and reduction of branching. There are contrasting interpretations of
the inter- and infrageneric relationships in Carex.
The morphology of the inflorescences, particularly of the spikes and peri-
gynia, has traditionally been most heavily relied upon in distinguishing species
and circumscribing sections. Anatomical and cytological features are also tax-
onomically useful. Anatomical evidence has long been applied to the system-
atics of Carex. Crawford described the stems and leaf blades of the British
species. Akiyama presented a systematic study of the eastern Asian species,
emphasizing anatomical differences. Several recent revisions have included
anatomical descriptions of culms and leaves. Standley (1985a), in her mono-
graph of the northwestern species of sect. PHAacocystis Dum. (sect. Acutae),
showed that related species differ in the distribution of sclerenchyma and sto-
mata in culms and leaf blades. In certain species stomata are present on one
or both surfaces (Standley, 1986). The importance of anatomical features has
also been discussed by Le Cohu (1972) and by Metcalfe.
Recent studies with the scanning electron microscope have revealed an in-
teresting variety of surface features in leaves, perigynia, and achenes of Carex.
The presence of tubercles (Hoshino, 1986) and papillae (Maloney & Evans)
and the distribution of stomata (Standley, 1986) are useful in distinguishing
species and circumscribing sections.
Cytological studies have been helpful in Carex, but chiefly at the specific
level. Chromosome numbers in the genus range from 1 = 6 to n = 56. The
base chromosome number is 5, and the commonest haploid numbers in North
American species are 10, 20, 30, and 40 (Wahl). In many instances related
pairs of species differ in chromosome number. Aneuploidy is prevalent within
the genus. Aneuploid series characterize many sections (Wahl; Davies; Dietrich;
Faulkner, 1972). Polyploidy is infrequent.
The pollination biology of Carex has received little attention. Most species
are anemophilous. Honey bees and beetles visit inflorescences to gather pollen
430 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
and thus may also be vectors (Leppik). Self-compatible and self-incompatible
species have been noted in the genus (Faulkner, 1973; Handel, 1976, 1978a;
Schmid, 1984b). It is not known whether the incompatibility is sporophytic
or gametophytic. Handel (1976) determined that pollen-flow distances in C.
platyphylla Carey and C. plantaginea Lam. were rarely more than 10 m.
Little 1s documented about the dispersal of fruits of Carex. It has been
assumed that species with inflated perigynia are dispersed by floating on water,
but experimental verification is lacking. Several North American species (e.g.,
C. communis Bailey, C. umbellata Willd., and C. pedunculata Willd.) have
elaiosomes at the base of the perigynia and are dispersed by ants (Handel,
1978; Gaddy, 1986). Carex pauciflora Michx., widespread in northeastern
North America, has subulate perigynia that at maturity spring away from the
rachis (up to 60 cm) when touched (Hutton).
Flavonoid profiles can be used to distinguish between closely related species.
Toivonen (1974) showed this in the Fennoscandian representatives of sect.
CANESCENTES (sect. Heleonastes). Manhart (1985) demonstrated that classifi-
cations based on occurrences of flavonoids were similar to relationships de-
termined by morphology.
The species of Carex fall into three broad ecological groups with regard to
habitat: wetland, forest, and ruderal. In general the species of a section are
ecologically similar. Several sections (e.g., sects. PALUDOSAE G. Don and
LUPULINAE Carey) include mostly wetland species. Section Acrocystis Du-
mort., however, contains species of dry to dry-mesic open or wooded habitats.
Several sections (e.g., sect. ALBAE Ascherson & Graebner) are composed mostly
of calcicoles.
Most species of Carex are rhizomatous perennials. Carex is the only large
genus of the family containing no annuals. Certain species reproduce mostly
vegetatively (e.g., C. Bigelowii Torrey, plants of which set abundant seed, with
little germination or recruitment of seedlings unless disturbance occurs). In the
boreal C. flava L. seedlings persist for several years until competition is removed
(by disturbance or herbivory) and then grow rapidly to fill in the available
space (Schmid, 1986).
The economic importance of the genus lies chiefly in providing fodder for
domestic and wild mammals, especially in colder regions. Many Russian species
are important in this way (Goncharov et a/.); Carex stans Drejer and C. discolor
Nylander provide good grazing for cattle and reindeer. In Iceland, meadows
of C. Lyngbyei Hornem. are managed and yield up to five tons per hectare.
The nutritional content is very similar to that of common pasture grasses such
as Kentucky bluegrass, Poa pratensis L.
The following is a synopsis of the southeastern species, with chromosomal,
systematic, and ecological references. The order and circumscription of sections
generally follows Mackenzie (1931-1935).
Subgenus ViGNeA (Lestib.) Kiikenthal, represented in the Southeast by species
belonging to ten sections, is characterized by lenticular achenes, dorsiventrally
flattened perigynia, two stigmas, and both carpellate and staminate flowers in
each spike of the inflorescence.
Species of sect. AMMOGLOCHIN Dumort. (Arenariae Kunth, including sect.
1987] TUCKER, CYPERACEAE 43]
Divisae) are small rhizomatous plants of grasslands and strands. Two Eurasian
species, Carex arenaria L., n = 29, 58, 60, 64 (Noble) and C. divisa Hudson,
are naturalized in our area. Both grow on coastal sands from eastern Maryland
to eastern North Carolina. Several others occur in Canada and the western
United States, where C. Eleocharis Bailey is an important forage in the Rocky
Mountain region (Hermann, 1970).
Section MACROCEPHALAE Kiikenthal comprises two eastern Asian species,
one of which, Carex Kobomugi Ohwi, 2n = 84, 88, 1s sparingly naturalized
from eastern Virginia (Norfolk Co.) north to Cape Cod; it should be looked
for in eastern North Carolina. Standley (1985b) studied its population biology.
Although previous authors had described the species as dioecious, she showed
that individual rhizomes of a clone were consistently either staminate or car-
pellate (monoecious).
Section PHAESTOGLOCHIN Dumort. (sect. Bracteosae (Kunth) Pax) is one of
the most diverse sections of Carex in North America; it includes 16 species in
our area, all with ranges that extend into the northeastern United States or to
Canada. Plants of these species are mostly caespitose, with one to five sessile
androgynous spikes. Webber & Ball revised the C. rosea complex and corrected
the application of the names C. rosea and C. convoluta. Chromosome numbers
are known for six southeastern representatives of this section: C. sparganioides
Muhl., 2 = 23; C. cephalophora Muhl. ex Willd., n = 24; C. retroflexa Willd.,
n = 20: C. rosea Schkuhr (C. convoluta Mack.), n = 26; C. appalachica Webber
& Ball (C. radiata auct., non (Wahlenb.) Sm.); and C. radiata (Wahlenb.) Small
(C. rosea auct., non Schkuhr), n = 29. David & Kelcey summarized the biology
of the European species, C. muricata L., C. spicata Hudson, and C. divulsa
Stokes, all 21 = 58. Carex spicata and C. divulsa are naturalized in the North-
east south to Virginia. They might be found in North Carolina.
Section MULTIFLORAE (Kunth) Mack. contains three species in our area. The
commonest of these, Carex vulpinoidea Michx., n = 26, 27, is known from all
of the southeastern states and ranges north into southern Canada. It is also
sparingly naturalized in England (Clapham et a/.). The other southeastern species
are C. triangularis Boeck. and C. annectens Bickn. Both occur in most of the
southeastern states, but neither is as common as C. vulpinoidea.
Section HELEOGLOCHIN Dumort. (sect. Paniculatae G. Don (Hort. Brit. 367.
1830; non Carey) is represented in the Southeast by Carex decomposita Muhl.,
n = 30, 32, 33, which occurs in every state in our area. Plants of this section
are the only North American representatives of Carex with paniculate inflo-
rescences. Certain extraregional species of the section appear to be cytologically
conservative (cf. Clapham et al.). Carex diandra Schrank, 2n = 60, is circum-
boreal, while C. paniculata L., 2n = 60, 62, 64, and C. appropinquata Schum.,
2n = 64, are European.
In the Southeast, sect. VULPINAE (Carey) Christ is represented by five species
of swamps, marshes, and wet meadows. The plants resemble those of the
preceding two sections but are distinguished by their long, slender perigynia
(1 cm long in Carex crus-corvi Shuttlew.). In several species the bases of the
perigynia are conspicuously enlarged with aerenchyma, which probably makes
the fruits buoyant and allows dispersal by water. Carex crus-corvi, n = 26, C.
432 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
laevivaginata (Kikenthal) Mack., 1 = 23, and C. stipata Willd., n = 26, occur
throughout our area.
Section GLAREOSAE G. Don (sect. Heleonastes (Kunth) Ktikenthal) is a group
of circumboreal species of wet woods and bogs. The plants are small and have
few-flowered inflorescences. Three species, Carex brunnescens subsp. sphae-
rostachya (Tuckerman) Kalela, n = 27, 28, C. canescens L., n = 27, 28, and
C. trisperma Dewey, n= 30, barely reach our area from the north and are
found in the mountains of North Carolina and Tennessee.
Section STELLULATAE (Kunth) Christ consists of perhaps 30 species world-
wide. The plants are caespitose and have gynecandrous spikes of spreading to
reflexed perigynia with serrulate beaks. Reznicek & Ball (1980) revised the
North American species and provided excellent keys and descriptions. There
are seven representatives in our area. Carex Ruthii Mack. is endemic to high
elevations in the southern Appalachian Mountains from West Virginia to Geor-
gia. Carex exilis Dewey is primarily northeastern, occurring from Newfound-
land to Ontario south to Maryland; it is also known from widely disjunct
stations in central North Carolina, southern Mississippi, and southern Ala-
bama. The other southeastern species are C. atlantica Bailey (including C.
Mohriana Mack.), C. Howei Mack., n = 27, C. incomperta Bickn., n = 22,
and C. angustior Mack., n = 26. The two European species for which counts
are available have similar numbers: C. elongata L., 2n = 56, and C. echinata
Murray, 2n = 56, 58.
Species of sect. DEWEYANAE (Tuckerman) Mack. are probably closely related
to those of sect. STELLULATAE but have fewer, narrower, and appressed rather
than spreading perigynia (Reznicek & Ball, 1980). Carex bromoides Schkuhr,
n = 31+(A4), is the sole southeastern representative; it occurs in every state in
our area. Carex Deweyana Schwein., the only other species of the section, occurs
in northeastern North America.
Section OvALES (Kunth) Christ contains about 50 species in North America.
It is the largest section in our area, and the 16 representatives occurring in the
Southeast have flattened, papery, appressed perigynia in dense, ovoid spikes.
The section is taxonomically difficult and needs revisionary work. Several taxa
recognized by Mackenzie (1931-1935) have been synonymized by later work-
ers. Most of our species are widespread in eastern North America. For example,
Carex tribuloides Wahlenb., n = 35, and C. reniformis (Bailey) Small occur in
all the southeastern States, while C. argyrantha Tuckerman and C. aenea Fern.
are northeastern and just enter our area in the mountains of North Carolina.
Carex vexans Herm. is endemic to central and southern Florida. Among our
representatives, chromosome numbers are known only for C. tenera Dewey
(n = 26, 27, 28), C. straminea Willd. (n = 34+(3)), and C. cristatella Mack.
(n = 35). The type species, the European C. ovalis Good., 2n = 64, 66, 68, is
cytologically similar to eastern North American species of the section.
Subgenus Carex (subg. Eucarex Cosson & Germ.) includes the remaining
sections of the genus, 26 of which are represented in the Southeast. The plants
are characterized by differentiated spikes in which the terminal spike is wholly
staminate and the others are wholly or partly carpellate. Except in the distig-
1987] TUCKER, CYPERACEAE 433
matic sect. PHAcocystis Dumort. (sect. Acutae), the ovaries and achenes are
trigonous and there are three stigmas.
Section PoLYTRICHOIDEAE (Tuckerman) Mack. contains only Carex leptalea
Wahlenb., n = 26, an eastern North American endemic growing in damp,
mossy woods, often in calcareous soils, from Florida and eastern Texas north
into Canada. These are small, thin plants bearing few slender, beakless perigynia
and narrowly oblong, truncate achenes
Section PHYLLOSTACHYAE (Tuckerman) Bailey has four North American
species, characterized by androgynous spikes and staminate scales with basally
fused margins. Two, C. Jamesii Schwein., n = 35, and C. Willdenovii Schkuhr,
n = 31, occur in the Southeast. In addition to features of the perigynia, these
species are distinguished by the distribution of micropapillae on the leaves and
culms (Maloney & Evans).
Section Acrocystis Dumort. (Montanae (Kunth) Carey) comprises ten species
in the Southeast and nearly 30 worldwide. The plants grow in the most xeric
habitats of any species of Carex in our area, typically dry woodlands and rock
outcrops. They are small and tufted, with the leaves stiff, the carpellate spikes
few flowered, and the perigynia globose to ovoid, closely covering the roundly
trigonous achenes. Chromosome numbers are reported for half of our repre-
sentatives and indicate an aneuploid series: C. communis Bailey, n = 14, C.
nigromarginata Schwein., n = 17, C. artitecta Mack., n = 18, C. pensylvanica
Lam., n = 18, and C. /ucorum Willd. ex Link, n = 20. The European species
are more diverse cytologically (n = 9, 15, 19, 33) but are similar ecologically.
The fruits of C. artitecta (Handel, 1978) and C. nigromarginata (Gaddy, 1986)
are dispersed by ants.
Section PicrakE Kiikenthal has two representatives in eastern North America,
Carex picta Steudel and C. Baltzellii Chapman ex Dewey. Both are local, dry-
woodland species of the unglaciated eastern United States. Carex Baltzellii is
endemic to Georgia and northern Florida. Carex picta, occurring from southern
Indiana to Georgia and Louisiana, is a curious species. It is the only native
dioecious representative of Carex in our area. The plants form “fairy rings”
as the rhizomes branch and proliferate dichotomously (see Martens for illus-
tration). Clones from individual rhizomes are consistently staminate or car-
pellate, and carpellate plants do not always flower every year.
Section CLANDESTINAE G. Don (Digitatae (Fries) Carey) consists of four
species of the North Temperate Zone. The plants have purple leaf sheaths and
perigynia with minute beaks and tapered bases. Carex pedunculata Muhl.,
which grows on wooded, mesic, calcareous slopes, is the only representative
of the section in the Southeast. Elaiosomes are borne at the base of the perigynia,
which are dispersed by ants (Handel, 1976; Gaddy, 1986). Mackenzie (1931-
1935) included the only tetrastigmatic species of Carex, C. concinnoides Mack.
of the Pacific Northwest, in this section. St. John & Parker established subg.
Altericarex for this unusual species, but aside from its tetramerous carpellate
flowers, C. concinnoides fits in sect. CLANDESTINAE rather well, both morpho-
logically and ecologically.
Section TRIQUETRAE (Carey) Kiikenthal comprises five species of temperate
434 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
North America. The plants are caespitose, and they have greenish sheaths and
short-beaked, pubescent perigynia. There are two representatives in the South-
east, Carex dasycarpa Muhl. and C. tenax Chapman ex Dewey. Both grow in
pine forests, mostly from South Carolina to southern Mississippi. Another
species of the section, C. hirtifolia Mack., n = 22+(3)+(3), of the northeastern
United States, reaches its southern limit in the mountains of Virginia.
Section ALBAE Ascherson & Graebner consists of two species, both boreal
calcicoles of dry soils. One of these, Carex eburnea Boott, the only North
American representative, is a stoloniferous plant with glabrous perigynia that
is recorded in our area only from Tennessee. The second is C. alba Scop., 2n =
54, of Eurasia
Section PANICEAE G. Don (non Christ) is a Eurasian and North American
section of 12 species, five of which occur in the Southeast. The stoloniferous
plants have purple to reddish scales subtending the flowers, and ascending to
spreading, more or less ovoid perigynia. A member of this section, the rare
southern Appalachian endemic Carex Biltmoreana Mack., occurs on wet, shad-
ed cliffs in the Blue Ridge Mountains of North and South Carolina (Gaddy,
1983). Three of our representatives, C. Woodii Dewey, n = 22, 26, C. tetanica
Schkuhr, n = 26, and C. Meadii Dewey, are mostly northern in distribution
and just reach our area in the mountains of Tennessee and North Carolina.
The fifth species, C. Chapmanii Steudel, is endemic to the Coastal Plain be-
tween Florida and North Carolina. The European C. panicea L. and C. vaginata
Tausch have lower chromosome numbers: both are 2” = 32.
Section LAXIFLORAE (Kunth) Kiikenthal, containing about 25 species in east-
ern North America (17 1n our area), one in the western United States, and a
few in eastern Asia, is the most diverse section of Carex in our area. The plants
grow in woodlands; they are caespitose and bear conspicuously two-nerved
perigynia. Our species have recently been studied by Bryson, and Manhart
(1986) has investigated their cytology. Handel (1978a), who investigated the
pollination biology of Carex plantaginea Lam. and C. platyphylla Carey, re-
ported that both are self-compatible and that apomixis is absent. He studied
the dispersal of pollen by wind and found that pollen was transported twice as
far from C. plantaginea as from C. platyphylla. This difference was attributed
to the greater average height above ground of the staminate flowers in C.
plantaginea. An aneuploid series is evident in those southeastern representa-
tives of the section for which chromosome numbers have been reported: C.
Manhartii Bryson, n = 14, C. purpurifera Mack., n = 17, 18, 19, C. leptonervia
(Fern.) Fern., n = 18, 19, C. blanda Dewey, n = 18, 19, 20, 21, 22, C. gra-
cilescens Steudel, n = 20, C. laxiflora Lam., n = 20, C. laxiculmis Schwein.,
n= 22, 23, C. digitalis Willd., n = 24, C. plantaginea, n = 25, and C. platy-
phylla, n = 33, 34, 35. Carex striatula Michx. and C. laxiflora are myrme-
cochorous (Gaddy, 1986).
ection GRANULARES (O. F. Lang) Kiikenthal includes five eastern North
American species, of which four are found in the Southeast. They are calcicoles
and have few-flowered pedunculate spikes and perigynia with many fine nerves.
Carex granularis Muhl. ex Willd., » = 16+(4), occurs in all the southeastern
states and is the widest-ranging species of the section. The other southeastern
1987] TUCKER, CYPERACEAE 435
representatives are C. rectior Mack., C. Crawei Dewey, and C. microdonta
Torrey & Hooker.
To sect. OLIGOCARPAE (Carey) Kiikenthal (including sect. Griseae Bailey)
belong nine species of eastern North America, of which six are present in our
area. Members of this section are ecologically and morphologically similar to
plants of sect. GRANULARES but have lower chomosome numbers. Carex flac-
cosperma Dewey (C. glaucodea Tuckerman), C. oligocarpa Schkuhr, n = 27,
and C. grisea Wahlenb. (C. corrugata Fern.), 7 = 28, occur nearly throughout
our area and are also found in the northeastern United States.
Species of sect. HYMENOCHLAENAE (Drejer) Bailey (including sects. Sy/vaticae
Boott and Gracillimae (Carey) Kiikenthal) are widely distributed in the tem-
perate regions of the Northern Hemisphere and in the East African Highlands
(Kiikenthal; Mackenzie, 1931-1935). The plants have slender, drooping spikes
and often strongly beaked perigynia. A European representative of this section,
Carex sylvatica Hudson, 2n = 58, is naturalized in southern New England and
Long Island. There are six species in the Southeast. The eastern North American
representatives are currently being revised (with particular attention to cytol-
ogy) by Waterway (in prep.). Reznicek (1986a) has provided a detailed illus-
trated study of the Mesoamerican species. Chromosome numbers have been
reported for C. gracillima Schwein. (n = 5, 27), C. flexuosa Muhl. ex Willd.
(n = 27, 28), C. aestivalis Curtis (n = 28), and C. prasina Wahlenb. (n = 30),
all of which occur in the Southeast. Carex cherokeensis Schwein., wolf-tail,
reported from every state in the Southeast, and C. Sprengelii Dewey, n = 21,
of the northeastern United States, are sometimes segregated into sect. Longi-
rostres Kiikenthal because of their longer perigynial beaks.
Section VIRESCENTES (Kunth) Carey is represented in temperate North Amer-
ica, Eurasia, and the mountains of northern South America. The plants have
densely cylindrical, stiffly erect spikes. There are six species in eastern North
America, and all occur in the Southeast. Our representatives for which chro-
mosome numbers are known (Carex Bushii Mack., n = 24, C. hirsutella Mack.,
n = 26, C. Swanil (Fern.) Mack., n = 27, and C. virescens Muhl. ex Willd.,
n = 30) provide yet another example of the aneuploidy so frequent in the genus.
Species of sect. CAREX (sect. Hirtae (Tuckerman) Christ) are widespread in
the Northern Hemisphere, and a few are disjuncts in temperate South America.
The plants are stoloniferous and have three to ten spikes of ascending, ovoid
perigynia. The section has only two representatives in the Southeast: the North-
eastern and midwestern Carex lanuginosa Michx., n = 39, is known in our
area only from Arkansas, while C. striata Michx. (non C. striata Gilib., nom.
illeg., C. Walteriana Bailey), of the Coastal Plain, ranges from Georgia north
to southeastern Massachusetts. The type species of this section and of the genus,
C. hirta L., n = 56, is sparingly adventive in the northeastern United States
(south to the District of Columbia).
Section ANOMALAE Carey includes many species in eastern Asia and Aus-
tralasia, one in the western United States, and another in the eastern United
States, Carex scabrata Schwein., n = 27, recorded in our area from North
Carolina, Tennessee, and northern Alabama. Plants of this species have dense,
436 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
cylindrical carpellate spikes and perigynia with bidentate beaks; they are sto-
loniferous and typically grow near woodland springs.
The monotypic sect. SHORTIANAE (Bailey) Kukenthal contains Carex Shor-
tiana Dewey, an uncommon but attractive species of the Ohio River valley
south to central Tennessee. The plants have culms each bearing four or five
gynecandrous spikes of nerveless, corrugated perigynia with stipitate bases and
entire beaks.
The species of sect. PENDULINAE (Fries) Christ have a circumpolar distri-
bution and are characterized by pedunculate spikes and closely spaced peri-
gynia. The type species is the European Carex pendula Hudson, 2n = 58 or
60. The three representatives in our area, C. Joorii Bailey, C. verrucosa Muhl.,
and C. glaucescens Ell., are all widely distributed.
Species of sect. LimosAE (Tuckerman) Christ have drooping, few-flowered
spikes and broadly elliptic, beakless perigynia. Many are circumboreal in dis-
tribution and grow in fens, bogs, or wet woods. A single, primarily northeastern
representative, Carex Barrattii Schwein. & Torrey, from the mountains of
Tennessee and North Carolina, 1s known in our area. The type species is the
circumboreal C. limosa L., 2n = 56
The diverse and heterogeneous sect. ATRATAE (Kunth) Christ!’ contains many
species of the arctic and alpine tundra. The plants are characterized by sessile,
erect or drooping spikes, dark pistillate scales, and beaked or beakless perigynia.
There are many representatives in the southern Rocky Mountains (Hermann,
1970; Murray), but none of these is shared with our area. The single species
of our area, Carex Buxbaumii Wahlenb., n = 37, ca. 50, reaches its southern
limit in North Carolina and Arkansas.
Section PHAcocystis Dumort. (sect. Acu/ae Fries) is also a diverse circum-
boreal group. The plants are moderately large and have drooping spikes and
distigmatic, lenticular achenes. Three northeastern species, Carex strictior
Dewey, n = 34, C. stricta Lam., and C. torta Boott ex Carey, n = 33, reach
their southern limits in the northern half of our area. Standley (1985a) revised
the 15 representatives of this section in the Pacific Northwest. While none of
the species she treated occurs in our area, her thorough investigation of inter-
specific differences in leaf and culm anatomy, cytology, morphology, and some
aspects of ecology is informative and provides a model for future studies.
Species of sect. CRYPTOCARPAE (Tuckerman) Kiikenthal are mostly wetland
plants. They have drooping, densely flowered spikes and trigonous achenes.
Carex gynandra Schwein., C. Mitchelliana M. A. Curtis, and C. crinita Lam.,
n = 33, occur in the Southeast. These have been treated as a single taxon under
the last name, but there is good evidence for their specific status (Bruederle &
Fairbrothers, 1986). Carex gynandra and C. crinita hybridize rarely. The hy-
brids produce aborted achenes (Standley, 1983).
Section CoLLinsiAE Mack. contains a single species, Carex Collinsii Nutt.,
that grows in swamps on the Atlantic Coastal Plain from Georgia to Rhode
"Carex sect. ATRATAE (Kunth) Christ, Bull. Soc. Bot. Belg. 24: 15. 1885.
1987] TUCKER, CYPERACEAE 437
Island (Tucker, 1978). It is characterized by few-flowered inflorescences and
subulate perigynia.
Species of sect. FOLLICULATAE Mack. also have subulate perigynia, but the
spikes are densely many flowered and the plants are taller. There are two
representatives in the Southeast, Carex lonchocarpa Willd. ex Sprengel, found
throughout our area, and C. folliculata L., n = 28, a northeastern species
growing only in the mountains of North Carolina and Tennessee.
Species of sect. PSEUDO-CYPEREAE (Tuckerman) Christ are tall, paludal plants
of circumpolar distribution. They have drooping, slenderly cylindrical spikes
and densely arranged, conspicuously bidentate perigynia. There are two rep-
resentatives in our area, Carex Schweinitzii Dewey, n = 30, and C. comosa
Boott, n = 32. Carex pseudocyperus L., 2n = 66, is widespread in the Northern
Hemisphere and is believed to be native to New Zealand (Clapham et ai.).
Section PALUDOSAE G. Don” has eight species in North America and several
in Eurasia. The plants are stoloniferous and bear firm, many-nerved, slightly
inflated perigynia. There are two representatives in our area, Carex hyalinolepis
Steudel, found in wetlands throughout the Southeast, and C. trichocarpa Muhl.
ex Schkuhr, » = 55, a boreal bog species known in the Southeast only from
the mountains of North Carolina (Core).
Dense spikes of conspicuously inflated perigynia characterize members of
sect. SQUARROSAE Carey, which are endemic to eastern North America. There
are three species in our area, Carex Frankii Steudel, C. typhina Michx., and
C. squarrosa L., n = 28, each occurring in all or most of the southeastern states.
Section VESICARIAE (Tuckerman) Carey is a group of perhaps 20 species,
mostly of eastern North America and Eurasia. The plants generally grow in
shallow water and are characterized by inflated perigynia. Five representatives
occur in our area, but only one, Carex /urida Wahlenb., n = 32, 33, 1s common
(reported from every state). The others are C. Baileyi Britton, n = 34, C. bullata
Schkuhr, C. Elliottii Schwein. & Torrey, and C. rostrata Stokes, n = 34. The
type species, C. vesicaria L., n = 41, and C. riparia Curtis, n = 36, are cyto-
logically similar Eurasian representatives.
REFERENCES:
Under oe renee 7 BARNARD; BARROS (1935); BEAL; BENTHAM; BERGGREN;
BLASER (1941c¢ Ses 1909); CLIFFORD & HARBORNE; Cook: EYLEs
& eee FASSETT: ae (1986); Gipps; GODFREY & WOOTEN; GONCHAROV ef
al.; Goon et al.; HARBORNE; HARBORNE et a/.; HARRIS & MARSHALL; Hesse; HOLTTUM;
Huanc; G. E. on J. HUTCHINSON; KOYAMA (1962a); KRAL; LE MAoutT &
OBLE & MURPHY; OGDEN; PATCH; RAYNAL (1972); RIKLI; SAVILE; SCHULZE-MOTEL
(1959, 1964); Stace; STANDLEY; TEERI et al.; TiETZ; TORREY; and WINFREY & SAMSEL.
Under Rhynchospora see LEPPIK.
Under Cymophyllus see KUKENTHAL: MACKENzIE (1931-1935).
20Carex sect. PALUDOSAE G. Don in Loudon, Hort. Brit. 367. 1830, non (Fries) Christ (1884). TyPE
species: C. paludosa Good. (= C. acutiformis Ehrh.).
438 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
AKIYAMA, S. On the systematic anatomy of the leaves of some Japanese Carices. (In
Japanese; English summary.) Bot. Mag. Tokyo 55: os 130. 1941. [Cross sections;
anatomical characters useful in distinguishing species. ]
ANDERSON, L. C. The life history and ecology of Carex tie a and C. purpurifera
Mackenzie (Cyperaceae). (Abstract. ) ASB Bull. 25: 66.
ARNAL, C. Essai sur la répartition des sexes chez les Carex. rahe orca Sarav. Naturwiss.
Sci. 1: 102-114. 1952
Baitey, L. H. A preliminary synopsis of the genus Carex. Notes on Mexico, Central
America, and Greenland, with the American bibliography of the genus. Proc. Am
Acad. 22: 59-157. 1886.
Notes on Carex. XI. Studies of the types of the various species of the genus
Carex. Mem. Torrey Bot. Club 1: 1-85. 1889. oe of types of North American
species In major American and European her
Beti, K. L., & L. C. Buss. Autecology of Kobresia "Bellardil why winter snow accu-
mulation ae local distribution. Ecol. Monogr. 49: 377-402. 1979.
BERNARD, J. M. The life history of shoots of Carex lacustris. Canad. Jour. Bot. 53: 256-
975
The life history and population dynamics of shoots of Carex rostrata. Jour.
Ecol. 64: 1045-1048. 1977.
& E. GorHAm. Life history aspects of primary production in sedge wetlands.
Pp. 39-51 in R. E. Goon, ed., Freshwater wetlands. London. 1978.
& G. HANKINSON. Seasonal changes in standing crop, primary production, and
nutrient levels in a Carex rostrata wetland. Oikos 32: 328-336. 1979.
B. A. Sotsky. Nutrient cycling in a Carex /acustris wetland. Canad. Jour. Bot.
55: pe 638. 1977
Berry, E. W.. Fossil grasses and sedges. Am. Nat. 39: 345-348. 1905. [Including C.
Clarkii Berry from the Tertiary of Maryland.]
B6cHerR, T. G. A study of the circum polar Carex Heleonastes-amblyorhyncha complex.
Acta Arct. 5: 5-29. 1952.
Brown, L. E., & C. D. PETERSON. Carex rosea (Cyperaceae), Trifolium lappaceum
(Fabaceae), and Aira caryophyllea (Poaceae) new to Texas. Sida 10: 263, 264. 1982.
BRUEDERLE, L. P., & D. E. FAIRBROTHERS. Preliminary chemosystematic investigations
of the genus Carex (Cyperaceae): methods. (Abstract.) Bot. Soc. Am. Misc. Publ.
162: 86, 87. 1982. [Summary of techniques; no species mentioned.
& . Variability and taxonomic usefulness of achene a perigynium char-
acters of the mee crinita complex (Cyperaceae). (Abstract.) Am. Jour. Bot. 70(5,
suppl. 2): 107. 3.
ystematic aa of Carex Mitchelliana (Cyperaceae). (Ab-
stract.) Ibid. a part 2): 1 1984.
. Genetic sane in populations of Carex crinita Lam. (Abstract.)
Thid. 72: 944. 1985.
Allozyme variation in populations of the Carex crinita complex
(Cyperaceae). Syst. Bot. 11: 583-594. 1986. [Support for recognition of four species;
prevalence of in sucasen noted from low intrapopulational variability. ]
Bryson, C. T. A new species of Carex (Cyperaceae: sect. Laxiflorae) from the southern
Appalachians. pee 50: 15-18. 1985. [C. Manhartii, closely related to C. pur-
purifera.]
BurpetteE, J. I., & J. F. CLovis. Seem etc e ee of section Laxiflorae - the genus
Carex _ pumenee): taxonomy. Proc. Va. Acad. Sci. 41: 97-101. 1970.
a E. R., & G. A. BUCHANAN. Control . wolftail (Carex oe Schwein.)
a erennae pastures. So. Weed Conf. Proc. 20: 75-82. 1967. [Chemical control.]
ae T. V. Growth and population dynamics of Carex Bigelowii in an alpine
environment. Oikos 27: 402-413. 1976. [All reproduction observed was vegetative:
seedling establishment not noted.]
1987] TUCKER, CYPERACEAE 439
. Growth and translocation in a clonal Southern Hemisphere sedge, Uncinia
meridensis. es Ecol. 72: 529-546. 1984.
CLAPHAM, A. R . G. Tutin, & E. F. Warsurc. Flora of oe British Isles. ed. 2.
xlviil + ee pp. Cambridge, England. 1962. [Carex, 1073-1115.]
CLaustres, M. G., & M. C. LE Conu. Interprét des éléments épidermiques de la feuille
et de l’utricle dans la taxonomie de Carex. Compt. Rend. Acad. Sci. Paris, D. 260:
4373-4376. 1965.
Core, E. L. The range of Carex trichocarpa Muhl. Castanea 33: 151, 152. 1968. [North
Carolina mountains. |
CRAWFORD, F. he Anatomy of the British Carices. 124 pp. Edinburgh. 1910.
Crins, W. J., & P. W. BALL. The taxonomy of the Carex pensylvanica complex (Cy-
eG in North Am erica. Canad. Jour. Bot. 61: 1692-1717. 1983.
Cusick, A. W. Carex praegracilis: a halophytic sedge naturalized in Ohio. Mich. Bot.
23: 103-106. 1984.
CusseT, F., & T. T. H. ey La ligule de la feuille végétative des Carex. Bull. Soc.
Bot. France 112: 42-54. 1965.
Davin, R. W., & J. G. ae ae flora of the British Isles. Carex muricata L.
aggregate. Jour. Ecol. 73: 1021-1039.
Davies, E. W. Cytology, evolution, and ane of the aneuploid series in the genus Carex.
Hereditas 42: 349-365. 1956.
Dewey, C. Caricography: index to species. Am. Jour. ee I. 42: 1-10. 1866. [All taxa
described in DEwey’s papers between 1824 and |
DieTrRicH, W. Die Cytotaxonomie der Carex-Sektion oe in Europa. Feddes Repert.
75: 1-42. 1967
Drury, W. H., Jr. The ecology of the natural origin of a species of Carex by hybrid-
ization. Rhodora 58: 51-72. 1956.
Duman, M. G., & D. Kryszczuxk. Introgressive ees in the Carex stans-
bigelowii complex. Bull. Torrey Bot. Club 85: 359-362. 1958.
Duncan, W.H. Preliminary reports on the flora of Georgia— i Notes on the distribution
of flowering plants including species new to the state. Castanea 15: 145-159. 1950.
[Fourteen species of Carex. ]
aa Jes: asa studies - aia section Acutae in northwest Europe.
r. Linn. Soc. Bot. 65: 271-301.
: Exp erimental hybridization of ee European species of Carex section
Acutae (Cyperaceae). Ibid. 67: 233-253. 1973.
Gappy, L. L. Notes on the Biltmore sedge, a Biltmoreana Mackenzie (Cyperaceae).
Bull. Torrey Bot. Club 110: 530-532. 1983.
GiLtLy, C. L. Phylogenetic development . the inflorescence and generic relationships
in Kobresiaceae. Iowa State Coll. Jour. Sci. 26: 210-212. 1952. [Caricoideae suffi-
ciently distinct to merit familial rank—an idea not receiving any acceptance.
the northernmost occurrence of any species of this primarily austral genus. ]
Haines, R. W., & K. A. Lye. Studies in African Cyperaceae tes Panicle morphology
and possible relationships in Scleriae and Cariceae. Bot. Not. 125: 331-343. 1972.
HALLIDAY, G., & A. O. ae Studies in the Carex glareosa oe 1. Fruit shape.
Feddes Repert. 80: 77-92.
HANDEL, S. N. Dispersal sa af Carex pedunculata (Cyperaceae), a new North
American myrmechochore. Am. Jour. Bot. 63: 1071-1079. 1976.
. Self-compatibility in Carex ee las and C. platyphylia (Cyperaceae). Bull.
meas Bot. Club 105: 233, 234. 1978a.
w and ant-dispersed species in the genera Carex, Luzula, and Claytonia.
Canad. Jour. Bot. 56: 2925-2927. 1978b.
440 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Harvitt, A. M. Phytogeography of the Carices of Virginia. Rhodora 75: 248-257. 1973.
[One hundred species discussed; generally applicable to the Southeast.]
Hemporn, O. Zur Embryologie und Zytologie einiger Carex-Arten. Sv. Bot. Tidskr.
12: 212-219, 1918.
Chromosome numbers and dimensions, species-formation and phylogeny in
the genus Carex. Hereditas 5: 129-216. 1924
HERMANN, F. J. Manual of the Carices of the Rocky Mountains and Colorado Basin.
USDA Handbook 374. 397 pp. 1970. [Keys, descriptions, information on forage
value; illustration of each species. ]
: variety of Carex Bicknellii from Arkansas. Sida 5: 49. 1972
—. Manual of the genus Carex in Mexico and Central America. USDA Handbook
467. 219 pp. 1974. [Keys, descriptions; illustration of each species.]
HyeLMQuist, H., & E. NyHoLM. Nagra sae oer artkaraktarer inom Carex-gruppen
Distigmaticae. (English summary.) Bot. Not. 1947: 1-31. 1947. [Anatomy of disty-
lous Fennoscandian species; hybrids ae anatomical features of parents. ]
Ho iM, T. Studies in the Cyperaceae. I. On the monopodial ramification in certain North
American species of Carex. Am. Jour. Sci. III. 151: 348-350. 1 unnumbered pl.
1896a.
II. The clado- and anthoprophyllon in the genus Carex. Ibid. 152: 214-220.
1896b.
. XI. On the abnormal development of some specimens of Carex stipata Muhl.,
caused by Livia vernalis Fitch. Ibid. 158: 105-110. 1899. [Hemiptera: Psyllidae.]
HosuHIno, T. Karyomorphological and cytogenetical studies on aneuploidy in Carex.
Jour. Sci. Hiroshima Univ. Bot. 17: 155-238. 1981. [Forty species.]
. Acytotaxonomical study of Carex Paxii and two allied species. Jour. Jap. Bot.
61: 161- es 1986. [SEMs of perigynia showing large tubercles on |
moto. Geographical distribution of two cytotypes of Carex conica in
Seto Inland ‘Sea area of Japan. Jour. Jap. Bot. 54: 185-189. 1979.
T. Summizu. Cytological studies of degenerative nuclei at pollen development
of Carex ciliato-marginata. Bot. Mag. Tokyo 99: 185-190. 1986.
& R. TANAKA. Karyomorphological studies of Carex siderosticta and its two
allied species. Kromosomo 7-8: 191-194. ec
Howe, E. C. New York species of Carex. Ann. . New York State Mus. 48: 118-
202. 1895. [One hundred and thirty-three oe many useful comments on tax-
onomy.]
Hu ten, E. The amphi-Atlantic plants and their phytogeographical connections. Sv.
Vet.-akad. Handl. IV. 7: 1-340. 1958. [C. echinata, 140; C. comosa, 168; C. Bux-
baumii, 272.)
Hutton, E. E. Dissemination of perigynia in Carex pauciflora. Castanea 41: 346-348.
19
76.
Incvason, P. A. The golden sedges of Iceland. World Crops 21: 218-220. 1969. [Carex
Lyngbyei Hornem.
Jermy, A. C., & T. G. TuTin. British sedges. 199 pp. London. 1968. [Illustrated guide
to Carex in the British Isles.]
JoHNSON, W. M. Vegetative apomixis in Carex. Jour. Range Managem. 19: 305, 306.
1966. [Bulbil formation on rhizomes.
KakeLa, A. Uber die Kollektivart Carex brunnescens (Pers.) Poir. Ann. Bot. Fenn. 2:
174-218. 1965.
Kiriv’tseva, A. A., & A. M. BABAEvV. Drying up of Carex physodes and Carex pachystylis
in relation to weather and ecological conditions. (In Russian.) Ekologiia 2: 90-92.
*
Kreczetowicz, V.I. Are the sedges of subgenus Primocarex Kiik. primitive? Bot. Zhur.
21: 395-425. 1936. [No.]
1987] TUCKER, CYPERACEAE 44]
KUKKONEN, I. Taxonomic studies on the genus ao ee (Ustilaginales). Ann. Bot.
Fenn. 34: 1-118. 1963. [Infects certain species of Car
: ert morphology and anatomy of Uncinia Pers. (Cyperaceae). Kew Bull.
21: 93-97. 1967.
: eee anatomy of Carex microglochin Wahl. and Carex camptochochia
Krecz. Jour. Linn. Soc. Bot. 63(suppl. 1): 137-145. 1970.
Kuntu, C. 8. Uber die Natur des schlauchartigen Organs (Utriculus), welches in der
Gattung Carex das Pistill und spater die Frucht einhiillt. Arch. Naturgesch. Berlin
2: 349-356. 1835. [Perigynium believed to be a modified prophyll of a spikelet.]
Le Conu, M. C. Remarques sur l’inflorescence femelle des Carex: interprétation des
faits el al Bot. Rhedonica, A. 5: 37- 68.
——.. Hi e comparative de Carex rostrata Stokes. Ibid. 8: 65-72. 1970.
Les Saar épidermiques des Carex de la section Acutae. Compt. Rend.
Acad. Sci. Paris, D. 272: 2075-2077. 1971. [Sect. PHAcocystTis.]
. Apports de la microscopie électronique a reas 4 l’étude des ornementations
stomatiques des Carex. Ibid. 275: 349-352.
Examen a la microscopie électronique a eae des cénes de silice chez les
Cypéracées. Ibid. 277: 1301-1303. 1973.
Levyns, M. R. A comparative study of the inflorescence in four species of Schoeno-
xiphium and its significance in relation to Carex and its allies. Jour. S. Afr. Bot. 11:
79-89. 1945
LoHAMMAR, G. Wasserchemie und héhere Vegetation schwedischer Seen. Symb. Bot.
Upsal. 3: 1-252. 1938. [Distribution of Carex lasiocarpa, C. rostrata, and C. pseu-
docyperus with respect to pH and calcium concentration; evidence for strong niche
differentiation between these emergent aquatics in Swedish lakes.]
Love, A., & A. Levyns. Different chromosome numbers within the collective species
Carex polygama. Hereditas 28: 495, 496. 1942.
, D. Léve, & M. RayMonp. Cytotaxonomy of Carex section Capillares. Canad.
Jour. Bot. 35: 715-761. 1957. [Both polyploidy and aneuploidy important in evo-
lution of this section.]
MACKENZIE, K. K. No rth American Cancese. 2 oe ba pp. New York. 1940. [Full-
Dp ted in MACKENZzIE (1931—1935).]
—.. ” Keys to North American species of Carex ae North American Flora, vol. 18,
pts. 1-7. 80 pp. New York. 1941.
Mapore, S. S. A. An ecological study of the genus Carex in eastern subarctic Canada.
Bull. Torrey Bot. Club 78: 44-50. 1951. [Distribution with respect to soil pH of 35
species in the Val David region, Quebec
Ma tory, M. R., & D. K. Evans. Leaf an aon as a basis for so ae in selected
species of Carex Salen (Abstract.) ASB Bull. 25: 65.
& n in leaf anatomy of selected species Carex (Cyperaceae)
representing three re sections. (Abstract.) Ibid. 27: 47. 0.
Ma onty, A. C., & D. K. Evans. A taxon omic study of local die ons of Carex
Jamesii Schweinitz and Carex Willdenowii Schkuhr (Phyllostachyae: Cyperaceae)
in the Ohio River valley. (Abstract.) ASB Bull. 32: 72. 1985
Manuart, J. R. Foliar flavonoids of the North American members of Carex section
Laxiflorae Kunth. (Abstract.) Am. Jour. Bot. 72: 962. 1985.
Cytology of Carex purpurifera Mack. (Cyperaceae). Rhodora 88: 141-147. 1986.
[Supports recognition of C. Manhartii Bryson (” = 4) as a new species distinct from
C. Pe (n = 17, 18, 19).]
Martens, J. Some observations on sexual dimorphism in Carex picta. Am. Jour.
Bot. 26: fae 88. 1939.
Marx, P. S. Chromosome ee on Carex section Lupulinae. (Abstract.) Bot. Soc.
Am. Misc. Ser. 156: 68.
442 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Menapace, F. J., & D. E. Wusex. Scanning electron microscopy as an aid to sectional
placement of taxa within the genus Carex (Cyperaceae): sections Lupulinae and
Vesicariae. Micron Microscop. Acta 16: 213, 214. 1985.
MoOHLENBROCK, R. H., & J. SCHWEGMAN. A new species of Carex sect. Bracteosae.
Brittonia 2: re 79. 1969. [C. socialis, described from southern Illinois; illustration :
Moors, D. M., & A. O. CHATER. Studies of bipolar disjunct species. I. Carex. Bot. Not
124: 317- on 1971.
Moore, R. J., & J. A. CALDER. Some chromosom e numbers of Carex species of Canada
and Alaska. Canad. Jour. Bot. 42: 1387-1391. 1964.
Murray, D. F. Taxonomy of Carex sect. Atratae (Cyperaceae) in the southern Rocky
Mountains. Brittonia 21: 55-76. 1969.
NANNFELDT, J. A. The species of Anthracoidea (Ustilaginales) on Carex subgen. Vignea
with special regard to the Nordic species. Bot. Not. 130: 351-375. 1977.
NeLMES, E. Facts and speculation on phylogeny in the tribe Cariceae of the Cyperaceae.
Kew Bull. 1951: 427-436. 1952.
Nos e, J. " oe flora of the British Isles: Carex arenaria L. Jour. Ecol. 70: 867-
886
ey ie Bee, & J. L. Harper. The population biology of plants with clonal
grow wth. I. The apes and structural demography of Carex arenaria. Jour.
Ecol. 67: 983-1008. 1979.
. MARSHALL. i population biology of plants with clonal growth. IJ. The
nutrient strategy and modular physiology of Carex arenaria. Jour. Ecol. 71: 865-
877. 1983.
NovozHiLova, N. N. Flowering and pollination of Kohresia of eastern Pamirs. Nauk
Dokl. Vyssh. Shk. Biol. 10: 63-67. 1974
Ou, Y.C. Taxonomic study of epidermal patterns on some American species [of] Carex
using scanning electron microscope. Korean Electron Microscopy 10: 7-14. 1980.*
So AUER, J., Ee WENHOVEN. Néhrstoffokologie von Molinia coer ulea und
Ca tifor uf baumfreien yr des Alpenvorlandes. (English sum-
mary. ) Flora (Jena) 178: 157-166. 1986.
Popov, A. A. An epiphytic sedge. (In ee ) Priroda (Moscow) 1960(5): 1 112.
1960 i
PRINGLE, W. L., & A. L. vAN Ryswyk. Response of water sedge in the growth room to
a and temperature treatments. Canad. Jour. Pl. Sci. 45: 60-66. 1965. [C.
aquati
REZNICEK, A A. Carex section [ymenochlaenae in Mexico and Central America. Syst.
Bot. 11: zee 87. 1986a.
w hypothesis for evolution in Carex and the tribe Cariceae. (Abstract.)
a ican Boe 73: 783. 1986b.
ALL. The taxonomy of Carex series Lupulinae in Canada. Canad.
fae Bot. 52: 2387-2399. 1974. [Six species, all of which also occur in the Southeast:
keys, porate chromosome counts; distribution maps showing Canadian por-
tions of ranges o
&
The taxonomy of Carex section Stellulatae in North America north
of Mexico. Contr, Univ. Mich. Herb. 14: 153-203. 1980. [Eight species, of which
five occur in the Southeast; keys, descriptions, distribution maps, illustrations of
perigynia; helpful in clarifying the long-confused taxonomy of this difficult section. ]
& The sedge Carex /oliacea in eastern North America. Canad. Field
Nat. 95: 89— 92. 1981. [Taxonomy, morphology, distribution; maps, illustrations. ]
P.M. Catuinc. Sectional limits and relationships in Carex sections Carex,
Paludosae, and io in eastern North America. (Abstract.) Bot. Soc. Am. Misc.
Publ. 162: 104.
: Cas shoots in Carex (Cyperaceae). Taxon (in press).
Rosarps, A. W., D. T. CLARKSON, & J. SANDERSON. Structure and permeability of the
1987] TUCKER, CYPERACEAE 443
en ee layers of the sand sedge (Carex arenaria L.). Protoplasma
101: 331-347. 1979.
RosBerRTSON, A. Variations in Carex (sect. Stellulatae Kunth) in Newfoundland. (Ab-
stract.) Bot. Soc. Am. Misc. Publ. 158: 95. 1980.
Rocers, K. E. Notes of plants of Mississippi. I. Castanea 38: 199-203. 1973. [C. picta,
new state record.]
& F. D. Bowers. Notes on Tennessee plants. Castanea 34: 394-397. 1969. [C.
venusta var. minor Buckley.]
Roster, S. J., & J. M. BERNARD. Seasonal changes in carbohydrate levels in tissues of
Carex lacustris. Canad. Jour. Bot. 57: 2140-2144. 1979.
Rueccer, R. Plantes melliféres et polliniféres: les Carex. Jour. Suisse Apicult. 50:
L7G ho SS
Russet, G. E., & W. H. DuNcAN. An annotated checklist of Carex (Cyperaceae) in
Georgia. Castanea 37: 200-214. 1972.
St. Jonn, H. A new Carex (Cyperaceae) of the section Stel/ulatae. Hawaiian plant
studies 116. Pacific Sci. 37: 25, 26. 1983. [C. hawaiiensis.
. PARKER. 7 as species, section, and subgenus of Carex. Am.
Jour. Bot. 12: 63-68.
SAVILE, D. B. O., & J. A. en Phylogeny of — in the light of parasitism by the
smut fungi. Canad. Jour. Bot. 31: 169-174. 1953.
ScHMID, B. W. Karyology and hybridization in ~ Carex flava complex in Switzerland.
Feddes Repert. 93: 23-59. 1982.
Notes on the aaa and taxonomy of the Carex flava group in Europe.
Watsonia 14: 309-319.
——.. Niche width and see within and between populations in colonizing species
(Carex flava group). Oecologia (Berlin) 63: 1-5. 1984a.
ife histories in clonal plants of the Carex flava group. Jour. Ecol. 72: 93-114.
1984b.
. Colonizing plants with persistent seeds and persistent seedlings (Carex flava
group). Bot. Helvetica 96: 19-26. 1986. [Seedlings of C. viridula respond to decreased
competition faster than those of C. flava.
SERNANDER, R. Entwurf einer Monographie ao etumaes Myrmechochoren. Sv.
Vet.-akad. Handl. 41: 1-410. 1906. [C. digi
SHAH, C. K. Studies in germination. I. Carex W aun Priesc. Jour. Indian Bot. Soc.
41: 551-556. 1962.
SHEPHERD, G. J. Experimental arene in the genus Carex section Vesicariae. Unpubl.
Ph.D. Thesis. Univ. Edinburg 3.
The use of anatomical carters in the infrageneric classification of Carex
(Cyperaceae). Hoehnea 6: 33-5
SHETLER, S. G. A catalog of the eens ce (Cyperaceae). Smithson. Contr. Bot. 12:
26-184. 1973. [Useful index of all type specimens of Carex in major American
herbaria; separate lists arranged by species name, author, place of collection, and
date of publication.]
Smit, D. L. Development of the inflorescence in Carex. Ann. Bot. 80: 475-486, 1966.
The experimental control of inflorescence development in Carex. Ibid. 81: 19-
30. 1967.
The growth of shoot eee and inflorescences of Carex flacca Schreb. in aseptic
culture. Ibid. 82: 361-370. 1968.
—. The role of leaves a roots in the control of inflorescence development in
Carex. Ibid. 33: 505-514. 1969.
_S. FAULKNER. The inflorescence of Carex and related genera. Bot. Rev. 42:
53-81. 1976.
SNELL, R. S. Anatomy of the spikelets and flowers of Carex, Kobresia, and Uncinia.
Bull. Torrey Bot. Club 63: 277-295. 1936
444 JOURNAL OF THE ARNOLD ARBORETUM [vVOL. 68
STANDLEY, L. A. A clarification of the status of Carex crinita and C. gynandra (Cyper-
aceae). Rhodora 85: 229-241. 1983.
Systematics of the Acutae group of Carex (Cyperaceae) in the Pacific Northwest.
Syst. t. Bot. Monogr. 7: 1-106. 1985a. [Sect. Phacocystis; 15 species; keys, descriptions,
illustrations, chromosome counts.]
. Paradioecy and gender ratios in Carex macrocephala (Cyperaceae). Am. Midl.
Nat. 113: 283-286. 1985b.
. Variation of stomatal distribution in Carex aquatilis (Cyperaceae). Am. Jour.
Bot. 73: 1393-1399. 1986. [Var. aquatilis has stomata on both surfaces, while var.
dives (Holm) Kiikenthal has them on the upper surface only; discussion of ecological
implications. ]
. Taxonomy of the Carex lenticularis complex in eastern North America. Canad.
Jour. Bot. 65: 673-686. 1987. [C. lenticularis Michaux, n = 43, 44, and C. nigra
(L.) Reichard, n = 42 (sect. Phacocystis), of northeastern North America.]
StanT, M. Y. The shoot apex of some monocotyledons. I. Structure and development.
Ann Bot. 66: 115-128. 1952.
Stout, A. B. The individuality of the chromosomes and their serial arrangement in
Carex aquatilis. Arch. Zellforsch. 9: 114-139. 1912. [Two plates.
Svenson, H. K. Carex foenea, C. straminea, and C. albicans in Willdenow’s herbarium.
Rhodora 40: 325-331. 1938.
TALLENT, R. C., & D. E. Wujsek. Taxonomy of several Carex species using microm
phological characters. (Abstract.) Am. Jour. Bot. 70: 103. 1983a. [Surface rae
similar in all species of sect. Ovales; in sect. Extensae these features varied between
species. ]
&
Scanning electron microscopy as an aid to eras of sedges (Cy-
peraceae: Carex). Micron Microscop. Acta 14: 271, 272. b.*
TANAKA, N. Chromosome studies in Cyperaceae. IV. ea numbers of Carex
species. Cytologia 10: 51-58. 1939.
. Chromosome studies in the rte Carex, with special reference to aneuploidy
and polyploidy. /bid. 15: 15-29. 1949,
THIELKE, C. Gerbstoffidioblasten in ee Scheide von Carex. Protoplasma (Wien) 47:
145-150. 1956.
——. Uber Rane aici bei Cyperaceen. II. Entstehung von epidermalen
Faserbiindeln in der Scheide von Carex. Planta 49: 33-46. 1957.
THomas, W. W cere of the species of Carex in Michigan’s upland deciduous
forests: a key stressing vegetative sua Mich. Bot. 21: 131-139. 1982. [Includes
many species occurring in the South
TrETEMA, T. Ecophysiology of the sand ners Carex arenaria L. U1. The distribution of
'C assimilates. Acta Bot. Neerl. 29: 165-178. 1980.
TIMONEN, T., & H. Torvonen. Gross and re te comparison of Carex
furva and C. lachenalii. Ann. Bot. Fenn. 16: 11-17, 1979,
Torvonen, H. Chromatographic comparison of the . of Carex section ao
and some Carex canescens hybrids in eastern Fennoscandia. Ann. Bot. Fenn. 11:
225- . ca
—— on the nomenclature and taxonomy of Carex canescens (Cyperaceae).
Ibid. a a -97. 1981.
MONEN. Perigynium and achene epidermis in some species of Carex
subgenus aie (Cyperaceae), studied by scanning electron microscopy. Ann. Bot.
Fenn. 13: 49-59. 1976.
Tucker, G. C. Notes on the flora of Rhode Island. Rhodora 80: 596, 597. 1978.
verlooked sectional names in Carex (Cyperaceae) from Loudon’s Hortus Brit-
tanicus (1830), (Abstract.) Canad. Bot. Assoc. Bull. 20(3): 16. 1987. [Authority for
sectional names is ““G. Don in Loudon’’—see p. i
1987] TUCKER, CYPERACEAE 445
Vonk, D. H. Biosystematic studies of the Carex flava complex, 1. Flowering. Acta Bot.
Neerl. 28: 1-20. 1979.
Voss, E. G. Additional nomenclatural and other notes on Michigan monocots and
gymnosperms. Mich. Bot. 11: 26-37. 1972. [Includes discussion of lectotypification
prompted by a query by Dr. Carroll E. Wood”; C. hirta is noted as the
correct lectotype; C. pulicaris had been selected previously.]
Wau_, H. A. Chromosome numbers and meiosis in the genus Carex. Am. Jour. Bot.
27: 458-470. 1940.
Wacter, K. S. A preliminary study of the achene epidermis of certain Carex (Cyper-
aceae) using scanning electron microscopy. Mich. Bot. 14: 67-72. 1975. [Features
of pinion of sects. Vesicariae and Pseudo-cypereae; supports placement of C. /urida
in former.
WATERWAY, M. J. Allozyme variation within Carex section Sy/vaticae. (Abstract.) Can-
ad. Bot. Assoc. Bull. 20(3): 16. 1987. [Allozyme data support relationships based
on morphological criteria.]
Wepsser, J. M., & P. W. BALL. The taxonomy of the Carex rosea group (section Phaes-
toglochin) in Canada. Canad. Jour. Bot. 62: 2058-2073. 1984. [Three species, all
extending southward to our area; keys, descriptions, distribution maps.
Wuirtkus, R. Chromosome numbers of some northern New Jersey Carices. Rhodora
83: 461-464. 1981. [Includes southeastern species.]
cker. A contribution to the taxonomy of the Carex Macloviana
appregate (Cyperaceae) i in western Canada and Alaska. Canad. Jour. Bot. 62: 1592-
1607. 1984.
WIEGAND, K. M. Carex laxiflora and its relatives. Rhodora 24: 189-201. 1922.
PERRY, PINUS 447
A NEW SPECIES OF PINUS FROM MEXICO AND
CENTRAL AMERICA
J. P. Perry, JR.!
A new species of Pinus iS described from Mexico and Central America.
Throughout most of its range, associated species were often P. pseudostrobus,
and P. montezumae; field observations indicate that natural
hybridization probably occurs between the new species and these taxa.
While carrying out field studies on species of Pinus growing in Mexico and
Central America, I discovered a number of populations of the genus in Mexico,
Guatemala, El Salvador, and Honduras that appear to belong to an undescribed
species.
Pinus nubicola J. P. Perry, sp. nov. Ficures 1-4.
Differt a P. oaxacana et P. estevezii foliis 25-40 cm longis, in fasciculo 5 vel
6 (interdum 7), cernuis vel pendulis; et squamis 20-25 mm latis, ad apicem
prominentiis disparibus et umbone parva cum margine depressa instructis.
Tree 25-30 m tall, d.b.h. 0.5-1 m, when mature with open, rounded crown.
Spring shoots uninodal; branchlets puberulent, soon becoming glabrous; young
trees with smooth bark. Leaves in fascicles of 5 or 6 (occasionally 7, rarely 8),
25-43 cm by 0.6-1 mm, flexible, very drooping, margin serrulate; stomata on
all surfaces; hypodermis 2- to 4-layered, with many slight penetrations into
chlorenchyma; resin canals 3 or 4, medial (occasionally | internal); endodermis
with outer cell walls thickened; vascular bundles 2, distinct; fascicle sheaths
persistent, 20-30 mm long, pale brown, not resinous. Cones subterminal, | to
4 together, subsessile, reflexed, asymmetrically ovoid to long-ovoid, 10-15 by
8-10 cm when open at maturity, peduncle and few basal scales remaining on
branch when cone falls. Scales 20-25 mm wide, thick, stiff, with apex obtusely
angled, generally with distinct, unequal marginal projections, apophysis 5-8
by 20-22 mm, transversely keeled, abaxial surface raised more than adaxial,
the umbo ashy gray, central, 2-3 mm long, margins often slightly depressed,
generally curved upward, terminating in small, persistent prickle. Seeds brown
or spotted to mottled black, 5-7 by 4-5 mm, with detachable, pale brown wing
20-25 by 8-11 mm; cotyledons (7 or) 8 to 10 (to 13).
1306 North Front Street, Hertford, North Carolina 27944.
© President and Fellows of Harvard College,
Journal of the Arnold Arboretum 68: 447-459. ae 1987.
448 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
ABS.
eA
AST RS
AS ak 7
Neqnsts Sars
Ficure 1. Pinus nubicola: A, mature cone, foliage, and branchlet, showing nonde-
current bases of foliage bracts; B, seed and seed wing; C, cone scale, showing apophysis,
umbo with depressed margins, and apex with unequal projections.
Type. Guatemala, Depto. Guatemala, about 40 km E of San José Pinula on
dirt road toward Las Nubes, 90°20’W, 14°33’30’N, alt. 2000 m, 25 Feb. 1979,
Perry GUA.32-79 (holotype, GH; isotypes, CHAP, E, GH, K, MEXU, NCSC).
TURPENTINE ANALYSES. Most trees had relatively large amounts of heptane,
nonane, and a-pinene; many also had sizeable quantities of limonene, while a
few had a great deal of terpinene-4-ol and methyl] chavicol. Results of individual
analyses performed on 31 specimens from Mexico, Guatemala, and Honduras,
as well as approximate mean percent composition, are shown in TABLE 1.
PHENOLOGY. Flowering starting late January, but mainly February and March.
HABITAT AND DISTRIBUTION. Mexico to Honduras (see Map 1), on cool, moist
mountain slopes, 1800-2400 m alt. (see FiGure 2),
In Veracruz state, Mexico, Pinus nubicola was growing at 1800 m on the
1987] PERRY, PINUS 449
Ficure 2. Pinus nubicola growing on slope of Mt. El Pital, Depto. Chalatenango, E]
dor showing characteristically drooping foliage.
humid eastern escarpment of the Sierra Madre Oriental. Associated species
were Pinus chiapensis (Martinez) Andresen, P. pseudostrobus Lindley, P. oa-
xacana Mirov, and Liquidambar styraciflua L. In Mexico (Chiapas), at a some-
what drier site, associated species were P. oaxacana, P. pseudostrobus, P. mon-
tezumae Lambert, P. rudis Endl., P. patula var. longepedunculata Loock,’ Pinus
oocarpa var. ochoterenae Martinez, Pinus Se Ehrenb., and Quercus
spp. In Guatemala associated species were P. oocarpa var. ochoterenae, P.
montezumae, P. rudis, P. maximinoi Moore, P. ie eee (rarely), and P.
Styles (1976) pointed out that there has been considerable confusion in the literature and in the
field rare the ena nea HOD of Pinus oocarpa var. CRs ie P. He sae var. longepedun-
culata. t t his b P. patula ae
& Cham. Although there is sadeed a great deal a confusion regarding Eee ee, of the tw a,
I do not believe the matter has been clarified by referring both varieties to P. patula. J prefer, svi
the results of further studies, to use the original varietal classification of these taxa.
450 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Sata! s4e@uag
rm eT AT
. a, 2 i'n 2 2
Ory
get
:
Ry
aes
2.:. .
80x
At
4) 4 |
VY if
FiGurRE 3. Cross section of leaf of Pinus nubicola.
tecunumanti Eguiluz & Perry. In El Salvador the species was found growing
with P. maximinoi, P. oaxacana, P. ayacahuite, P. oocarpa var. ochoterenae,
P. tecunumanii, Abies guatemalensis var. tacanensis Martinez, Cupressus lu-
sitanica Miller, Liquidambar styraciflua, and Quercus spp. In Honduras as-
sociated species were P. maximinoi, P. oaxacana, and P. tecunumanii. At most
locations epiphytes were growing in large numbers on the branches and trunks
of the trees. In Guatemala and El Salvador some of the larger oaks with massive,
horizontal branches were almost covered with orchids and bromeliads. Un-
fortunately, in most locations—particularly in Guatemala and El Salvador—
the forests were rapidly disappearing as the trees were cut for lumber and
firewood and the land was converted to pasture and crops.
SPECIMENS EXAMINED.* Mexico. VERACRUZ: ca. 15 km W of Jalapa, 1800 m alt., Perry
M96-81, M96-81A. Cutapas: ca. 18 km § of San Crist6bal de Las Casas, 2200 m alt.,
Perry MEX.24-79, 15 km N of Comitan, 2200 m alt., Perry MEX.25-79; 20 km S of
San Cristobal de Las Casas, 2200 m alt., Perry MEX.26-79; E of San Cristobal de Las
Casas, vic. of Las Piedrecitas, 2400 m alt., Perry MENX.151-83; ca. 10 km W of San
Cristobal de Las Casas, near Hwy. 190, 2300 malt., Perry MEX.74-83; S of San Cristobal
de Casas, vic. of Teopisca, 2300 m alt., Perry MEX.84-84. Guatemala.
QUEZALTENANGO: Vic. of Quezaltenango, 2300 m alt., Perry GUA.3-78. SOLOLA: ca. km
140 W of Guatemala City, 2400 m alt., Perry GUA.17-78, GUA.19-78; W of Quezal-
tenango on hwy. toward San Marcos, ca. km 232, 2300 m alt., Perry GUA.24-78. JALAPA:
on dirt road from San José Pinula to Mataquescuintla, 2300 m alt., Perry GUA.112-78
(Ncsc), GUA.112-78A; E of San José Pinula on dirt road, vic. of Las Nubes, 2200 m
alt., Perry GUA.113-78; on dirt road from San José Pinula to Las Nubes, ca. km 38,
2250 m alt., Perry GUA.28-79; E of San José Pinula on dirt road near Soledad Grande,
ca. 2200 m alt., Mittak 8299 (BANSEFOR‘):; E of San José Pinula on dirt road, vic. of La
‘Specimens listed are in addition to those collected as vouchers for the trees tapped for oleoresin.
Unless indicated otherwise, they are located in the author’s personal herbarium.
‘Banco Nacional de Semillas Forestales, Guatemala.
1987] PERRY, PINUS 451
Cc D
iGURE 4. Cone scales: A, Pinus nubicola, B, P. estevezii; C, P. oaxacana; D, P.
pseudostrobus.
Lagunilla, ca. 2100 m alt., Mittak 9017 (BANSEFOR). El Salvador. CHALATENANGO: near
Miramundo, 2200 m alt., Perry SAL.7-77, near El Aguacatal, 2000 m alt., Perry SAL.8-
77. Honduras. La Paz: vic. of Las Trancas, Perry H-8, H-10 (ESNACIFOR).°
RELATIONSHIPS OF PINUS NUBICOLA
Pinus nubicola, with its slender, pruinose branchlets and its smooth-barked
young trees, readily falls into the Pseudostrobus group of Mexican pines, which
has been variously called a section (Martinez, 1948), a “group” (Loock, 1950;
Stead, 1983a; Stead & Styles, 1984), and a “complex” (Mirov, 1967; Stead,
1983b). The other species in the group are P. douglasiana Martinez, P. maxi-
minoi H. Moore, P. pseudostrobus, P. oaxacana, and P. estevezii (Martinez)
Perry.® As determined through chemotaxonomic studies of many of these taxa
by Mirov (1958, 1961, 1967), Coyne and Critchfield (1974), Brimmer (1978),
Mittak and Perry (1979), and Perry (1982), P. oaxacana and P. estevezii are
most closely related to P. nubicola.
‘Escuela Nacional de Ciencias Forestales, Siguatepeque, Honduras.
Stead and Styles (1984) criticized the use of resin chemistry by Mirov (1958) and Perry (1982) in
elevating var. oaxacana and var. estevezii to specific status.
452 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
TABLE 1. Turpentine composition* of xylem oleoresin from Pinus nubicola.
ce
: ae
QO i) HW cv) Tt > Lol
E a vu i=] I © oO
2 2 yg a9 ¢ Ho &§ oa? 4B
v fs] e o © €& -& ©£ oOo @ A
o cy v9 @ a @ 4 eo GY vA a ¢€ & 4 A
2 8 §€R 2 8 $G 5 2 Baa BSE
& «a uw eG a i=] a. l wow i=} [=5 cs) u HW woo
iss) co} io) ie) i] i} i on tal i} od 1 1 i) vu uv {
POPULATION nF OC 4 8 O 2 4 =. 4h ee Oe = = 3
ME
Chiapas 1B 25 12 28 10 #1 3 18 2 #1
ta 2B 10 12 31 9 3 30 2 1
Wi of San 3B 34 14° 6 o 2 39 1
Cristobal de 4B 34 15 31 6 7 1 oS -d
5B 5 2 44 2 2 84 tr 1
6 6 is s 64 8
7B 13 7 44 5 7 YT 6. Tr 3 1 Tr
eB. 2 12 43 9 3 4 21 #6 Tr
7 #8 rt. 8 16 2 4 6 9
10B 27 il 36 6 2 a 12 4
11B 39 12 34 Tr 3 3 3 o ode An
12B 12 54 16 4 = 2) 3
13B 2. 81 6 Tr 10 Tr
Station San M6083 51 Tr 17 12 1 1 6 bi ag 7 #65
Jos M7283 15 8 36 5 3 24 1 8 1
M7383 8 5 39 1 #4 31 1 7 #4 #‘Tr
Station Las M12483 9 6 30 8 15 26 1 3 1
Piedrecitas M14283 6 12 13 al 40 3.18 4 1
M15283 12 11 Tr 5 K| 53 6 11
M14883 - a £ 64 1 Tr 4 Tr
GUATEMALA
East of 2A 26 4 8 Tr 1 2 2 1 4 4
San Jose 3A 19 12.18 8 co ae > 2 1
P 1 4A 42 8 39 6 2 1
6A 31 Tr 11 16 ae: 2 4 17 6 1 2
TA 40 ie} 1 a 20 Hie a
8A 39 io S 6 1 4 23 2
10A 21 10 38 4 3 8 3 13 Tr
11A 27 13 24 oa 2 6 4 5 10 1
124 32 18 28 4 5 8 4 1
HONDURAS
Las Trancas H8 42 3 30 5 3 3 Tr 4 2 Tr:
H10 7 3 65 3 #5 2 1 8 al
Mean 21 1 #12 27 Tr 5 3 Tr 3 19 Tr Tr 1 5 3 1
*Percent of total turpentine.
Tr - trace.
At a number of locations in Mexico (Chiapas), Guatemala, and El Salvador,
I have observed trees with cones and foliage that appeared to be intermediate
P. pseudostrobus, the trees were five needled, and the bases of the fascicle bracts
were not decurrent. In many instances P. nubicola, P. pseudostrobus, P. mon-
tezumae, P. oaxacana, and occasionally P. rudis formed a part of these mixed
stands. It hae that hybridization and back-crossing had been occurring
for many years among these pines. Mirov (1961) stated that P. oaxacana
apparently crosses naturally with P. pseudostrobus and probably also with some
varieties of P. montezumae. Martinez (1948) pointed out that P. pseudostrobus
and P. montezumae are very closely related. Extensive sampling and analyses
1987] PERRY, PINUS 453
95 90
Map |. Distribution of Pinus nubicola.
of oleoresins from carefully selected trees could, with morphological studies of
the cones and foliage, reveal the extent and nature of the hybridization that is
occurring in many of these mixed stands.
Because the chemical composition of turpentine is inherited (Squillace, 1976),
I believed that information about this character would provide valuable knowl-
edge regarding identification and possible relationships of the new species.
Accordingly, I took samples of xylem oleoresin from Pinus nubicola trees in
Mexico, Guatemala, and Honduras. Results of the analyses are shown in TABLE
1. Information regarding collection and analysis of the oleoresin is given in the
APPENDIX.
Although Pinus nubicola fits Martinez’s and Loock’s original concepts of a
Pseudostrobus group of the Mexican pines, it (like P. oaxacana and P. estevezii)
differs markedly from P. pseudostrobus in the chemical composition of its
turpentine. Heptane and nonane are consistently present in the turpentine of
P. nubicola, P. estevezii, and P. oaxacana but usually absent in P. pseudostrobus
(see TABLE 2). Mirov (1958) stated that the gum turpentine of P. oaxacana
contained 21 percent heptane, 51 percent d- and dl-a-pinene, 15-16 percent
dl-limonene, 1.3 percent n-undecane, and 7.5 percent d-longifolene.’ There
™Mirov’s data were obtained from one sample, in which oleoresin from 25 trees (from near Rancho
Nuevo, 25 km SW of San Cristobal de Las Casas, Chiapas, Mexico) was combined. In other samples
the percentages may be different. The presence of large quantities of heptane is significant
454
JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
TaBLe 2. Turpentine composition of xylem oleoresin from Pinus nubicola, P. estevezii,
P. oaxacana, and P. pseudostrobus.
P. nubicola P. estevezii P, oaxacana* P. pseudostrobus
(n 3 n 13) (n = 26) (n = 10)
TERPENE Mean’ % Hight Mean % High Mean % High Mean % High
Heptane 21 61 38 100 16 54
Octane 1 Z 4 15
Nonane 12 61 11: 55 10 35
a -pinene ar 65 19 61 37 Haff 80 100
Camphene th 3 } 12 1
8 -pinene 5 29 4 4 15 2
\*-carene 3 16 Hipg 4 4 4
Myrcene Vie ag 9 30 1 40
® -terpinene 3 6 Tr
Limonene 19 48 3 8 10 27 1
8 -phellandrene Tr 4 15 Tr
P-cymene ex Tr
Terpinolene HT Tr 1 4 TRF
& —~fenchol Tr Tr
Terpinene-4-ol 5 16 2 8
8 -caryophyllene Tr
Methyl] chavicol 3 16 6 15 4 12
a -terpineol i 3 1 4 x
*Samples collected by the author in Mexico (Puebla, Oaxaca, and Chiapas states) and
Guatemala.
‘Mean percent of total turpenti
*Percent of trees having rela
et al., 1980).
Squillace
ne
tively high amounts. (For mathematical procedure see
thus appears to be a cluster of species within the Pseudostrobus group of
Mexican pines that differ from typical P. pseudostrobus in the morphology of
their cones and in the presence of heptane and nonane, usually in high amounts,
in their turpentine. Further studies of oleoresins from the remaining taxa in-
cluded in Martinez’s sect. Pseudostrobus are required in order to clarify these
relationships.
DISTINCTION AMONG PINUS NUBICOLA, P. ESTEVEZII,
P. OAXACANA, AND P. PSEUDOSTROBUS
Although the principal identifying characteristics of the Pseudostrobus group
(i.e., the smooth stems of young trees and the nondecurrent bases of the needle
TABLE 3.
Summary of differences among Pinus nubicola, P. estevezii, P. oaxacana, and
P
. pseudostrobus.
CHARACTER
TAXON
P. nubicola
P. estevezii
P. oaxacana
P. pseudostrobus
FORM OF MATURE
LEAVES
Number per
fascicle
Habit
Dimensions
Anatomy
CONES
25-30 m all
stem clear;
open, rounded
5 6
(occasionally 7,
rare 8)
Flexible, very
drooping to
pendent
25-40(-43) cm x
6-1 mm
Hypode
ec with
many shal w
penetrations into
10-15 x 8-10 cm,
asymmetric,
reflexed
12-20 m tall:
broadly onned
5 (rarely 4)
Stiff. erect
not flexible
20-30(-35) cm x
1 mm
Hypoderm
irregular, with
many shallow
penetrations
int
chlorenchyma:
nals 3
internal
10-13 x 7-8 cm,
asymmetric,
usually reflexed
25-30 m tall;
stem clear;
crown moderately
dense, rounded
5 (rarely 6)
Flexible.
slightly
drooping
20-30(-33) cm x
ca mm
ypoderm
uniform, with
few slight
penetrations
into
chiorenchyma;
resin canals
(to 4). medial
a 11 cm.
asymmetr
usually Po eee
30-40 m tall;
Bs
rown narrow,
rounde
Flexibie,
drooping
20-25(-30) cm x
ca
Hypoder
uniform, with
few slight
penetrations
into
chlorenchyma
S=10° 57
nee curved.
not refl
[L861
SONId ‘AWUdd
SSP
TABLE 3.
(continued).
TAXON
P. nubicola
P. estevezii
20-25 mm wide,
prickle; margin
of apex wi
unequ
projections
Usually large
mount of heptane
and sma r
amoun of nonane
sometimes large
amount o
terpinene- Ls
very small amount
d
chavicol
1800-2400
P. oaxacana
P. pseudostrobus
12-15 mm wide,
hard, strong,
thick; apophyses
transversely
keeled: umbo
ise th
Usually large
f
®2-phellandrene
800-1800
12-20 mm wide,
r ng.
thick; apophyses
with pronounced,
22 mm long);
margin
of apex smooth
Usually large
and sm
amoun of
nonane; usually
r
methyl chavicol
1500-3200
15-18 mm wide;
apophyses
slightly raised
(e) is
sli ly
transversely
Ke d; um
small
occasiona
depressed, wit
smal] deciduous
prickle; margin
fe) ape smooth
Heptane, octane,
usually
large amount of
-pi ne,
occasionally
with arge
amoun of
myrcene; usually
smal] amounts of
a-terpineol
1600-3200
WOLAYOUAUV GIONAV AHL AO TYNUNOSL 9ST
89 10a]
1987] PERRY, PINUS 457
bracts) are shared by Pinus nubicola, P. estevezii, P. oaxacana, and P. pseu-
dostrobus, the four species can be readily separated by combinations of char-
acters (see TABLE 3)
Pinus nubicola is easily distinguished in the field from the other three species
by its long, very drooping needles (see FiGureE 2) five or six (occasionally seven)
in a fascicle, and its large, ovoid to long-ovoid cones with unusually wide, thick
cone scales having unequal apical projections and a small depressed umbo
(FicuRE 4). Cones of P. oaxacana are readily identified by their thick, stiff
cone scales with unusual prolongation of the apophysis and umbo. Pinus es-
tevezii can be distinguished from the other three taxa by its stiff, erect needles
and its cones with thick, hard scales armed with a persistent up-curved prickle
on the umbo. Pinus pseudostrobus is easily separated from the other three
species by its much smaller cones having thin, flexible scales with a flat to
slightly raised apophysis and a small umbo tipped by a small deciduous prickle
(FIGURE 4).
A comparison of the oleoresin components z also reveals aan differences
among the four species (see TABLE t g these was the presence
of high amounts of limonene in 48 percenit of Pinus nubicola trees. In addition
there were trees of P. nubicola with high amounts of terpinene-4-ol (16%) and
methyl chavicol (16%).
There appeared to be some population differences, but samples were too few
for this to be determined with certainty. For example, all of the Pinus nubicola
trees in the Guatemalan population had high heptane levels while only about
half of the trees in the Mexican population did (see TABLE 1). It is interesting
to note that the two trees in Honduras had the highest nonane levels of all the
trees sampled.
As in most species, individual trees varied greatly in monoterpene compo-
sition. It would have been helpful to have oleoresin samples from Pinus nu-
bicola trees growing in El Salvador. Unfortunately, the very unsettled political
situation in that country, particularly in Depto. Chalatenango, made it unwise
to attempt any resin collections there.
ACKNOWLEDGMENTS
I would like to express my appreciation for the help received from a number
of friends and colleagues during this study. Suggestions and comments on the
manuscript from P. C. Mangelsdorf, B. Zobel, J. W. Duffield, W. B. Critchfield,
and P. F. Stevens were invaluable, as were P. F. Stevens’s assistance with the
Latin diagnosis and E. B. Schmidt’s editing of the manuscript. Suggestions from
A. E. Squillace and E. C. Franklin regarding the turpentine analyses and the
presentation of the data were very useful. I am also grateful for J. Drew’s advice
and comments on analyses of the oleoresins. My thanks are expressed here for
V. G. Wright’s drawings, and for W. L. Mittak’s help with the collection of
oleoresins and herbarium specimens in Guatemala. Similar collections in Mex-
ico and Honduras were made possible by the assistance of W. S. Dvorak,
Director of CAMCORE, and staff members J. K. Donahue and E. G. Ponce;
my deep appreciation for their help is expressed here.
458 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
This study was supported in part by a grant from the Rockefeller Foundation
made through North Carolina State University to J. P. Perry, Jr. A grant from
the American Philosophical Society assisted with payment of resin-analysis
costs.
LITERATURE CITED
Brimne_r, U. 1978. Zur re ail von Kiefernharzen. Unpubl. M.A. thesis,
ones Hamburg, Ham
Coyne, J. F., & W. B. oa +1974, _ Identity and terpene composition of Hon-
duran sae attacked by the bark beetle idae). Turrialba
14: 327-331.
Loock, E. E. M. 1950. The pines of Mexico and British Honduras. S. Africa Dept.
Forestry Bull. 35: 1-244.
Martinez, M. 1948. Los pinos mexicanos. ed. 2. Ediciones Botas, Mexico.
Mirov, N. T. 1958. Pinus oaxacana, a new species from Mexico. em 14: 145-
150.
1961. Composition of gum turpentines of pines. U.S.D.A. Forest Serv. Tech.
Bull, 1239: 1-158.
967. The genus Pinus. Ronald Press Co., New
Mitrak, W. L., & J. P. Perry, Jr. 1979. Pinus maximinoi: its taxonomic status and
distribution. J. Arnold Arbor. 60: 386-395.
Perry, J. P. 1982. The taxonomy and chemistry of Pinus estevezii. J. Arnold Arbor.
63: 187-198.
SQUILLACE, A. E. 1976. Analyses of monoterpenes ene by gas- -liquid chromatog-
raphy. Pp. 120-157 in J. P. MIKSCHE, ed., ern methods in forest genetics.
Springer- pels Berlin and Heidelberg.
,O. is, & D. L. Rockwoop. 1980. Inheritance of monoterpene com-
aa in ere oleoresin of loblolly pine. Silvae Genet. 29: 141-152.
STEAD, J. 1983a. Studies of variation in Central American pines V: a numerical
study variation in the Pseudostrobus group. Silvae Genet. 32: 101-115
983b. A study of variation and taxonomy of the Pinus pseudostrobus complex
Commonw. Forest. Rev. 62: 25-35.
& B. T. Sryces. 1984. Studies of Central American pines: a revision of the
*pseudostrobus’ group (Pinaceae). J. Linn. Soc., Bot. 89: 249-275.
Styes, B. T. 1976. Studies of variation in Central American pines I. The identity of
Pinus oocarpa var. ochoterenai Martinez. Silvae Genet. 25: 109-118.
APPENDIX. Collection and analysis of oleoresins.
COLLECTION
In Guatemala samples of xylem oleoresin were collected from nine trees of Pinus
nubicola (d.b.h. 30-60 cm) growing near San José Pinula at the location described for
At about 75 cm above the ground, a hole ca. | cm in diameter was drilled into the
stem of each tree (22 February 1979) and a threaded glass vial was immediately screwed
ughtly into the hole. Three days later the vials were collected and each one covered with
a threaded, gasketed cap. Perry GUA.28-79 was rato as a composite voucher for
these trees and has been deposited in the herbaria at GH a ic.
In Chiapas, Mexico, samples of xylem oleoresin were nee from 13 trees of Pinus
nubicola (d.b.h. 35-90 cm) about 10 km west of San Cristobal de Las Casas, near highway
190. Vials were placed on the trees 2 March 1979 and collected two days later. Perry
1987] PERRY, PINUS 459
en 7B-12B and Perry MEX.1B-14B were collected as composite vouchers for these
or of San Crist6bal de Las Casas, near Las Piedrecitas, oleoresin was collected from
four trees of Pinus nubicola (d.b.h. 32-50 cm). Vials were placed on the trees 31 January
and | February 1983 and collected the following day. Perry M-12483, M-14283, and
M-15183 were collected as vouchers for these trees
West of San Crist6bal de Las Casas, near San José, oleoresin was collected from three
trees of Pinus nubicola (d.b.h. 35-80 cm). Vials were placed on the trees 27—28 January
and ee 30 January 1983. Perry M-6083 and M-7283 were collected as vouchers
for these tre
In Honduras oleoresin was collected from two trees of Pinus nubicola in the Depar-
tamento de La Paz, near the village of Las Trancas. Collections were made in October
1982 by W. S. Dvorak, Director of CAMCORE (Central America and Mexico Coniferous
Resources Cooperative), and E. G. Ponce, staff member of ESNACIFOR (Escuela Na-
cional de Ciencias Forestales), Honduras. Ponce H-8 and H-10 were collected and de-
posited in the herbarium at ESNACIFOR, Siguatepeque, Honduras.
The sampling procedure described for collections in Guatemala was followed for all
collections in Mexico and Honduras.
ANALYSIS
Analyses of the pine-resin samples were performed by a chemical consulting laboratory,
with the following gas-chromatographic conditions and equipm
Turpentine from each sample was separated from the resin and extraneous matter by
steam distillation (alkalinity was maintained to prevent acid isomerization
The chromatograph used was a Varian Series 1700 with a thermal conductivity de-
tector. A 10’ x %” diameter stainless-steel column packed with 15% carbowax 20M on
“chromosorb W” was injected with 1.5 ul of sample. The injector temperature was 210°C,
the detector temperature 225°C, and the column oven programmed from 75° to 220°C
with a 4°C per minute temperature rise. The carrier gas was helium.
Components were identified by comparison of elution times against standard chro-
matographs made from combinations of pure compounds. When a question arose as to
the identity of a compound, the sample was reshot with known components added until
the presence of overlapping peaks or increase in peak size eliminated any uncertainty.
CHU, ARCHIATRIPLEX 461
ARCHIATRIPLEX, A NEW CHENOPODIACEOUS
GENUS FROM CHINA
GE-LIN CHU!
A new genus of Chenopodiaceae (4rchiatriplex) and its sole species (A.
nanpinensis, from northern Sichuan Province, China) are described. The genus
is characterized by unisexual flowers, foliaceous bracts subtending the carpel-
late flowers, and annular embryos; it therefore belongs in tribe Atripliceae. Its
Agata and morphology are discussed, and a key to the genera of this
tribe is give
In 1974, as I was finishing the manuscript of the Chenopodiaceae for the
Flora Reipublicae Popularis Sinicae (Kung & Tsien, 1979), my attention was
drawn to an unidentified fragmentary specimen (K. 7. Fu 2/66) in the her-
barium of the Institute of Botany, Academia Sinica, Beijing. Its surprising floral
morphology —unisexual flowers with the staminate ones fasciculate in terminal,
interrupted spikes and the carpellate ones below—suggested that the plant could
be placed in the tribe Atripliceae C. Meyer, but in floral and inflorescence
morphology it matched no genus in the tribe. Although I located another
specimen of the same taxon in the herbarium (7. P. Wang 7967), it was also
fragmentary.
In 1980 I had the opportunity to visit Nanping, on the northern flank of the
Tsinling mountain range in Sichuan Province (Map 1), where both of the
specimens had been collected. While there, I was fortunate to re-locate the
population and was able to re-collect more complete specimens and make field
observations. Study of the ample material gathered at that time has shown that
the plant is a new species that also comprises a new genus. I propose the new
enus and species below, followed by a discussion of its relationships and
morphology
Archiatriplex G. L. Chu, gen. nov.
Proximum Microgynoecio J. D. Hooker sed in floribus femineis basi et in
stipitibus bractearum insertis, perianthio evoluto, et staminibus differtibus,
dissimilis.
Monoecious herbs. Leaves opposite or alternate, petiolate, complanate, slightly
succulent, serrate, with unicellular inflated trichomes. Flowers unisexual. Sta-
minate flowers in interrupted spikes at apexes of branchlets, lacking bracts;
perianth 5-parted, segments membranaceous, slightly succulent on back near
apex, lacking nerves; stamens 5, inserted on disc. Carpellate flowers under
‘Institute of Botany, Northwest Teachers’ College, Lanzhou, Gansu, People’s Republic of China.
© President and Fellows of Harvard College, 1987
Journal of the Arnold Arboretum 68: 461-469. October: 1987.
462 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
0 104 10:
103 ooo
i)
Zhouqu 7
e
ie! GANSU <S
N ad
y )
i Bailong\River go
f ¢
Nanpin, i (
'p 1g \ :
* c
33 Longkang eee ae
e
Song
SICHUAN
Jialing iver
\
Map 1. Archiatriplex area, showing major river-system involving tributaries of Huanghe
angjiang rivers, and 4 towns on border of Sichuan and Gansu provinces, with *
indicating type locality of A. nanpinensis.
32
staminate inflorescences, attached to base and petiole of bracts; bracts folia-
ceous, short-petiolate or nearly sessile, smaller than leaves; perianth 3- or
4-parted, the segments with longitudinal midrib, slightly enlarged in fruit; ovary
obovoid, smooth, with 2 stigmas, style inconspicuous. Utricle slightly com-
pressed, papillate; pericarp membranaceous, adnate to seed. Seeds laterally
compressed, lenticular, testa crustaceous; embryo annular, perisperm copious.
Type SPECIES: Archiatriplex nanpinensis G. L. Chu.
Archiatriplex nanpinensis G. L. Chu, sp. nov. FIGuRE |.
Herbae annuales, usque ad 1.2 m altae; caulis erectus vel ascendens, ramosus,
leviter tetragonus, striatus; rami ascendens ramosi, ramulis 1-5 cm longis, saepe
gracilibus. Folia late ovata vel triangulari-hastata, 2-10 cm longa, latitudine
longitudinem fere aequante, supra viridia, subtus pallide viridia, apice breviter
acuminata, basi cordata, margine irregulariter laxe dentata; petiolus tenuis,
0.5-8 cm longus. Inflorescentiae masculinae graciles, interdum ramis brevibus
praeditae; flores masculini multi in glomerulis dispositi; segmenta perianthii
obovata vel oblanceolata, circa 1 mm longa, basi tantum connata, prope apicem
leviter succulenta, apice paulo cucullata; stamina 5, filamentis filiformibus,
1987] CHU, ARCHIATRIPLEX 463
e, seed,
lateral view, x 7.5, showing position of radicle and hilum; f, seed, longitudinal section,
x 7.5, showing testa, curved embryo with radicle, 2 cotyledons, and central endosperm.
(a-c drawn by Xia Quan.)
464 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
planis, segmentis perianthii fere aequilongis, antheris late oblongis vel late ova-
tis, circa 0.3 mm longis. Flores feminei 4—7 simul in glomerulo, basim bracteae
inserti; bracteae ovatae vel cordatae, 4-20 mm longae, margine integrae vel
serratae; eines perianthii basi fructificationis lineari-elliptica vel obovata,
./-1 mm longa, basi tantum connata, patentia, margine integra vel leviter
lacerata; stigmata circa 0.2 mm longa. Utriculus oblique ovatus, tuberculatus.
Semen rubiginosum vel nigrum, nitidum, circa 1-1.5 mm
Annual herbs to 1.2 m tall; stems erect or ascending, ramified, slightly te-
tragonal, striate, the branches ascending, ramified, with the branchlets 1-5 cm
long, usually gracile. Leaves with petiole 0.5—8 cm long; blade broad-ovate or
triangular-hastate, 2-10 cm long and nearly as wide, short-acuminate at apex,
cordate at base, irregularly coarsely dentate at margin, dark green above, light
green below. Staminate inflorescences slender, sometimes short-branched:
flowers several in glomerules; perianth segments obovate or oblanceolate, ca.
1 mm long, connate at base, slightly succulent and somewhat cucullate near
apex; stamens 5, the filament filiform, complanate, nearly as long as perianth
segments, the anther broad-oblong or broad-ovate, ca. 0.3 mm long. Carpellate
flowers 4 to 7 per glomerule, inserted at base and petiole of bracts; bracts ovate
or cordate, 4-20 mm long, entire or serrate; perianth segments in fruit linear-
elliptic or obovate, 0.7—1 mm long, connate at base, patent, entire or slightly
lacerate; stigmas ca. 0.2 mm long. Utricles oblique-ovate, the pericarp mem-
branaceous, papillate. Seeds red-brown or black, ca. 1-1.5 mm in diameter.
Type. People’s Republic of China, Sichuan Province, Nanping, Longkang, 2100
m alt., at edge of bush-wood, 30 September 1980, G. L. Chu 80040 (holo-
type, herbarium of the Institute of Botany, NW Teachers’ College, Gansu;
isotype, A).
ADDITIONAL SPECIMENS EXAMINED. People’s Republic of China. SICHUAN PROVINCE
Nanping, Longkang, 2100 m alt., K. 7. Fu 2166 (pe), T. P. Wang 7967 (pe); on banks
of terraced farm, G. L. Chu 80041, 80073, 80086 (all at Herb. NW Teachers’ College,
Gansu)
MORPHOLOGICAL OBSERVATIONS
SEEDLINGS. Approximately 25 seeds were taken from unfumigated isotypes and
were sown on 28 May 1982. Germination was first observed on 2 June and
proved to be epigeal. On the eighth day after germination, the first pair of
photosynthetic leaves appeared; the cotyledons were then ovate-elliptic,
4-6 x 1-1.5 mm, and light green above and purplish beneath. At the first eight
nodes the photosynthetic leaves were opposite, but at the ninth node only one
emerged, and thenceforth the leaves were alternate.
Potten. Pollen of Archiatriplex nanpinensis was taken from fresh material and
prepared for examination with a scanning electron microscope. The tuberculate,
punctate ektexine of the spherical, polyporate grains corresponds to the general
pattern of chenopodiaceous pollen. The grains are ca. 26 um in diameter and
have approximately 60 circular apert scattered on the tuberculate and finely
punctate surface (FiGURE 2a). Each aperture is ca. 2 um in diameter, with six
1987] CHU, ARCHIATRIPLEX 465
Figure 2. Scanning electron micrographs of pollen grains: a, b, Archiatriplex nan-
pinensis, showing numerous circular apertures, and tuberculate and punctate ektexine;
c, d, Microgynoecium tibeticum, showing more numerous and larger apertures and smooth
ektexine (a, c, x 1240; b, d, x 6200)
to nine free or coalescent tubercles (FIGURE 2b). Compared with the pollen of
Microgynoecium tibeticum Hooker f. (FIGURE 2c, d), the grains of A. nanpinensis
have fewer apertures but more tubercles.
CYTOLOGICAL OBSERVATIONS
Very young buds of staminate flowers from greenhouse-grown plants of
Archiatriplex nanpinensis were fixed in Carnoy’s solution, and pollen mother
cells were stained and prepared in the normal manner for microscopic obser-
vation. It was determined that the species is a diploid with 2n = 18. At meiosis
bivalent pairing is regular (see FIGURE 3).
GENERIC RELATIONSHIPS OF ARCHIATRIPLEX
Including Archiatriplex, the tribe Atripliceae consists of 13 genera, of which
Atriplex L. is the largest, with more than 100 species widely distributed in Asia,
North America, Europe, Australia, and Africa. 4xyris L. and Ceratoides (Tourn.)
Gagnebin are represented in the floras of Eurasia and North America, while
466 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Ficure 3. Chromosomes of dividing microsporocyte of Archiatriplex nanpinensis,
n=9, nee 1 (voucher specimen, G. L. Chu 80084).
Spinacia L. and Ceratocarpus L. are confined to Eurasia. The remaining genera
ares Ti
chiatriplex), North America (Endolepis Torrey, Suckleya A. Gray, Zuckia
dley, and Grayi ker & Arn.), Africa (E-xomis Fenzl ex Mog.), or
pees (Theleophyton (Hooker) Mogq.).
Archiatriplex has close affinities to Microgynoecium. The two species are
characterized by similar foliaceous bracts subtending the carpellate flowers,
with each bract containing several flowers; the carpellate flowers of Microgy-
noecium, however, lack perianths. The other genera in the tribe differ from
Archiatriplex in having a single carpellate flower included between two opposite
and specialized bracts and (except for Exomis and Endolepis) in lacking a
perianth, or in the stellate hairs covering the plant
jab}
=S
PS
a
°
}
Ficure 4. Inflorescences in tribe Atripliceae, showing possible evolutionary changes
that led to present forms. a, hypothetical prototype with numerou an anches, each with
staminate flowers at distal end, carpellate at eae (dark bracts subtending lateral
branches indicate key area of evolutionary change). b, Archiatriplex fasci cles of 1 to 7
—
tended by enlarged bracts: g, Endolepis, carpellate Howe with perianth; h, Atriplex,
carpellate flowers lacking perianth
CHU, ARCHIATRIPLEX 467
1987]
468 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
KEY TO THE GENERA OF THE ATRIPLICEAE
—_
. Plant glabrous, or covered with eae or ramified inflated hairs.
2. Carpellate flowers with perian
3. Carpellate flowers sbtended by single foliaceous bract, axil of each bract
usually with several flowers. ......0.00000 00.0000 ccc eee Archiatriplex.
3. Carpellate flowers each abe by 2 opposite, separate bracts, these not
foliaceous.
4. Radicle oriented downward; perianth with 5 segments; bracts cee
Babee aes a nce acai g hea eae Ge eases apes eteie nade rece ORS busta grates Exom
4. Radicle oriented upward; perianth with 3 or 4 segments; _ not suc-
TVET, 5 face ccs te cist eel een SG deat es a BE tae ee Endolepis.
Carpellate dowers lacking perianth.
5. Stigmas 2; plants monoecious, rarely dioecio
6. Gnesi flowers ao by single fouaceis bract, axil of each bract
tN
usually with several NOWEIs: 6.2.2 .65e3s- hove weaeideee ee Microgynoecium.
6. ee ee flowers each ere by 2 opposite bracts, these partially or
totally fus
7. Seeds Sed vertically in fru
. Bracts of carpellate flowers raided saclike; inflated poe se
MEG Ui eis ass cases vara whe eaaiens ophyton.
co
8. Bracts of carpellate flowers compressed; inflated hairs Rees when
dry.
9. cena hairs not ramified; radicle oriented upward, rarely down-
ard.
10. Bracts of carpellate flowers bilobed at apex. .... Suckleva.
10. Bracts of carpellate flowers entire or serrate, not bilobed.
Shatadse dt cate ecagt ah aad sete eon Geese gee dk ped triplex
9. Inflated hairs ramified; fat oriented downward. ... Grayia.
Ts fae oriented yhieaeaeie MMA. 6 hoc bees ee eae tot ne Zuckia.
5. Stigmas 4 or 5; plants aan Soe dis ee eeee aati pap aaa Spinacia.
1. Plant peer with stellate hair
11. Carpellate flowers with Seat perianth, each subtended by 2 separate, foliaceous
GAGIS: cahetda dd enuace eran pine ee ata ok eas oa Seine ge Geo deste AXyYris.
. Carpellate flowers lacking perianth, each subtended by 2 opposite bracts, these
partially or totally fused.
12. Shrubs or subshrubs; bracts of carpellate flowers fused below middle, forming
Ce
l
a"
tube furnished with 4 fascicles of villose hairs. .............. ratoides.
12. Annual herbs; bracts of carpellate flowers fused their entire length, furnished
with acicular appendage on both sides near apex. ......... Ceratocarpus.
Compared with the other genera in tribe Atripliceae, the most distinctive
primitive character of Archiatriplex is its large, loose panicles (FIGURE 4b)
branches and the carpellate ones below (FiGurRE 4a). Evolutionary change from
the Archiatriplex type of inflorescence led to fasciculate carpellate flowers lack-
ing a perianth and to a reduction in the length of the rachis and in the number
of flowers, leaving small bracts as in Microgynoecium (FIGURE 4c). It seems
that also through reduction of the rachis, the fasciculate Endolepis- and Atri-
plex-type inflorescences (FIGURE 4g, h) evolved from the Archiatriplex type.
In the Endolepis type of inflorescence, the flowers have a perianth, while in
the Atriplex type they do not. Another trend in the inflorescence can be traced
1987] CHU, ARCHIATRIPLEX 469
from the prototype: through reduction in flower number and rachis length and
by fusion of the bracts, the Axyris type of inflorescence (FIGuRE 4d) resulted.
Here, two bracts subtend a single carpellate flower with a segmented perianth.
Further evolutionary changes led to the Eurotia and the Ceratocarpus types
(Ficure 4e, f). In these the carpellate flowers lack a perianth, and the subtending
bracts have become highly specialized and fused.
From the above interpretation, it is clear that the discovery of Archiatriplex
provides a better understanding of the evolutionary changes in tribe Atripliceae.
LITERATURE CITED
Kuna, H. W., & C. P. Tsien. 1979. Chenopodiaceae. Fl. Reip. Popul. Sin. 25(2): 1-
194.
HOWARD, COOLEY 47]
SOME BOTANICAL REMINISCENCES OF
GEORGE R. COOLEY, 1896-1986
RICHARD A. HOWARD!
Several years ago I suggested to George Cooley that he divide his life into
chapters and begin dictating his memoirs. He had led such a diverse and
interesting life and had contributed so much to so many people and organi-
zations that only he could supply a complete accounting. I then knew little of
his early years, his short career at Colgate University, his service in World War
I, or his employment in investment banking firms, which led to the start of
his own financially successful company. Why he began collecting botanical
specimens was never clear to me, although it was somewhat illuminated by
his statement years later: “Plants and the people who study them are both
intriguing organisms.”
I met George in 1951, when he came to the Gray Herbarium seeking aid in
identifying specimens of Solidago from New York State and a miscellany of
plants from Florida. I remained associated with him and knew of his botanical
pursuits for 35 years; I saw him last at his favorite spot, his home at Hickory
Hill, Rensselaerville, New York, a month before his death. I would divide the
botanical life of this devoted amateur into several chapters: the flora of Albany
County, New York; the reprinting of Small’s Manual; the flora of Sanibel Island,
Florida; the initiation of the Generic Flora of the Southeastern United States
and his Research Fellow appointment at Harvard; Chinsegut Hill and the
University of South Florida; the AETFAT meeting and our trip around the
world; the flora of St. Vincent, West Indies, and the two-hundredth anniversary
of the establishment of the Botanic Garden; his continuing generosity to botany
and botanists of the United States; and the Cooley prizes of the American
Society of Plant Taxonomists. I offer vignettes of some of these.
SOUTHEASTERN UNITED STATES
George Cooley’s greatest impact came from his interest in the flora of the
southeastern United States. To the best of my knowledge, he visited Florida
first in 1951 and was attracted to Sanibel Island. For many years after that, he
spent several winter months on the island. One of his early collections on
Sanibel was Eragrostis traceyi A. Hitche., which had not been collected since
the original gathering in 1901. He even grew seeds of this plant to have ad-
ditional herbarium specimens for distribution. The available ‘wild flower’
‘Arnold Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138.
© President and Fellows of Harvard College, 1987.
Journal of the Arnold Arboretum 68: 471-478. October, 1987.
472 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
books did not meet his needs, and when Small’s Manual was recommended
to him he learned it was out of print, out of date, and controversial in its
treatment of genera. He was unable to find any botanist or department willing
to undertake an immediate revision, so he financed the reprinting of the manual
(1954) to make it available. After prolonged discussions he was finally con-
vinced that additional collections of both native and cultivated plants of Flor-
ida were needed. To this end he supported the collecting of others and did a
considerable amount himself. The idea of an annotated checklist of the Sanibel
vegetation appealed to him. In 1954 he published a more elaborate study, ““The
Vegetation of Sanibel Island, Lee County, Florida” (Rhodora 57: 269-289),
profusely illustrated and definitive as to the location and abundance of each
species. Perhaps this work was the start of his concern for conservation and
the preservation of endangered species and habitats. The paper clearly shows
Cooley’s broad interests and keen observation. He noted 672 plants of poison
ivy in a quadrat 20 x 20 feet. On one branch four feet long he counted 6130
“blossoms,” not learning until later that it was a staminate inflorescence. To
do this work on the Sanibel vegetation, he perused the library and herbarium
of the Gray Herbarium/Arnold Arboretum, with an appointment as a research
fellow (1954) to use the facilities. George and his wife, Myra, moved to Cam-
bridge for six months and took full advantage of his Harvard appointment,
both intellectually and socially.
The lack of a proper manual for the southeastern United States still bothered
him, but for ten years he gave financial support to the work his Harvard
colleagues felt was basic: a consideration of the generic limits of the flora, the
assembly of a bibliography, and the preparation of new, accurate illustrations
of each genus. This work, under the direction of Dr. Carroll E. Wood, was to
become the Generic Flora of the Southeastern States. The first paper, ““The
Genera of the Woody Ranales in the Southeastern United States,” was pub-
lished in 1958 (J. Arnold Arbor. 39: 296-346) and has been followed by 115
papers by 38 authors. After Cooley’s initial grants, generous support for this
work has been received from the National Science Foundation.
CHINSEGUT HILL AND THE
UNIVERSITY OF SOUTH FLORIDA
When an old friend, Dr. James Allen, was appointed president of the new
University of South Florida, George Cooley “adopted” the school and directed
his support and energy to the establishment of a botany department with a
herbarium, a library, and a botanic garden. The university acquired, presum-
ably with his help, a property known as Chinsegut Hill, near Brooksville, which
Cooley decided could be a center for botanical studies. His first chore was to
supply a new roof for the building, and then he started local collecting. Cooley
invited old friends and acquaintances to Florida and put them to work col-
lecting, mounting, and inserting specimens in the young herbarium. Henry
Gleason, Stanley Pease, Lily Perry, Leonard Brass, Richard Eaton, William
Weston, and Mackenzie Lamb were among the hard-working “volunteers.”
Cooley sought advice on books he bought for the library, and he cajoled curators
of major herbaria to work over collections long in storage to find duplicates to
1987] HOWARD, COOLEY 473
send to South Florida. Among the rare plants he collected at Chinsegut was
the new species Justicia cooleyi Monach. & Leonard. He also gathered the
abundant and pestiferous Dioscorea bulbifera. Cooley and his volunteers filled
a dump truck with the unwanted “‘bulbs” of this species, which he proclaimed
was a suitable entry for the Guinness Book of Records. He admitted defeat,
however, in his efforts to prepare an edible dish from the bulbs—perhaps the
only time when his persistence did not succeed. Before the first undergraduate
class had graduated, the University of South Florida had a biological study
area, a botanical garden, a creditable botanical library, a herbarium, and an
enthusiastic supporter. Cooley was honored at the university’s first convocation
with an honorary Doctor of Science degree.
SOUTH AFRICA AND AROUND THE WORLD
In 1963 the National Botanical Gardens of South Africa invited about 50
botanists to a celebration of their fiftieth anniversary, followed by a month-
long bus tour of the country. For a taxonomist teaching a course in plant
families, it was an opportunity not to be missed, and I planned to go. George
Cooley applied and received an invitation, and we traveled together. Our first
stop was the AETFAT meeting in Florence, Italy. Then we visited Cairo, Addis
Ababa, Nairobi, and Capetown before taking the country-wide tour of South
Africa. Our wives joined us in Johannesburg for the flight to Mauritius, then
to Perth, Melbourne, Canberra, Brisbane, Port Moresby, and Sydney. There
we parted company, the Cooleys visiting New Zealand and my wife and I
continuing to Fiji, Hawaii, and home to Boston. In two months of travel with
a companion, you learn all of his social tricks, foibles, and moods. George was
always cheerful, energetic, and ready to go. At each new social encounter with
a botanist, he offered a dollar bill to anyone who could spell Rensselaerville
and paid only once to an Afrikaans-speaking person in South Africa. His pants
pocket always had a “hole” that surreptitiously dropped shining coins on the
grass, to the delight of children. He treated an assembly of Masai gathered at
a store to soda pop and gave bubble gum to the young police officers in sarongs
at Port Moresby. He joined local botanical societies and natural history clubs,
usually with a life membership, asking that the publications go to the University
of South Florida. In each country visited he gathered herbarium specimens,
depending on the local botanists to identify, press, dry, and ship them to South
Florida. He left money behind, knowing that there was more than enough to
cover costs. Artifacts he purchased in local markets decorated the guest room
in his home, and each had a story, often embellished with the passage of time.
ST. VINCENT
In my work on the vegetation of the Lesser Antilles, George Cooley often
asked how he could help, stating that he and Myra needed a winter escape.
Finally, I suggested that they might like St. Vincent and that I needed plant
specimens from the island and someone to search the archives and botanical-
garden records for data on plant introductions. George and Myra traveled to
St. Vincent in November, 1961, armed with plant presses and a list of taxa
474 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
previously collected by H. H. and G. W. Smith. George was determined to
regather all the species and add to the known flora. As exact localities for the
Smiths’ collections were not in the list published in Kew Bulletin in 1898, the
Cooleys interrupted their stay and took a boat to England to search out the
original specimens in the Kew herbarium for locality data. The information
was not on the sheets, so the search was futile. Undaunted, Cooley returned
to St. Vincent and succeeded beyond my wildest dreams in preparing over
3000 excellent specimens, well documented and with duplicates. He insisted
on working up the material himself on his return to Cambridge, and his col-
lections remain some of the best ever assembled from St. Vincent.
While George was collecting plants, Myra searched the archives for the
historical material I wanted and carefully copied in notebooks records of plant
introductions, trials, and successes. Only a small part of this important detail
has yet been published.
In September, 1963, I wrote to the governor of St. Vincent, calling to his
attention the bicentennial of the founding of the St. Vincent Botanic Garden
two years later. I suggested that the garden might be spruced up, the plants
newly labeled, and the attention of tourists directed to the anniversary. When
the Cooleys returned to the island in 1964, I asked George to determine what
plans were being made for the anniversary. He learned that the idea had local
appeal, but little was being done about it. He approached the colonial governor
(whom he knew personally), the Department of Agriculture, area naturalists,
the press, and local businessmen. Soon committees were actively planning,
assured of some financial support from George Cooley.
The celebration was held in March, 1965, and the Cooleys were present. The
theme was the arrival of Captain Bligh on the Providence with the breadfruit
and other plants. ‘““Captain Bligh’? came ashore in a small boat with some
breadfruit plants in pots. These were “headed” by local volunteers in the same
manner as they had been 200 years previously. Some were planted with cer-
emony. There were special greetings sent from the director of the Royal Botanic
Gardens, Kew, and from the president of the International Association of
Botanic Gardens and Arboreta. A set of postage stamps commemorated the
founding of the garden, the arrival of the breadfruit, and Captain Bligh and
the Providence. A parade featured floats, bands, and marchers in the prize-
winning costumes of the recent carnival. Picnics and flower shows were held
in the botanic garden, which was also lighted for nighttime visitation. Historical
booklets had been reprinted and were distributed. Two cruise boats were in
the harbor for the occasion. In all this George enjoyed his supportive role.
In 1950 I had climbed the Soufriére of St. Vincent, collected specimens,
taken many photographs, and made observations on the growth of the vege-
tation since the last eruption in 1902 had decimated the plants on the eastern
slope. On the talus of the Soma, a fragment of an earlier volcanic mass north
of the present crater, I had collected an herb thought to be a member of the
Gesneriaceae. Conrad Morton examined the material and was not certain of
the family assignment, and E. C. Leonard could not accept it as a member of
the Acanthaceae; both held decision or description pending the collection of
more material. I had asked local naturalists to return to the site for more
1987] HOWARD, COOLEY 475
specimens, but none could find the population. There was also a problem with
another species from the area. Solanum urens Dunal was described in 1852
from material collected 50 years earlier by Alexander Anderson, the second
director of the St. Vincent Botanic Garden, as ‘“‘Bonhomme de Saint Vincent.”
In 1909 O. Schulz, unaware of Dunal’s species, described Solanum lobulatum
on sterile material collected by H. H. & G. W. Smith at Morne Garu, a name
inaccurately associated with the eruptive massif including the Soufriére.
A trip was planned to St. Vincent in early 1971 to record the summit vege-
tation 70 years after the volcano had erupted and to look for the two problematic
species. George Cooley, then 75 years of age, wanted to go along; he would set
his own pace, I was told. The two episodes that occurred on nearly successive
days revealed his extraordinary courage and poise
To search for the Solanum, we decided to ascend Richmond Peak, starting
from the Richmond River valley. Our local companion, Con de Freitas, de-
termined that the only approach was a stiff climb, essentially up a rock face,
to a shoulder where we could climb a ridge to Richmond Peak. There was no
trail, and the face of the cliff required climbing in a crevasse, which was ex-
hausting. From the shoulder the going was easier in some spots and required
machete clearing in others, but the collecting was good and the stops frequent.
The final few hundred yards were a dense tangle of Clusia and intertwined
shrubs, which meant we left packs, field presses, lunches, and water behind.
George trailed the party for a while, but after we found the plant we sought in
flower and fruit and stopped to prepare specimens and take photographs, he
announced he was turning back and would wait for us lower down. Over an
hour later we started down, arms loaded with specimens to be put in the field
presses that were with our packs, water, and lunches. We passed another hour
arranging our presses before we continued down the mountain, wondering
where we would overtake George. At the base of the ridge and near the cliff
face, we made a shift of several hundred yards to the crevasse we had ascended,
still without encountering him. We yelled, to no avail, and assumed he had
found his way down the cliff in spite of our suggestion that he wait for us to
descend together. We returned to our car at dusk and found no sign that George
had been there. Again we called and waited. We knew ofa small store several
miles down the road and drove there to ask if George had stopped for a cold
drink. No one had seen him, so we returned to our starting point. In the path
of our headlights we saw a very tired, dirty George Cooley walking toward us
along the road. He said little beyond the fact that he had not found the crevasse
and so came down the cliff face anyway. How, we will never know.
Two days later we planned to climb the Soufriére from the east, search for
the unidentifiable plant, circle the crater to take photographs of the vegetation
on close radii, and descend the west slope. We left Kingstown before sunrise
to drive to Orange Hill and as far as possible up the mountain. The climb was
fairly easy up to the area of cinders. We reached the rim and walked clockwise
to the point of eventual descent, where we left our packs. We reversed direction,
photographing as we walked to the northern point of the crater rim, and then
traveled north across the dry crater to the talus slopes of the Soma wall. On
the third or fourth talus slope we explored, we rediscovered the unknown plant
476 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
that was to be described later as Lindernia brucei Howard (Scrophulariaceae).
It was then after lunchtime, and we knew several additional hours would be
required to complete the circuit of the rim. George Cooley decided not to
accompany us on the circuit but to return to our packs and wait for us there.
Again we admonished him to be careful, and we started off in different direc-
tions. Clouds had already formed and periodically obscured the visibility, so
we often had to wait to take our photographs. About 5 p.m., later than we
expected, we returned to our packs. There was no sign of George’s having been
there. We had settled down to eat something and wait when we heard a faint
yell of what we thought was “hello” across the crater. We replied and realized
the answering shout was “help!” By then some of us had stiffened up from the
walk and the wait. Con de Freitas started off at a run, while the rest of us
followed as fast as we could. Con outdistanced us but from the clouds soon
called “bring the rope,” for we had carried a stout climbing rope with us. One
local aide went back for the rope, and the rest of us continued until we found
Con de Freitas flat on his stomach at the fragile edge of the crater.
George Cooley, returning alone, had become lost in the clouds, put down
the package he was carrying, and approached the rim to determine where he
was in relation to the crater. The edge gave way, and he fell down the crater
wall. Somehow he managed to turn on his stomach in the fall and grasp for
something to stop his slide. Several bromeliads growing in the cinders served
the purpose. It was truly a miracle, for the steep slope became vertical a few
yards below, with a straight drop of several hundred feet to the crater lake.
While holding onto the Guzmania plants, Cooley was able to scrape a toehold
in the cinders of the slope and eventually made it wide and deep enough to
support his weight. As he could not climb upward, he widened the shelf he
had created and eventually released his hold on the bromeliads and perched
on his small ledge. So he was found, about 30 feet below the crater rim. Only
the bag he had left above indicated where he was. He reported afterward that
he was so tired from his exertions he had trouble staying awake, but he didn’t
dare fall asleep, so he recited all the poetry he recalled, prayed, and sang the
hymns of his childhood. His periodic calls were eventually heard. We estimated
that he must have been there three hours before we located him.
The retrieval process was not an easy one. Although one end of the rope was
tossed over repeatedly, the strong winds up the crater wall invariably placed
it outside of George’s reach. Finally it was decided to tie the rope in a bowline
and lower the lightest member of our party over the crater rim. An unnamed
St. Vincentian accepted the role. I, being the tallest and heaviest of the party,
anchored the rope around my waist and spread-eagled on the cinder slope
outside of the crater. The others lowered the young man, who reached Cooley
and joined him on the shelf he had created. The rope was tied around Cooley
under his arms, and he was hauled back to safety. Fortunately, the next toss
of the rope reached the volunteer, who was also pulled up. Our transport was
to meet us at the coast on the leeward side of St. Vincent, so we had no choice
but to walk back to our packs and descend the Soufriére. Our one flashlight
gave out at this point, and we were forced to proceed in total darkness. We
had Cooley on the trail, his left arm over my shoulders and his right over Dick
1987] HOWARD, COOLEY 477
Weaver’s, as we worked our way around the crater rim and then down the
several miles to the coast. Once there, we still had two miles of beach ahead
of us and a pair of rivers to ford. Cooley was completely exhausted by the time
we reached the contact point, but fortunately our driver had waited. We re-
turned to our hotel at midnight, grateful for our beds.
A knock on my door early the next morning awakened me. There was George
Cooley in sparkling white shirt and shorts, white socks, and clean sneakers. He
looked ready for a tennis match and was his usual joking self for a few minutes
but then admitted he was having severe chest pains. Hastily we located a doctor
and rushed George to her office, fearing the worst. The pains proved to be
from a chest bruise derived from the bowline knot and the drag on his chest
as we pulled him from the crater wall of the Soufriere. Characteristically, George
had packed his bags before awakening me and wanted to return to New York
immediately. We put him on a plane for Barbados in less than two hours.
When we returned to Boston a week later and called him, he said he was fine
but added, “Don’t tell Myra.” To my knowledge, she has never known this
story of true courage, the result of foolish behavior on his part and negligence
on mine for letting him start back alone. Anyone who knew George Cooley
will understand that he wasn’t to be dissuaded when he had made up his mind.
The story of “the man who fell into the crater and survived” persists on St.
Vincent in several versions. One of these was written in Boy’s Life magazine
(August, 1974) as a legend of St. Vincent.
In the winter of 1971-1972, extruded cinders from the bottom of the crater
lake formed a cone in the crater, and the rising hot water destroyed vegetation
at the lake edge but not that of the crater rim. A massive eruption of the crater
in 1979, however, did destroy all plants on the upper levels of the volcano,
and our photographic record is useful now only to document the regrowth of
vegetation in the 1902-1971 period.
CONTINUING GENEROSITY
The adventures of 1971 may have been George Cooley’s last field trip. He
returned to Sanibel Island many times. There he helped to create the Sanibel-
Captiva Conservation Foundation and to establish a nature preserve, cutting
brush and establishing trails in his energetic way. He was the “local” guide for
many of the visitors. His interest and support turned to the Nature Conser-
vancy, where he served on the national board of governors and was awarded
the Conservancy’s Oak Leaf Award in 1984. He spearheaded the protection
of Florida’s Tiger Creek near the Bok Tower and helped establish reserves 1n
several areas, including the Big and Little Bear swamps in New York State.
He was generous to Colgate University (which he attended for only six months),
where a library and herbarium, as well as a chair in Peace studies, are named
for him. To encourage botanical studies in the southeastern states, he supported
local floras, herbarium development, and lecture series, many named for him.
Nationally, he funded the Cooley prizes of the American Association of Plant
Taxonomists. One was for the outstanding paper published during the previous
year, but this was not continued beyond the initial five-year period for lack of
478 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
suitable papers. The other was for the best paper presented at an annual meeting
of the American Society of Plant Taxonomists. Since its inception in 1956,
this prize has been awarded to over forty individuals, who are eligible to win
but once. After Cooley’s death, when it seemed the prize would lapse, his wife
and daughters added to the available funds to permit the award to be endowed
and the prize continued. Former award winners have added to this fund with
gifts in his memory. In addition, Justicia cooleyi Monach. & Leonard, Thal-
ictrum cooleyi Ahles, and Thelypteris cooleyi Proctor honor his role in plant
taxonomy
George Ralph Cooley was born May 29, 1896, in Troy, New York, and died
September 27, 1986, at his home in Rensselaerville, New York. He was a
valued, sincere, and thoughtful friend to many of us and a devoted, loving
husband, father, and grandfather.
1987]
INDEX
479
INDEX
Abdra, 211
— brachycarpa, 212
Abelmoschus esculentus, 125
Abies, 289
_— Ere ee var. tacanensis, 450
Abietina
Sree 363, 370, 391, 393-395
— mexicana, 395
— RG 395
— ovata, 395
Asu-Asab, Monss S., and Puitip D. CAN-
TINO. Phylogenetic Implications of Leaf
Anatomy in Subtribe Melittidinae (La-
biatae) and Related Taxa, 1-34
Abutilon indicum, 125
— umbellatum, 125
Acacia, 316
— acantholoba, 309, 3
— adenantheroides, a 311
— anegadensis,
— arabica, 349, 350, 352, 353
— auriculiformis, 349, 350, 352, 353
09
— leucophloea, 349, 350, 352, 353
— macracantha, 109, 123
— nilotica, 123
Acalypha amentacea subsp. wilkesiana, | 22
, 352
Achyranthes aspera, 118
— obtusifolia, 118
Acmopyle, 285
Acorellus, 396
Acrocephalus capitatus, 10
Actinostrobus, 284, 287, 292, 293
Acuneanthus, 135
Adenanthera pavonina, 349, 350, 352, 353
Adina, 170
Aduseton, 203, 204
Aesculus flava ‘Solander Ce eerie
ae), Status of the Name, 335-34
— glabra, 337, 339
— hippocastanum, 337
ene lutea, 340
a, 337
— ae 340
— octandra, 335, 339, 340
— pavia, 337, 33
— sylvatica, 339, 340
— karat
1,2
_ ens 12. 14902219332
Albizia amara, 349, 350, 352, 353
— lebbeck, 123, 349, 350, 352, 353
Alocasia plumbea, 116
he Genera of
Alysseae (Cruciferae; See, in the
Southeastern United States, 185-240
AL-SHEHBAZ, IHSAN A., and VERNON
Bates. Armoracia lacustris (Brassicace-
ae), the Correct Name for the North
9
Alternanthera brasiliana, 118
— caracasana, 118
— repens,
Althenia, 260, 261
Alysseae (Cruciferae; Brassicaceae) in the
Southeastern United States, the Genera
of, 185-240
Alyssoides, 197, 216, 227, 228
Alyssum, 186-188, 190, 195- 205, 208, 227,
228
— subg. Tetratrichia, 196
— sect. Alyssum, 196, 199
— sect. Gamosepalum, 197, 199
— sect. le 197, 198
— aizoides
— ene 186, 196, 198
— — var. alyssoides, 196
480 JOURNAL OF THE ARNOLD ARBORETUM
Alyssum alyssoides var. depressum, 196
— americ 196-198
yf
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So
<<
3 oe
oOo 3
Se
8s
——
Nomte)
oon
|
[ei
oO
wn”
OQ
a
< Oo
=e
Be.
BEN
—
So
~)
— gracile, 223
|
=
=
n
i=
=
=
55
o
co
|
°
2%
ie}
<
p
2
=
5
pk
oO
ox
I
\o
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_ Fier 200
— shortii, 224
— cue 198
— spinosum, 197-199
— strigosum, 197
— szowitsianum, 197, 199
Ambrosia hispida, 121
Amentotaxus, 288
Amomum,
Amyris elemifera, 129
Anacardiaceae, 118
Anacardium occidentale, 118
da, 336
Anethum graveolens, 131
Angiospermae, 115
Anguilla and Adjacent Islets, Contribu-
31
tions to a Flora of, 105-1
Anisomeles, 2—4, 18, 23
— iar 10
— ov
ree glabra, 177
— muri si 119
squa , 119
Seer 119
Anotis, 150
Antigonon leptopus, 126
Antirhea acutata, 109, 127
Aploleia monandra, 116
Apocynaceae, 119
119
ae 275, 280, 286
— heterophylla, 115
Araucariaceae, 115, 269-271,
282-284, 286, 288, 293, 294
Araucarites, 293
Archiatriplex, a New Chenopodiaceous
69
s from China, 461-4
Archiatriple, 461-469
26
Argusia enaphalodes, 108, 119
Argythamnia candicans, 122
Aristida adscensionis, 116
Armoracia lacustris eee, the
Correct Name for the North American
9
Asparagus setaceus, 116
— sprengeri, 116
Asperuginoides, 188
Asystasia gangetica, 118
Athrotaxites, 293
Atriplex, 465, 468
— pentandra ]
Aurinia, 188. 197, 199, 208
— corymbosa,
— halimifolia, 197
— petraea, 197
— saxatilis, 197
Austocedri 284, 287, 292
Austrotaxus, 288
Avicennia germinans, 108
Axyris, 465, 468, 469
Azadirachta indica, 108, 125
1987]
Baeothryon, 374
Basanacantha, 173, 174
Bastardia viscosa, 125
Bataceae, 119
BATES, VERNON, and IHSAN A. AL-SHEHBAZ.
Armoracia lacustris oo the
Correct Name for the h American
Lake Cress, ce 359
Batis maritima, 119
Berteroa, 190, “197, 207-210
— gintlii, 208, 20
— incana, 186, 208, 209
— macrocarpa, 208
Bertiera, 181, 183
Bidens cyanapiifolia, 121
Bignoniaceae, 119
Black-headed sedge, 417
Black-potato, 6
Bladderpod, 2 se
Blechum browne), 118
Blephilia ee 12, 14, 20, 21, 23, 32
Boerhavia coccinea, 126
Boraginaceae, 119, 135
ee 204
arborescens, 121
Bothrochloa ischaemum, 116
7
— pe
Boneainvilles glabra, 126
urreria succulenta, i6o 119
Brachiaria adspersa, 117
— ears 117
— reptans, 117
Brassica a
— ae var. ae 122
r. capitata, 12
Bekteaee: Actiomeia lacustris, the Cor-
rect Name for the North American Lake
Cress, 357-359
Brazoria, 1, 4-6, 9, 13, 15, 17, 18, 26-28
— scutellarioides, 5, 12-16, 20, 21, 32
— truncata, 12, 14-16, 20, 21, 32
INDEX
481
Breynia disticha, 122
Bromeliaceae, 37, 38, 116
Bryophyllum pinnatum, 122
Buchingera, 18
Buffalo-grass, 57
Bulbostylis, 363, 370, 391, 393-395, 397
— barbata, 393, 394
— capillaris, 393, 394
_ ciliatifolia, 393
_ stenophylla, 393
Bulrush, 373
Bumelia obovata, 130
— salicifolia, 130
Buriadia, 293
Bursera simaruba, 120
Burseraceae, 120
Buttonbush, 168
Buxaceae in aa Sean United States,
The, 24
Buxaceae, ae oe
3
Bees lucida, 109, 124
Cactaceae, 120
Gacsaipints bonduc, 123
— coriaria, 123
— crista, 110
— divergens, 110, 123
Cajanus c
Callisia cian f 16
9
Calyptranthes bling 111
— kiaerskovii,
Calyptrostylis, ‘i
Camelina, 188, 190, 234-240
— sect. Pseudolinum, 234
— alyssum, 236
— anonale. 234, 235
— glabrata, 235
482
Camelina hispida, 235, 236
— laxa, 235
— microcarpa, mee A 236
— rumelica, 235,
Canavalia rosea, 124
Canella,
- alba. 120
— winterana, 109, 120
Canellaceae, ie
“ANTINO, PHIL d Moness S.
ABU- ASAB. Phylogenetic Implications of
Leaf Anatomy in Subtribe Melittidinae
(Labiatae) and Related Taxa, 1-34
tata, 120
Capraria ee 130
Capsella, 235
Capsicum frutescens, 130
Cardaria draba, 2
Cardiospermum corindum, 130
Carex, 362-364, 371, 373, 396, 397, 424—
445
— subg. Altericarex, 433
— subg. Carex, 427-429, 432
sect. Acrocystis, 430, 433
sect.
— sect. Anomalae, 435
sect.
sect. Carex
sect.
sect.
sect. C
sect.
sect.
sect.
sect.
sect.
sect.
sect. Paludosae, 430, 437
sect. i
sect.
sect.
sect.
sect.
oO
Granulares, 434, 435
Hymenochlaenae, 435
34
Phacocystis, 429, 433, 436
Phyllostachyae, 433
Pictae, 433
JOURNAL OF THE ARNOLD ARBORETUM
Carex subg. Carex sect. Polytrichoideae, 43
— sect. Pseudo-cypereae, 437
— sect.
— sect.
— sect.
— sect.
— sect.
. Heleoglochin, 426, 431
. Macrocephalae, 431
. Multiflorae, 426, 431
: hese 427
: haestoglochin, 426, 431
Stelluatae, 427, 432
sect. Sylvaticae, 435
— aenea,
aestivalis, 435
alba, 434
angustior, 432
— blanda, 434
[VOL. 68
3
1987]
Carex brevicollis, 364
— bromoides, 426, 432
— brunnescens subsp. sphaerostachya, 426,
432
_ bullata, 437
— collinsii, 436
| |
aad
ae
z 8
Ss =
es)
pot
a
Ww
©
— hyalinolepis, 437
INDEX
Carex incomperta, 432
— ee 434
— limosa, 4
_ ceo 437
— mitchelliana, 436
— mohriana, 432
— muricata, 431
— nigromarginata, 433
— oligocarpa, 43
— panicea, 434
, 430
pedunculata, 430, 433
— pendula, 436
— pensylvanica, 433
— picta, 433
— plantaginea, 430, 434
— platyphylla, 430, 434
— prasina, 435
pseudocyperus, 437
427
|
SS
E
=
te)
et.
nw
— rectior, a
—
“a
g
°
i")
oO
Fal
&
ap”
a
-
WwW
— riparia, 43
| |
=
e 8
2.4
BFS
s
BS
we
~
| |
ae
g3
a3
t
iana, 4
— sparganioides, 431
— spicata, 4
— sprengelii, 435
483
484 JOURNAL OF THE ARNOLD ARBORETUM [voL. 68
Carex squarrosa, 437 Casuarina equisetifolia, 109, 120
— stans, 430 Casuarinaceae,
— stipata, 432 Catesbaea, 138, 142, 173, 182, 183
— straminea, 432 — elanocarpa, 82
— striata, 435 — parviflora, 182
— stniatula, 434 — spinosa, 182, 183
— stricta, 43 Catharanthus roseus, 119
— strictior, 436 Cathaya, 2
— swanil, 43 Cedrus
sylvatica, 435 Ceiba pentandra, 119
— tenax, Celastraceae, 120, 242
— tenera, 432 — ser. Buxaceae, 242
— tetanica, 434 Celosia nitida, 118
orta, Celtis iguanaea, 13
— triangularis, 431 Cenchrus echinatus, 117
— tribuloides, 432 — gracillimus, 117
— trichocarpa, 437 — incertus, 117
— trisperma, 432 — tribuloides, 117
— typhina, 437 Central America, A New io eae of Pinus
— umbellata, 430 from Mexico and, 447-459
— vaginata, 434 Centrosema virginianum, 124
— verrucosa, 436 Cephalanthus, 138, 139, 142, 167-172
— vesicaria, 437 — angustifolius, 168, 171
— vexans, 432 — glabratus
— virescens, 43 — natale l
— vulpinoidea, 426, 431 — occidentalis, 168-171
— walteriana, 435 _ subsp. californicus, 169
— willdenovil, 433 — — var. californicus, 169
— woodii, 434 — — var. pube 169
Carica papaya, !20 — — f. lanceolatus, 169, 171
Caricaceae, 120 — salicifolius, 168, 170
Caricopsis, 364 — tetrandra, 168
Casasia, 138, 139, 142, 176-178 Cephalocereus nobilis, 120
— acunae, 176 Cephalotaxaceae, 269-271, 275, 280, 283,
- canna 176 288
— chiapensis, 176 Cephalotaxus, 275, 280
- clusiifolia, 176, 177 Ceratocarpus, 466, 468, 469
— domingensis, 176 Ceratoides, 4
— ekmanii, 176 Cereus intortus, 120
— haitiensis, 176 Chamaecrista glandulosa var. swartzil, 124
— jacquinioides, 176 Chamaecyparis, 284, 2
— longipes, 176 sega as oD
— nigrescens, 176 — eee 2
— parviflora, 176 — hirt
— piricarpa, 176 — ee aye
— samuelssonii, 176 — mesembrantflia 122
Cassia obovata, 124 — multinodis
— occidentalis, | — pilulifera i
Cassytha fihiounis. 110, 123 — prostrata, 122
Castanea, 76, 102 CHANNELL, R. B., and C. E. Woop, Jr. The
Castanopsis, 74—76, 102 Buxaceae in the Southeastern United
— sect. Pseudopasania, 75 States, 241-257
uminatissima, 7 Cheirolepidiaceae, 291
Gastcla erecta, 109, 130 Chelonopsis, 4-6, 9, 28
1987]
Chslonepe's forrestii, 12, 14, 20, 21, 32
moschata, 12, 14, 32
Ghanonedieeie 121, 461
— tribe Atripliceae, 461, 468
Chenopodiaceous Genus from China, a
New, Archiatriplex, 461-469
Chenopodium murale, |
China, a New Chenopodiaceous Genus
from, Archiatriplex, 461-469
Chloris gayana, 117
— inflata, 117
Chlorocharis, 388
1
Chrysobalanus icaco, 109, 121
Chrysochamela, 235
Cuu, Ge-uin. Archiatriplex, a New Cheno-
podiaceous Genus from China, 461-469
Cinchona, 139, 145, 166
— subg. Exostema, 165
Cinchonoideae (Rubiaceae) in the South-
eastern United States, The Genera of,
137-183
Cissus verticillatus, 110, 131
ee fruticosum, 131
subserr
eae eis, 129
ntium, 129
_ — paradisi 129
— sinensis, 129
Cladistic Analysis of Conifers, A: Prelim-
inary Results, 269-307
Cladium, 363, 364, 370, 418-420
— fragrans, 1 l
— umbellatum, 11
Club-rush, 373
Clusia rosea, 123
Clypeola, 188
_ ao 196
— aspera, 188
_ Eas 196
INDEX
485
Clypeola lappacea, 188
Coccoloba, 114
— diversifolia, 126
_ ce 109, 126
x Coccoloba uvifera, 126
_ pie ee
— uvifera, 108, 109, 126
ae barbadensis, 110, 118
_ , 110
Cochleatia, 235, 357
57
r. aquatica, 358
Cocos nucifera, 118
Codiaeum variegatum, 122
Coleus ak nicus, 10
— blum
Sica. oe 116
Colquhounia, 5
Colubrina arborescens, 127
Commelinaceae, 37, 38, 116
Comocladia So ia 109, 118
— ilicifolia,
peers Study of Root and Stem
ods of Some Members of the Mim-
osoideae (Leguminosae), A, 349-355
Compositae, 121, 138
Coniferae, 269
— suborder Pinineae, 269
— suborder Taxineae, 270
Conifers, A Cladistic Analysis of: Prelim-
inary Results, mea
Conocarpus erecta, 108,
Contributions to a Flora ‘of Anguilla and
Adjacent Islets, 105-131
Q
°
s
<
2.
<
S
rs)
°
oO
s
a)
, 1896-1986, Some Bo-
tanical Reminiecerces of, 471-478
Cooley, George R., 471-478
Corchorus hirsutus, 130
Cc
Crataegus, 336
486 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 68
tone cujete, 119 Cyperaceae subfam. Caricoideae tribe
Crinum, 115 Scleriae, 363, 420, 427
Ce ci rhacoma, 120 — subfam. Cyperoideae, 363, 371
Crotolaria incana, 124 — — tribe Cypereae, 363, 395, 416
— retusa, 124 — — tribe Hypolytreae, 363, 427
— verrucosa, 124 — — tribe Schoeneae, 363, 364, 413
Croton, 108, 242 — — tribe Scirpeae, 363, 371, 391
— betulinus, 122 — subfam. Rhynchosporoideae, 429
— fishlockii, 111 — subfam. Scirpoideae,
— flavens, 109, 122 — tribe Abildgaardieae, 391
— lobatus, 122 — tribe Dulichieae, 416
— microcarpus, 123 — tribe Fimbristylideae, 363, 391
— nummulariaefolius, 123 — tribe Mapanieae, 363
— ovalifolius, 123 — tribe Rhynchosporeae, 363, 413
Cruciferae; Brassicaceae: The Genera of Cyperus, 362, 364, 370, 373, 395-407, 409,
Alysseae in the Southeastern United 411, 416, 421, 424, 429
States, 185-240 — subg. Cypenis. 396- 398, 400, 402
Cruciferae, 122, 187, 188, 191, 192, 204, — — sect. Compressi, 401
211, 216, 217, 226-229 — — sect. Cyperus, 400
— tribe Alysseae, 185-240 — — sect. Irioidei, 401
— — subtribe Lunariinae, 191 — — sect. Laxiglumi, 400
— tribe Alyssineae, 185 — — sect. Umbellati, 400
— tribe Arabideae, 187, 359 — — sect. Viscosi, us
— tribe Camelineae, 186, 235 — subg. Fimbricyperu
— tribe Drabeae, 186, 187, 228, 359 — subg. Juncellus, 396, 397, 399
— tribe Heliophileae, 187 — subg. Kyllinga, 408
— tribe Lepidieae, 187, 228, 235 — subg. Mariscus, 397, 398
— tribe Lunarieae, 186, 187, 191 — subg. Protocyperus, 396
— tribe Schizopetaleae, 228 — subg. a 396-398, 402
— tribe Sisymbrieae, 235 — — sect. Fusci, 399
— tribe Thelypodieae, 191, 228 — — sect. Haspani, 399
Cryptomeria, 284, 286 — — sect. Luzeoloidei, 398, 402
Cryptostegia erandifior, 126 — subg. Pycreus, 396-399
Cucumis anguria, 122 — subg. Torulinium, 396-398
Cucurbita nar 122 — — sect. Diclidium, 402
Cucurbitaceae, 122 — — sect. Remirea, 402
Cunninghamia, 284, 286 — sect. Esculenti, 400
Cupressaceae, 269-271, 278, 280, 283, 284, — sect. Glutinosi, 401
286-291, 293, 294 — sect. Iriae, 401
Cupressus, 278, 280, 284, 287 — sect. Rotundi, 400
— lusitanica, 4 — albomarginatus, 399
uscuta a americana, 110, 121 — bipartitus, 399
Cyclobalanops — brevifolioides, 408
Cymbopogon ak 117 — calcicola, 116
Cymodoceaceae, 116, 260 — compressus, 401
Cymophyllus, 363, 371, 422-425 — correllii, 402
— fraseri, 423, 424 — croceus, 400
Cynanchum anegadensis, 110 — dentatus, 399
— parviflorum, | — var. multiradiatus, 399
ge in the Southeastern United — aia 396, 399
s, The Genera of, 361-445 — diffusu
Cyperaceae 116, 361-445 oa dipeacilornie 400
subfam. Caricoideae, 363, 420 — echinatus, 400
— — tribe Cariceae, 363, 422, 427 — elegans, 401, 402
1987]
Cyperus engelmannii, 402
— eragrosti
s, 398
= eiculentus 396, 397, 399-401
Pp
— houghtonii, 398
— hystricinus, 400
>
laevigatus, | 16, 399
!
lentiginosus, 400
longus, 396
jouisianensis; 399
pedunculatus, 402
planifolius var. brunneus, | 16
plukenetii, 400
pollardii, 400
polystachyos, 399
wilburii, 401
Dacrycarpus, 285
Dacrydium, 285
INDEX 487
Dactyloctenium aegyptium, 117
Datura stramonium, 13
Decussocarpus, 285
Delonix regia, 124
Dendropemon caribaeus, 110, 124
Desmanthus virgatus,
Desmodium frutescens var. angustifolium,
124
Dichromena, 413
Dichrostachys cinerea, 349, 350, 352-354
Dieffenbachia seguine, 116
Digitaria bicornis, 117
— decumbens, 117
Diplazium legalloi, 113
Diselma, 284, 287, 292
Dithyrea, 228
Dog’s-hair grass, 384
Dolichostylis, 211
minica, Lesser Antilles, A New Species
of Lantana (Verbenaceae) from, 343-348
Dorella, 234
Draba, 186-188, 190, 191, 210-223
— sect. Leucodraba, 214
— sect. Phyllodraba, 214
— t. Tomostima, 214
—_ a. 216
— alpina, 216
|
9
3
3
°
so)
St
cy
i‘)
—c
— cuneifolia, 213- 216
— — var. cuneifolia, ra 215
— — var. foliosa,
— — var. helleri a7)
— — var. integrifolia, 215
_ =~ densttolia. 216
488
Draba dentata, 212
— exunguiculata, 216
— macrocarpa, 217
— micrantha, 214
— nemorosa, 217
_ ie 217
— olgae, 217
= ieee 216
|
|
<
rt)
ca)
pik
ra
om
=F
S
5
i)
N
a, 216
— verna, 211, 215-217
_ Aaa 215
Drabella, 21
Dulichium, oa 370, 415-417
— arundinaceum, 416
Duranta erecta, 131
— plumieri, 10
Dyssodia tenuifolia, 121
Eichhornia, 36, 38, 40, 41, 49-57, 65, 67,
68
,71
— sect. Eichhornia, 50
— sect. Eueichhornia, 50
— sect. RE a aeE 50
— azurea, 41, 49, 50, 53
— crassipes, 38, o 41, 49-54, 65, 67
— natans, 49,
— paniculata, 41, 50, 67
— par a, 50
Elaeodendron xylocarpum, 120
Eleocharis, 363, 370, 373, 384—390
— ser. Eleocharis, 385, 388
— ser. Maculosae,
— ser. Mutatae, 385
— ser. Ovatae,
— ser. Palustriformes, 385, 388
JOURNAL OF THE ARNOLD ARBORETUM
[VOL. 68
Eleocharis ser. Pauciflorae, 385
— cellulosa, 385, 386
— confervoides, 388
— dulcis, 385
— elongata, 385
— engelmannu, 387, 388
— equisetina, 3
— equisetoides, 385
— fallax, 385
— flavescens, 387, 388
=]
oF
Oo
5
n
c.
So
OQ
So
»
ie)
oo
a
— palustris, 384
— subsp. palustris < subsp. vulgaris,
— parvula, 385
— ee 385
— radicans, 387
— sane 385
— — subsp. anisms. 388
— wolfii, 387
Eleusine indica, 117
468
Enterolobium saman, 349, 350, 352-354
Ephedra, 282
Epidendrum bifidum, 118
— kraenzlini, 118
Epiphyllum oxypetalum, 120
Eragrostis ciliaris, 117
— tenella, 117
Sn 1, 23
riope, 9
1987] INDEX 489
Eriophorum, 370, 380-382 Fauria crista-galli, 134
— sect. Eriophorum, 380 Fever tree,
— sect. Phyllanthela, 380 Fibigia, 187, 208
— sect. Mager 380 Ficus emery 108, 125
— alpinum, 381 — elastica
— aNeNea ole 380, 381 Fimbristylis, aa 364, 370, 384, 385, 390-
— vaginatum, 380, 381
— virginicum, 380, 381 — sect. Dichelostylis, 392
— viridicarinatum, 381 — sect. Fimbristylis, 392
Erithalis fruticosa, 108, 127 — sect. Trichelostylis, 392
Ernestiodendron, 278, 282 — annua, 392
Ernodea littoralis, 127 — autumnalis, 392
pases 211, 215, 216 — caroliniana, 392
— verna, 215 — castanea, 392
_ SEA 215 — complanata, 392
Erythrina variegata var. orientalis, 124 — cymosa subsp. spathacea, 116
Eucyperus, 39 — decipiens, 39
Eugenia axillaris, 109, 125 — dichotoma, 391, 392
— foetida, 109, 126 — ferruginea, 11
— monticola, 126 — miliacea, 392
Eupatorium odoratum, 121 — monostachya, 116
Euphorbia cyathophora, 123 — ovata, 116
_ eee 123 — perpusilla, 392
— lactea, 123 — puberula, 392
_ ei 123 — schoenoides, 392
— pulcherrima, 123 — spathacea, 116, 392
— tirucalli, 123 — thermalis, 391
Euphorbiaceae, 122, 242, 243 — tomentosa, 392
Eurotia, 466, 469 — vahliu, 392
Eurystemon, 37, 57, 71 Firebush, 179
Evolvulus antillanus, 121 Fishlockia anegadensis, 110
— argyreus, 121 Fitzroya, 284, 287, 292
_ Soa oases 121 Flagellariaceae, 362
— glaber, 121 Flat-sedge, 396
— sericeus, 121 Flaveria bidentata, 121
Exallage, 154 Flaxweed,
Exomis, 466, 468 Flora of Anguilla and Adjacent Islets, Con-
Exostema, 109, 142, 165-167 tributions to a, 105-131
— caribaeum, 109, 127, 165, 166 Fokienia, 284, 287
— longiflorum, 165 Forestiera eggersiana, 126
— parviflorum, 165 Fraser’s sedge,
Freziera (Theaceae), Taxonomic Studies in,
Fagaceae: Reproductive Structure of with Notes on Reproductive Biology,
Lithocarpus Sensu Lato: Cymules and 323-334
Fruits, 73-104 Freziera, 323-334
Fagaceae, 73, 74, 77 — candicans, 324
Fagara trifoliata, 129 — canescens, 324
Fagus sylvatica, 102 — carinata, 325-328
Falcatifolium, 285 — chrysophylla, 324, 330
False DD: — echinata, 328
Farsetia, 187, 204 — euryoides, 333
incana, guianensis, 328
— somalensis, 187 — microphylla, 333
— undulicarpa, 187 — minima, 331-33
490
Freziera parva, 333
— tomentosa, 330
— umbellata, 382, 383
Furcraea, 108
Galactia dubia, 124
Galeobdolon, 1, 9, 11, 23, 28
— luteum, 12, 14, 20, 21, 32
Galingale, 396
Galitzkya, 208
Galphimia gracilis, 125
Gamosepalum, 195
Gardenia, 174
Genera of Alysseae (Cruciferae; Brassica-
ceae) in the Southeastern United States,
The, 185-240
Genera of Cinchonoideae (Rubiaceae) in
the Southeastern United States, The,
137-183
Genera of Cyperaceae in the Southeastern
United States, The, 361-44
Genera of Pontederiaceae in the =
eastern United States, The, 35-7
Genipa,
— clusifolia, 176
Georgia bark, 143
Ginkgo, 278, 280, 282, 290
Gliricidia sepium, 124
Glyce, 204
Glyptolepis, 278
Glyptostrobus, 284, 286
Gnetu
Gold- of pleasure 234
Gomphostemma,
Pee bee 123
Gossypium barbadense, 125
Graellsia, 22
Gramineae, 116, 362, 364
Grayia, 466, 468
GRETHER, Rosaura. Taxonomic and No-
JOURNAL OF THE ARNOLD ARBORETUM
[VOL. 68
menclatural Notes on the Genus Mi-
mosa (Leguminosae), 309-322
Grubbia rourkei, 353
Guaiacum officinale, 131
Gyminda latifolia, 109, 120
Gymnanthes lucida, 123
ymnospermae, 115
Haemodoraceae, 37, 39
— tribe Conostylideae, 40
— tribe Haemodoreae, 40
Halocarpus,
Hamelia, 138, 139, 142, 178-182
9
— — var. patens, 179
HARDIN, JAMES W., and FREDERICK G.
MEYER. Status of the Name Aesculus fla-
va Solander (Hippocastanaceae), 335-
341
Hart, JEFFREY A. A Cladistic Analysis of
Conifers: Preliminary Results, 269-307
Haynes, Ropert R., and Lauritz B.
LM-NIELSEN. The Zannichelliaceae in
ie Southeastern United States, 259-268
Hedyotis, 138, 139, 142, 146-163
— subg. Houstonia, 150, 152
_ subg. ae 152, 163
— affinis,
_ surcularia, 147, 153-155
15
153
corymbosa, 148, 150, 152, 153, 155
= Graciela | 3
— diffusa, 155
— fasciculata, 152
— fruticosa, 153-155
— halei, 163
— herbacea, 148, 153, 155
— lancifolia, 14
— longifolia, 153
— nuttalliana, 153
— ouachitana, 153
— procumbens, 153
1987]
Hedyotis purpurea, 151-153
— rosea
— salzmanii, 152
— scandens, 155
— uniflora, 148, 152
Heliotropium angiospermum, 119
— curassavicum, 119
10
— micrantha var. aristulata, 410
— occidentalis, 410
Hibiscus rosa- sinensis, |
— sabdariffa, oe
et ener Sta of the a
s flava ade. 335-34
aioe mancinella, 108, a
Hoary alyssum, 208
Hoffmannia, 138, 180, 181,
Holargidium, 211
Hotm-NIE.Lsen, LAuritz B., and ROBERT
. Haynes. The Zannichelliaceae i in the
Southeastern United States, 259-268
ee on sanguinea, | |
Honesty, 19
Cee EST 187, 198
Horned pondweed, 264
183
INDEX
ee 147- 152
_ serpyllacea, 148
— a eet scaphia: 149
Howarp, RicHAarD A. Some Botanical
Reminiscences of George R. Cooley,
1896-1986, 471-478
Howarp, RICHARD A., and
KELLOGG. Contributions to a Flora of
Anguilla and Adjacent Islets, 105-131
Howarb, RICHARD A., ELIZABETH A.
Ke_Locc. Unusual Pollen Dimorphism
in Rondeletia anguillensis (Rubiaceae),
133-136
Hybanthus portoricensis, 131
Hydrocharitaceae, 1
ae 35-37, 40, 71
— gardne
ices undatus, 120
Hymenocallis caribaea, 115
Hypelate trifoliata, 130
Hyptis suaveolens, 10
ELIZABETH A.
Indigo berry, 173
oe suffruticosa, 124
— tinctoria, |
Ipomoea eee 122
— batatas, 12
— carnea subsp. fistulosa, 122
— eggersii, |
— nil, 122
— pes-caprae subsp. brasiliensis, 122
— triloba, 12
Ixora, 168
— casei, 127
— coccinea, 127
Jacquemontia cayensis, 122
, 122
Jacquinia ab bores. 109, 130
— berterii, 109, 1
492
Jasminum fluminense, 126
Juncaceae, 362-364
Juncellus, 396
Juncus, 375
Juniperus, 284, 287, 292
Kalanchoé blossfeldiana, 122
— tubiflora,
Kallstroemia maxima
KAUL, ROBERT B. ie Structure
of Lithocarpus Sensu Lato (Fagaceae):
Cymules and Fruits, 73-104
KELLOGG, ELIZABETH A., and RICHARD A.
Howarp. Contributions to a Flora of
Anguilla and Adjacent Islets, 105-131
KELLOGG, ELIZABETH A., and RICHARD A.
Howarp. Unusual Pollen Dimorphism
in Rondeletia anguillensis (Rubiaceae),
and K. RANJANI.
A Comparative Study ‘of Root and Stem
Woods of Some Members of the Mim-
osoideae (Leguminosae), 349-355
Krugiodendron ferreum, 127
Kyllinga, 364, 370, 408, 409, 411
— brevifolia, 408
— brevifolioides, 408
— squamulata, 408
— tibialis, 408, 409
— vaginata, 408
Labiatae: sy a Implications of Leaf
e Melittidinae and
34
Labiatae, 1, 3, 9, 10, 12-15, 23, 25, 26, 33,
123
— subfam. Lamioideae, 2, 3, 5, 10, 12, 14,
20, 21, 23
JOURNAL OF THE ARNOLD ARBORETUM
[VOL. 68
Labiatae subfam. Lamioideae tribe Lam-
ieae, 2-5, 10-12, 14, 18, 20, 21, 26-28
— — — subtribe Melittidinae, 1- 34
— — tribe Prasieae, 2—4, 18
— subfam. Nepetoideae, 2, 3, 10-12, 14,
26
— — tribe Ocim
— — tribe Salvieae, 10
— subfam. Stachyoideae, 3
— — subtribe ae 1, 23
— tribe Prostanthereae, 2
Laguncularia racemicsa: 108, 121
Lake Cress, Armoracia lacustris (Brassi-
caceae), the Correct Name for the North
American, 357-359
Lamium, 5, 9, 23
— purpureum, 12, 14, 20, 21, 32
Lantana (Verbenaceae) from Dominica,
Lesser Antilles, A New Species of, 343-
34
mise, 343, 346
ect. Camara, 344, 346, 347
— ere
_ camara, 10, 131, 343, 344, 346, 347
8
a, 347
— nHicifolla. 343, 344, 346-348
Larix, 289
Lauraceae, 74, 123
Lavandula burmanu, 10
Leaf Anatomy in ore Melittidinae
(Labiatae) and Related Taxa, Phyloge-
netic Implications of, 1-34
Lebachia, 278, 279, 282, 295
Lebachiaceae, 278, 280, 282, 294
Leea, 337
Leguminosae: A Comparative Study of
Mem-
tural Notes on the Genus Mimosa, 309-
322
1987]
Leguminosae, 123
— subfam. Mimosoideae, 349-355
Leonotis, 23
— nepetifolia, 10, 123
Leonurus,
— cardiaca, 12, 14, 20, 21, 32
ee 57-59
— sect. Alysmus,
|
9
3
i
oI
n
2.
o
=
a
Ss
to
ms
1 |
Cee
5.5
Bum Pie
Ee
per)
a oe
eae
Nubv
NO
Ww
iw)
No
Ne}
a
— densipila, ie 229
— — var. maxima, 226
— — xX Lesquerella stonensis, 226
— douglasii, 229
— engelmannii, 228, 229
— fendleri, 230
| |
ga
ag
ae
“oS
Siu
SS
Se
YS
)
Np
way
— — subsp. gracilis, 223, 224
— — subsp. nuttallii, 224
— var. repanda, 224
— grandiflora, 228, 229
|
<
6s
=
o
oO
ee
©
=]
Q
—_:
oO
A
No
nN
~
— lescurii, 223, 225-227, 229
——x eee has densipila, 226, 227
— ludoviciana, 22
— lyrata, 334, 225, 227, 229
— macrocarpa, 229
5222.
— ovalifolia subsp. ovalifolia, 228
— palmeri, 23
— paysonii, 227
INDEX 493
Lesquerella peninsularis, 228
— perforata, 225-227, 229
— polyantha, 223
— repanda, 224
— rubicundula, 229
— stonensis, 225-227, 229
— — x Lesquerella lescuri, 226
— thamnophila, 2
Lesser Antilles, A New Species of Lantana
(Verbenaceae) from Dominica, 343-348
Leucaena leucocephala, 124, 349, 351-353
Leucas, 9, 23
saree pores 164
— diffus
ae
Linum isi atiseimun 236
Lipocarpha, 370, 409-411
— schomburgkii, 410, 411
— senegalensis, 410
Lippia, 9
— lanceolata, 9, 10
Liquidambar styraciflua, 449, 4
Lithocarpus Sensu Lato (Fagaceae), Re-
roductive Structure of: Cymules and
Fruits, 73-104
76
— subg. rc selante 74, 76, 78, 85, 93,
99, 101
— subg. Cyrtobalanus, 76
— subg. Liebmannia, 78, 82
— subg. Lithocarpus, 76, 78, 81, 101
— — sect. Costatae,
— subg. Oerstedia,
— subg. Pachybalanus, 76, 78, 82
— subg. Pasania, 76, 78, 91, 99- 101
494
wee subg. Pseudocastanopsis, 75,
76, 79, 97, 100
— ae Pecudoamaedoe: 76
— subg. Synaedrys, 76, 79, 82, 101
— sect. Gymnobalanus, 76, 78, 82,99, 101
— aggregata, 78, 86, 91
— amygdalifolia, 78, 82, 83, 99
— beccariana, 78, 80-82, 97
— blumeana, 75
— buddii, 77, 78
— bullata, 78, 87, 88
— caudatifolia, 78
— celebica, 77, 78
— clementiana, 78
— conferta, 78
a, 78
— dealbata, 77, 78, 94, 97, 100
— densiflora, 73, 78, 96, 97, 100
— edulis, 7
— eichleri, 78
— elegans, 77, 79, 95, 98, 100
— elephantum,
— encleisacarpa, 75, 78, 87, 89, 102
— elles 78, 87,
— falcone i, 79
— ae 77, 79, 93, 100, 102
— fissa, 75, 79, 10
= Haviland. 78, 82, 83
— hendersoniana, 78, 80, 82, 97
i, 78
= lampadaria 78, 84, 91, 99
— lapp
Reser 78, 82, 83
— longispina, 7
ee a 78, 84, 85, 99
— lut
— ie 78, 87, 88, 99
— maingayi,
JOURNAL OF THE ARNOLD ARBORETUM
Lithocarpus mariae, 78
— meyeri, 78
— nantoensis, 78, 82, 83, 99
— neorobinsonii, 78, 87, 89, 99
— nieuwenhuisii, 78
— papillifer, 79, 93, 94
— pattaniensis, 78, 90, 91, 99, 102
— pulchra, 79, 81, 82, 97, 101
r J
— recurvata, 75
— reinwardtil, 77, 78, 85, 86, 99
— rufovillosa, 79, 90, 91
— sabulicola, 79, 93, 97, 100
— scortechinil, 79, 92, 95, 97, 100
— sericobalanus, 78
— soleriana, 79, 93, 96, 100
— sootepensis, 77, 79, 91, 93
— truncata, 78, 82, 83, 99
— turbinata, 78, 80, 81, 97
— wallichiana, 79, 95, 98, 99
— wrayi, 79, 92, 93, 99, 100
Lithophila muscoides, 118
Lithospermum, 135
Lobularia, 187. 189, 198, 203-208
— arabica, 204, 20
— intermedia, 204, 205
— libyca, 204, 205
— marginata, 205
— maritima, 186, 188, 204—206
palmensis,
— spath 05
— alpina, 191
— annua, 186, 191-193
— — subsp. annua, 191
— subsp. pachyrhiza, 191
l
ata, 191
es 191-193
— telekiana, 19]
Lycium americanum, 130
Lycopersicon lycopesicum, 130
Lythraceae, 40,
[VOL. 68
1987]
INDEX
Macbridea, 1, 4-6, 9, 13, 15, 26-28
— alba, 12, 13, 17, 19-21, 28, 32
— caroliniana, 12, 13, 19-21, 32
Machaerina, 419
Macropodium pterospermum, 191
Madwort, 19
Malpighia emarginata, 109, 125
M
M
M
2
angifera indica, 108, 118
anihot esculenta, 123
arantaceae, 63
Mariscus, 396, 397
MEYER, FREDERICK G., and JA
subsect. Laxiglumi, 400
arrubium, 9, 13, 23
vulgare, 12-16, 19-21, 32
aytenus élliptica, 120
egacarpaea, 216
elia azedarach, 125
eliaceae, 125
eliococcus bijugatus, 108, 130
elittis, 4, 6,
melissophyllum, 12, 14, 16, 20, 21, 32
elocactus intortus, 120
elochia pyramidata, 130
tomentosa, 130
eniocus, 196
entha viridis, 10
enyanthaceae, 134
erremia dissecta, 110, 122
etabolos, 154
Stascquola: 284, 286, 291
exico and
Central America, A New
Species of Pinus from, 447-459
M
=
Mi
Mi
MES W. HAR-
DIN. Status of the Name Aesculus flava
Solander (Hippocastanaceae), 335-341
icrobiota, 284, 287
icrocachrys, 285
tetragona, 291
erogynoeciom, 466, 468
tibeticu
icromeria een 10
Microstrobos, 285
Microtoena,
Mimosa (Legum
3
Mimosa, 309-322
var. intermedia, 313
Sita ee 314
var. horrida, 313, 314
ealicioni. 318
Ce)
a
=:
}
lacerata, 313, 314
lactiflua, 314, 315
langlassei, 315, 316
mexiquitensis, 312
mixtecana, 314, 315
rekoana, 321
495
inosae), Taxonomic and
Nomenclatural Notes on the Genus, 309-
22
496
Mimosa remota, 310
— stipitata, 318, 319
— ursina
_ vazquezii, 314, 315
— vepres,
— watsoni, 320-322
anti, 316
Mimosoideae (Leguminosae), A Compar-
ative Study of Root and Stem Woods of
Some Members of the, 349-355
— glutinosa, 313, 314
— rhodocarpa, 320
Mirabilis jalapa, 126
Moenchia, 196
Momordica charantia, 122
Monarda fistulosa, 12, 14, 20, 21, 23, 32
— vaginalis, 39
Monocotyledoneae: 115
Monstera acuminata, 116
ceae, 125
Moringa oleifera, 125
Moringaceae, 125
Moschosma polystachyum, 10
Mud-plantain, 57
Murraya paniculata, 129
Musa era 117
Musaceae, | 17
eons sativum, 234, 235
8
, 125
Myrtus anguillensis: 110, 126
Najadaceae, 260
s, 260
ajas
Mechuitinn 357, 359
358
Neocallitropsis, 284, 287
Neomammillaria nivosa, | 20
Neomazaea, 135
JOURNAL OF THE ARNOLD ARBORETUM
[VOL. 68
Neomimosa colimensis, 309
— donnell-smithii, 312
— eurycarpoides, 309
— russellii, 309, 310
Neotchihatchewia, 187
Neptunia pubescens, 124
Nerisyrenia, 228
Nerium oleander, 119
2
New Species of Lantana (Verbenaceae) from
Dominica, Lesser Antilles, A, 343-348
New Species of Pinus from Mexico and
Central America, A, 447-459
Nomenclatural Notes on the Genus Mi-
mosa one aeae and,
309-322
Notobuxus, 242
Nut-rush, 4
Nyctaginaceae, 126
Ocimum, 9
— adscendens, 10
— basilicum, 10, 26
— canum,
— gratissimum, 10
— Pie ate 10
— micranthum 3
— sanctum, 10
Odontarrhena, ae
Odontocyclus,
Oldenlandia, vn 7 153
— affinis, 15
— corymbosa, 150
Oplismenus. hirtellus subsp. setarius, 117
— setarius, 117
Oplonia spinosa, 118
Opuntia cochenillifera, 120
— dillenii, 120
Origanum, 24
Ochosiphon pallidus, 10
Oxalidaceae, 40
Pachysandra, 242, 243, 249-257
— procumbens, 242, 249-252, 254
1987]
ia tee stylosa, 249, 255
— — var. errima, 255
_ eae 242, 243, 249, 254, 255
Paleotaxus jurassica, 288
8
m,
— pani Soniiiuris 117
Passiflora edulis, 126
n
oo
<
oO
img
oO
—
v4
Se
Ww”
Pedilanthus tithymaloides, 123
Peltana,
Pentodon, 138, 139, 142, 150, 162-165
— meine 163
— halei
ee 162, 163
— pentandrus, 163, 1
_ a
Perry, J. P., Jk. A New Species of Pinus
from Mexico and Central America, 447-
459
Persea ee 123
Petrea volubilis
Philydraceae, 37, on 40, 71
INDEX
Phlomis, 1, 9, 23
— bracteosa, 10
Phoenix dactylifera, 118
horadendron trinervium, 110, 124
Phyllanthus amarus, |
— epiphyllanthus, 109, 123
Phyllocladus, 270, 271, 285, 290
Phyllostegia, 5
Phylogenetic Implications of Leaf Anato-
my in Subtribe a ae (Labiatae)
and Related Taxa,
Physalis angulata, Bo
— ore
Physoptychis, 187
Physostegia, 1, 4-6, 9, 13, 15, 17, 18, 26-
28
— angustifolia, 12, 20, 21, 23, 32
— digitalis, 12, 20, 21, 32
— godfreyi, 12, 20-23, 26, 32
— intermedia, 17
— leptophylla, 12, 20-22, 32
— longisepala, 12, 20, 21, 32
— purpurea, 12, 20, 21, 32
— virginiana, 11, 12, 20, 21
— — subsp. praemorsa, 12, 19-21, 32
ubsp. virginiana, 12, 20, 21, 32
Phytolaccaceae, 126
Piaropus, 49
Picea, 289
Pickerel- weed, 63
Pickerel-weed Family, 35
Pilea serpyllifolia, 131
Pilgerodendron, oe 287, 292
Pimenta racemosa,
Pinaceae, 269-272, a. 280, 283, 284,
28
9-292
Pinckneya, 138, 142-146
— bracteata, 143, 144
— pubens, 143, 144
Pinus from Mexico and Central America,
A New Species of, 447-459
Pinus, 278, 280, 289, 447
— sect. Pseudostrobus, 454
451
— estevezil, 447, 451, 453-457
l
— nubicola, 447-4
— oaxacana, 447, 449-457
498 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Pinus oocarpa var. ochoterenae, 449, 450 Pontederia cordata var. ovalis, 65, 66
— patula, 449 — hastata, 63,
— — var. longepedunculata, 449 — lanceolata, 38, 66
— pseudostrobus, 447, 449, 451-457 — lancifolia, 66
— — var. apulcensis, 45 — ovata, 63
— — var. estevezil, 451 — parviflora, 63, 65, 66, 68
— — var. oaxacana, 451 — rotundifolia, 41, 65, 67, 69
— rudis, : — sagittata, 4 5. 8
— tecunumanii, 450 — subovata, 41, 68
Pisonia subcordata, 108, 126 Pontederiaceae in the Southeastern United
Pithecellobium dulce, 349, 351-353 States, The Genera 35-71
— unguis-cati, 109, 110, 124 Pontederiaceae, 35—7
Platycladus, 284, 287 — tribe Seen ae 36, 39, 49
Plectranthus, 9 — tribe Heteranthereae, 36, ag 39, 57
— amboinicus, 123 — tribe Pontederieae, 36, 39, 63
— australis, 10, 26 Portulaca halimoides, 126
— blumei, 123 — oleracea, 126
— incanus, 10 Portulacaceae, 126
— mollis, | Posidoniaceae, 260
Pluchea odorata, 1 Possum
— symphytifolia, 121 Potamogeton, 60
Plumbaginaceae, 126, 135 Potamogetonaceae, 260
Plumbago auriculata, 126 Princewood,
— scandens, 110, 126 Priva lappulacea, 131
Plumeria alba, 109, 119 Prosopis ieee 349, 351-353
— rubra, 119 Prostanthera, 9, 13, 27
Poa pratensis, 430 _ andl. 12, 13, 20, 21, 23, 28, 32
Podocarpaceae, 269-272, 275, 278, 280, Prumnopitys, 28
282-285, 289, 290, 292-294 Prunella, 11
Podocarpus, 275, 280, 285, 293 Pseudalthenia, 260, 261
Podranea ricasoliana, 119 — aschersoniana, 260
Pogonopus, 144, 145 Pseuderanthemum carruthersii var. reticu-
— exsertus, 144 latum, 118
— speciosus, 144 Pseudobraya, 2
— tubulosus, 144 Pseudogynoxis one 121
Pogostemon, 11, 15, 17, 23, 34 Pseudolarix, 283, 289
— cablin, 12, 14, 20, oe 23, 32, 34 Pseudotaxus, 272, 288, 291
— heyneanus, 34 Pseudotsuga, 283, 28
— parviflorus, 10 Pseudovoltzia, 278
— purpurascens, 10 Psidium guajava, |26
Pollen Dimorphism in Rondeletia anguil- © — longipes var. orbicularis, 110, 126
lensis mere oo 133-136 Psilocarya, 41
Polygonaceae, 126, 3 Psilonema, 196
Polyscias fruticosa, ae — alyssoides, 196
— guilfoylei, Ptilotrichum, 19
Pontedereae, 3 Punica granatum, 127
Pontederia, 36-39, 41, 49, 63-71 Punicaceae, |
— subg. Pontederia, 65-67, 69 Pycreus, 396
Bh a Quercus, 73-76, 81, 101, 102, 450
ibis seins. — subg. Cyclobalanopsis, 74, 75
sain a4 — subg. Quercus, 74, 75
— cordata, 38-41, 63-69 ene 3
— — var. cordata, 65-68 Radicula, 357
— — var. lancifolia, 65-68 — aquatica, 358
1987]
Randia, 138, 139, 142, 172-175, 183
— subg. Basanacantha, 174
— aculeata, 109, ne 127, 173, 174, 183
— clusiifolia, 17
— formosa, 174
— mitis, 173
— rhagocarpa, 173
ANIJANI, K., and K. V. KRISHNAMURTHY.
A Comparative Study of Root and Stem
Woods of Some Members of the Mim-
osoideae (Leguminosae), 349-355
Rauvolfia viridis, 119
Reed, 373
Remirea, 396, 402
— maritima, 402
Reproductive Biology, Taxonomic Studies
in Freziera (Theaceae), with Notes on,
Reproductive Structure of Lithocarpus
Sensu Lato (Fagaceae): Cymules and
04
a, 39
Reynosia uncinata, 109, 110, 127
Rhacoma crossopetalum, 120
Rhamnaceae, 127
Rhaphidophora aurea, 116
Rhizophora mangle, 109, 127
0, 124
Rhynchospora, 363, 370, 373, 388, 413-
416, 424
— subg. Eurhynchosporae, 414
— subg. Rhynchospora, 41
— sect. Dichromena, 409, 412, 414
— alba, 413
— colorata, 412
— floridensis, 412
Ricinus communis, 123, 230
Rochefortia acanthophora, 120
GERS, GEORGE K. The Genera of Cin-
chonoideae (Rubiaceae) in the South-
eastern oo States, 137-183
Roigella, |
INDEX
499
Rondeletia aan (Rubiaceae), Un-
n Dimorphism in, 133-136
Rondeletia, 135
— anguillensis, 105, 110, 127-129, 133-
136
Root and Stem Woods of Some Members
of the Mimosoideae (Leguminosae), A
Comparative Study of, 349-355
Rosa indica, 127
Rosaceae, 127
RosaAtTtI, THOMAS J. The Genera of Pon-
tederiaceae in the Southeastern United
States, 35-71
Rubiaceae: The Genera of Cinchonoideae
in the Southeastern United States, 137-
183
Rubiaceae: Unusual Pollen Dimorphism
in Rondeletia anguillensis, 133-13
Rubiaceae, 40, 127, 133, ee 137-139,
144, 145, 151, 170, me
— subfam. Cinchonari -
— subfam. encciene se 137-183
— — tribe Catesbaeeae, 138, 183
— — tribe Cephalantheae, 170
— tribe Condamineeae, 138, 142, 144,
145
— — — subtribe Pinkneyeae, 144
— — tribe Gardenieae, 138, 172-174, 179,
183
— — tribe Hedyotideae,
]
138, 146, 150,
5
— tribe Naucleeae, 138, 167, 170
Ruellia tuberosa, 118
Ruppia maritima, 118
Ruppiaceae, 118
Rutaceae, 129
Saccharum officinarum, 117
Salazaria, 3, 4, 18
500
Salicornia bigelovii, 121
— herbacea,
Salvia occidentalis, 123
— ple
arn 10
— serotina, se
ANDERS, ROGER W. A New Species of
Lantana ea from Dominica,
Lesser Antilles, 343-348
Sansevieria hyacinthoides, 115
— trifasci 115
— guatemalensis, 242
_ eae 242
— salign
Scag domingensis, 127
91
8
Scaevola plumieri, 107, 109, 123
Schaefferia frutescens, 120
Schlechtera, 187
Schoenoplectus, 374
Schoenus, 370, 417, 418
— albus, 413
— ferrugineus, 418
|
=
io)
s
wn”
QO
g
Seholleropss, 36, 37, 71
_ a, 39
ice eae 269, 283
Sciadopitys, 269, 278, 280, 283, 284, 286,
28
Scirpus, 363, 364, 370-381, 384, 388, 411,
429
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— acutus, 376
— americanus, 375, 376
Actaeogeton, 377
76
Bolboschoenus, 375, 385
Isolepis, 377
Junco-scirpus, 375, 376
Oxycaryum, 375
Pterolepis, 375, 376
ren ee 375
Scirpus, 37
oe 374
Trichophorum, 375
Vaginati, 380
JOURNAL OF THE ARNOLD ARBORETUM
Scirpus atrocinctus, 375
_ lea 375
— deltaru
— etuberculatus, 375
— expansus, 374
— flaccidifolius, 374
— fontinalis, 375
— georgianus, 374
— hallu, 377
— hattorianus, 374
— lacustris, 375
—- ea 373, 374
— curtissi, 421
— flagellum-nigrorum, 420
— foliosa, 421
1987]
Scleria georgiana, 421
— hirtella, 421, 422
— leptostachya, 421
— lithosperma, 116, 421
— tesselata, 421
— triglomerata, 421, 422
— verticillata, 42
Scrophulariaceae, 130
Scutellaria, 2-5, 9, 11, 13, 15, 17, 18, 23,
2
— elliptica, 12-14, 16, 20, 21, 26, 32
— incana, 12, 14, 19-21, 33
_ integrifolia, 9, 12, 14, 20, 21, 33
— laterifolia, 7, 12, 14, 20, 21, 33
— nervosa, 12-14, 16, 20, 21, 33
— ovata, 12-14, 20, 21, 26, 33
— serrata, 12-14, 20, 21, 33
Sedge, 427
Sedge Family, 361
Sedge-grass, 396
Selenia, 187, 191
Senna bicapsularis, 124
2
_ oe ere 124
— siamea, |
Sequoia, 284, eh
Sequoiadendron, 284, 286
Sesbania grandiflora, 124
Seven-year-apple, 176
Solanaceae, 130
INDEX 501
Solandra guttata, 130
Solanum melongena, 130
— racemosum, 130
Solidago microglossa, 121
Some Botanical Reminiscences of George
R. Cooley, 1896-1986, 471-478
Sonchus oleraceus, 121
Southeastern United States, The Buxaceae
in the, 241-257
Southeastern United States, The Genera of
a (Cruciferae; Brassicaceae) in the,
185-240
Southeastern United States, The Genera of
Cinchonoideae (Rubiaceae) in the, 137—
3
Southeastern United ie The Genera of
Cyperaceae in the, 361-445
Southeastern United States, The Genera of
Pontederiaceae in the, 35-71
Southeastern United States, The Zanni-
Spigelia anthelmintha, 124
Spike-rush, 384
Spinacea, 466, 468
ene oe 118
18
Sporobolus indicus 117
Stachytarpheta j eee 10, 131
Status of the Nam er sake Solan-
der (H ENERO —34]
Stem Woods of Som toes of the
Mimosoideae cee A Com-
parative Study of Root and, 349-355
Stenophyllus, 393
Sterculiaceae, 130
Stigmaphyllon diversifolium, 110, 125
— -periplocifolium, 125
rumpfia maritima, 129
aes 242
Stylocerataceae, 242
502
Stylosanthes hamata, 124
Suckleya, 466, 468
Suriana maritima, 109, 130
Sweet alyssum, 20
Swietenia mahagoni, 108, 125
Synandra, |, 4-6, 15, 28
= hispidula. 12-14, 16, 19-21, 33
8
Syringodium filiforme, 116
Tabebuia heterophylla, 119
— pallida, 108, 11
Tabernaemontana divaricata, 119
286
, 130
Tamarindus ndicg. 108, 124
Tamarix chinensis, 130
Taonabo, 334
— stuebelii, 333, 334
Taxaceae, 269-272, 275, 278-280, 283,
286, 288, 291-293
Taxodiaceae, 269-271, 278, 280, 283, 284,
286, 288-290, 293
Taxodium, 278, 284, 286
Taxonomic and Nomenclatural Notes on
the Genus Mimosa (Leguminosae), 309—
Taxonomic Studies in Freziera (Theaceae),
s on Reproductive Biology
323-334
Taxus, 272, 275, 280, 288, 291
goin leucoxylon, 119
— stans, 119
pen cinerea, 124
Terminalia catappa, 121
Ternstroemia, 334
— stuebelii, 333, 334
Tetraclea, 5
Tec 284, 287
Tetramicra canaliculata, 118
Teucrium, 11,
— canadense, on 14, 19-21, 33
— chamaedrys, 12, 14, 20, 21, 33
Thalassia testudinum, 117
Theaceae: Taxonomic Studies in Freziera,
with Notes on Reproductive Biology,
323-334
Theaceae, 323
Theleophyton, 466, 468
Theophrastaceae, 130
Thespesia populnea, 109, 125
Three-square,
Thrinax morrisii, 110-113, 118
JOURNAL OF THE ARNOLD ARBORETUM
[VOL. 68
Thuja, 284
Thujopsis, 2 4, 287
Thunbergia fragrans, 118
Thysanocarpus, 191
Tiliaceae, 130
Tillandsia recurvata, 116
— usneoides, 116
— utriculata, 116
6
Tournefortia gnaphalodes, 119
— volubilis, 110, 120
cea pallida, 116
Triplopetalum, 196
Tsuga, 289
gin GorDON C. The Genera of Cy-
eraceae in . Southeastern United
es 361-445
Turnera oe 131
Turneraceae
Twig-rush, 419
Ullmannia, 278
Ulmaceae, 131
Umbelliferae, 131
Umbrella-sedge, 396
Unusual Pollen Dimorphism in Rondele-
tia anguillensis (Rubiaceae), 133-136
Urechites lutea, 110, 119
— suberecta, 119
Urticaceae, 131
Vaginaria, 382
Verbena hastata, 10
eno
Verbenacene’ A New Species of Lantana
from Dominica, Lesser Antilles, 343-348
1987]
Verbenaceae, 3, 4, 9-11,
34
— subfam. Verbenoideae, 10
— — tribe @ierodendicae. 11
— tribe Viticeae, 11
Vernonia albicaulis, 12]
_ short, 224
Jil
Vleisia ae hemoniaia: 260
Voltziaceae, 278
Walchiostrobus, 278
Walkomiella, 293
Waltheria americana, 130
Water hyacinth, 49
Water-hyacinth Family, 35
WEITZMAN, “ANNA L. Taxonomic Studies
in Freziera (Theaceae), with Notes on
Reproductive Biology, 323-334
INDEX
15, 25, 26, 131,
503
Whitlow grass, 211
Widdringtonia, 284, 287, 292, 293
Wild-gentian,
Woop, C. E., Jr., and R. B. CHANNELL.
The Buxaceae in the Southeastern United
States, 241-257
Woods of Some Members of the Mimo-
soideae (Leguminosae), A Comparative
Study of Root and Stem, 349-355
Wool-grass, 373
Xanthium strumarium, 121
Xanthosoma sagittatifolium, 116
Xerococcus, 181
Yucca guatemalensis, 115
gies ay 260, 261, 264-268
andina, 264
— major, 264
— palustris, 260, 264, 265
Zannichelliaceae, 259-268
Zanthoxylum flavum, 110, 129
— punctatum, 110, 129
— spinifex, 110, 129
Zea mays, 117
Zephyranthes candida, 115
Zhumeria, 23
Zingiberaceae, 40
Zinnia multiflora, 121
, 468
Soe 131
JOURNAL oF tHe
ARNOLD ARBORETUM
HARVARD UNIVERSITY VOLUME 68 1987
Dates of Issue
No. | (pp. 1-136) issued 6 January 1987.
No. 2 (pp. 137-268) issued 9 April 1987.
No. 3 (pp. 269-359) issued 8 July 1987.
No. 4 (pp. 361-503) issued 9 October 1987.
Contents of Volume 68
Phylogenetic Implications of Leaf Anatomy in Subtribe Melitti-
dinae (Labiatae) and Related Taxa.
Mones S. ABu-ASAB AND PHILIP D. CANTINO ................
The Genera of Pontederiaceae in the Southeastern United States.
THOMAS J. ROSATTI ... 0.000000. ccc cee een ene eees
Reproductive Structure of Lithocarpus Sensu Lato (Fagaceae): Cy-
mules and Fruits.
RoBERT B. KAUL
Contributions to a Flora of Anguilla and Adjacent Islets.
RICHARD A. HOWARD AND ELIZABETH A. KELLOGG ...........
Unusual Pollen Dimorphism in Rondeletia anguillensis (Rubi-
aceae).
ELIZABETH A. KELLOGG AND RICHARD A. HOWARD ...........
The Genera of Cinchonoideae (Rubiaceae) in the Southeastern United
States.
GEORGE Ki ROGERS o5:2u doe ueeseeunvdanuscuceeeaweswuse Kies
The Genera of Alysseae (Cruciferae; Brassicaceae) in the South-
eastern United States.
IHSAN A. AL-SHEHBAZ .... 00000000 ee eee
The Buxaceae in the Southeastern United States.
R. B. CHANNELL AND C. E. Woop, JR.. 0.000000 000000 eee
The Zannichelliaceae in the Southeastern United States.
Rospert B. HAYNES AND LAuRITz B. HOLM-NIELSEN ..........
A Cladistic Analysis of Conifers: Preliminary Results.
UEEPREN et ART, pi van hv owe ei vo ead seat eee ee ees
Taxonomic and Nomenclatural Notes on the Genus Mimosa (Le-
guminosae).
ROSAURA’GRETHER » ¢o0005 b400 2244060 veiw bbeeansandowered ees
Taxonomic Studies in Freziera (Theaceae), with Notes on Repro-
ductive Biology.
ANNA L, WEITZMAN ... 0.0000 ete eee
Status of the Name Aesculus flava Solander (Hippocastanaceae).
FREDERICK G. MEYER AND JAMES W. HARDIN ............-.-.
A New Species of Lantana (Verbenaceae) from Dominica, Lesser
Antilles.
ROGER W. SANDERS .......0.00000 0000 cece eterna ee
73-104
105-131
133-136
137-183
185-240
241-257
259-268
269-307
309-322
323-334
335-341
343-348
A Comparative Study of Root and Stem Woods of Some Members
of the Mimosoideae (Leguminosae).
K. RANJANI AND K. V. KRISHNAMURTHY ....................
Armoracia lacustris (Brassicaceae), the Correct Name for the North
American Lake Cre
IHSAN A. AL-SHEHBAZ AND VERNON BATES.................-.
The Genera of Cyperaceae in the Southeastern United States.
TOR DON (UCR eto paces thea he eerie otes Coes
A New Species of Pinus from Mexico and Central America.
Jes ER RNS Jaa d poesia ge pre deae eae eae opal ee ee ets
Archiatriplex, a New Chenopodiaceous Genus from China.
BCI A go a ore ed ep ay eerie etd e evanianeaan a Ae
Some Botanical Reminiscences of George R. Cooley, 1896-1986.
RICHARD A. HOWARD .......00....0.00000 0 ccc cece eee eeee
349-355
357-359
361-445
447-459
461-469
471-478
479-503
Journal of the Arnold Arboretum October, 1987
CONTENTS OF VOLUME 68, NUMBER 4
The Genera of Cyperaceae in the Southeastern United States.
CSORDON Ca FUER: aint nuh id Gide de eee eetaees act ddes 361-445
A New Species of Pinus from Mexico and Central America.
PS CO S50 "eb a Re ee Re aeieena at os 447-459
Archiatriplex, a New Chenopodiaceous Genus from China.
GE-LIN CHU ........... oT ee eee Oe ee nee eee ey eee 461-469
Some Botanical Reminiscences of George R. Cooley, 1896-1986.
RICHARD A. HOWARD: 64.4 ui5¢id60d26o0)seunbsaureraceeu's 471-478
Tie ea gracias asa ek es ee es bee eee 479-503
Volume 68, Number 3, including pages 269-359, was issued July 8, 1987.