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JOURNAL oF tre 
ARNOLD ARBORETUM 


HARVARD UNIVERSITY VOLUME 68 NUMBER 1 


ISSN 0004-2625 


Journal of the Arnold Arboretum 


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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 \ 


I 
+ 
I 
! 
! 
| 


I 
+ 
1 
1 
1 
i 


I 
tetteetteer+eeees 
i} 
i} 
I 
I 


look smeoinsoibosdnoowns-dne®) 


Toc ats 


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. 


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1987] ABU-ASAB & CANTINO, LEAF ANATOMY 29 


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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 
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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). 
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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 
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cological genetics of breakdown in tristyly. Pp. 267-275 in J. Haeck & J. W. 
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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- 
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Barton, L. V., & J. E. Horcuxiss. Germination of seeds of Eichhornia crassipes Solms. 
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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- 
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Borescu, K. Die Gestalt der Blattstiele ae Eichhornia crassipes (Mart.) Solms in ihrer 
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Britton, N. L. Piaropus azureus. Addisonia 2: 67, 68. pl. 74. 1917. [Color plate.] 

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56 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68 


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Curtiss, A. H. The water hyacinth in Florida. Plant World 3: 38-40, 1900. [Recounts 
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Francois, J. Observations sur l’hétérostylie chez Eichhornia . (Mart.) Solms. 
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. Habenaria repens and Piaropus crassipes in Leon County, Ace ee ie 

267-270. 1916. [Piaropus = Eichhornia; introduction thought t 


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Jounson, A. M. The mid-styled form of Piaropus paniculatus. Bull. Torrey Bot. Club 
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. 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). 
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maries in Spanish and French. ) vii + 174 pp. Washington, D. C. 1976. [Numerous 
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18: 447-472. 1948. [Distribution, nature and extent of damage, morphological/ 
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seenaney vegetative and sexual reproduction (including pollination and seed 
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Pettet, A. Seedlings of Eichhornia crassipes: a possible complication to control mea- 
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PIETERSE, A. The water hyacinth (Eichhornia crassipes),; a review. fae Trop. 
Agric. res ae 42. 1978. [666 citations covering biology, control, and u 

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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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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- 
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tribes, and genera with alkaloids.] 

McVauGu, R. The vegetation of the gramuc ot ocks ore southeastern United States. 
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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 
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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 


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JANKUN, A. /n: ar SKALINSKA & E. PoGAn, Further studies in chromosome numbers 
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JANOoT, M.-M., & J. LEMEN. Sur les alcaloides de Lunaria biennis Mnch. (Cruciféres). 
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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 
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Miwa, T. K. Gas chromatograms of synthetic liquid waxes prepared from seed triglyc- 
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& I. A. Worf. Fatty acids, fatty alcohols, wax esters, and methyl esters from 

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3 


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fatty acids.] 

OLESEN ee P. Amino acids and y-glutamyl derivatives in seeds of Lunaria annua 
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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 
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—. Regeneration, vernalization and flowering in Lunaria annua L. in vivo and in 
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ventitious root formation in oe ae segments of Lunaria annua L. 
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POLATSCHEK, A. Cytotaxonomische Beitrage zur Flora der Ostalpenlinder, I. Osterr. 
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1987] AL-SHEHBAZ, ALYSSEAE 195 


Uxricn, R. Observations biométriques sur la croissance des fruits de lunaire (Lunaria 
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. K. Hicazy. The juvenile see for flowering in Lunaria biennis. Proc. 
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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- 
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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. 
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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. 
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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. 
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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. 
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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 
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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. 
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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. 


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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 
» 


Soy 
COSI LE 


x) 


nee 


US 
y 
Sty 


He 


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. 
f- 8, oie 
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. } 
a ca tye a i tong ne due ioe \ 
. AG ; ‘m . ; i 
a ees 4 to mins ee es a — fe owt wes et : ! 
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4 , = \ 
rs Aa cere Cac, pebpr dl, Foie 38 
(Sabon cals warrenty 
ACG ees. ee ees Los ale. [Care on 


Cy : 
Cees! Fass ; Koy , bi sia rte pty fever 
Corte Pe de gr poly. eeeee ee ; 
(On foer. he, ea P isaewe Oe Le a. a) : 


+ Ly 


ae tnvere ay F hal yy hae ALC Co na TR > 
Ea plore 4 faa apt ; falwhe FR / 
. a 


ae e« for from o> anehs he. , fe ates 


; “ 
’ SY, 
wiceya , a L wee aJ ; coca 


Pe - - 
, Z, i ey S35 
a7. Pan ian, , 7 ees Moe 7 y the on hc, 
i Ae \ 


U7 len ~ aaa Vazecsnae yee, Ce8, 


oe imclr nah, ~ 
fhe - ; iae 
Nie eo aoe ou yurey 
= ae . 
dh ey ee a 


Dt. Korey nwleo Alt, Lore 


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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national Code (all editions through 1972) should be followed. When possible, 
reference should be made to past issues of the Journal for form. Titles should 
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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 


Journal of the Arnold Arboretum 


The Journal of the Arnold Arboretum (ISSN 0004-2625) is published quarterly in 
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EDITORIAL COMMITTEE 

S. A. Spongberg, Editor 

E. B. Schmidt, Managing Editor 
P. S. Ashton 

K. S. Bawa 

P. F. Stevens 

C. E. Wood, Jr. 


Printed at Allen Press, Inc., Lawrence, Kansas 


COVER: The stylized design appearing on the Journal and the offprints was drawn 
by Karen Stoutsenberger. 


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 
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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. 


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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. 


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20Carex sect. PALUDOSAE G. Don in Loudon, Hort. Brit. 367. 1830, non (Fries) Christ (1884). TyPE 
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438 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68 


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440 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68 


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: 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- 
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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 

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& 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 
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Cytology of Carex purpurifera Mack. (Cyperaceae). Rhodora 88: 141-147. 1986. 
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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 
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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 
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Moore, R. J., & J. A. CALDER. Some chromosom e numbers of Carex species of Canada 


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Murray, D. F. Taxonomy of Carex sect. Atratae (Cyperaceae) in the southern Rocky 
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NANNFELDT, J. A. The species of Anthracoidea (Ustilaginales) on Carex subgen. Vignea 
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NeLMES, E. Facts and speculation on phylogeny in the tribe Cariceae of the Cyperaceae. 
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Nos e, J. " oe flora of the British Isles: Carex arenaria L. Jour. Ecol. 70: 867- 
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ey ie Bee, & J. L. Harper. The population biology of plants with clonal 

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NovozHiLova, N. N. Flowering and pollination of Kohresia of eastern Pamirs. Nauk 
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Ou, Y.C. Taxonomic study of epidermal patterns on some American species [of] Carex 

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So AUER, J., Ee WENHOVEN. Néhrstoffokologie von Molinia coer ulea und 

Ca tifor uf baumfreien yr des Alpenvorlandes. (English sum- 
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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 
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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.) 
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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 
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RosBerRTSON, A. Variations in Carex (sect. Stellulatae Kunth) in Newfoundland. (Ab- 
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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 
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St. Jonn, H. A new Carex (Cyperaceae) of the section Stel/ulatae. Hawaiian plant 
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SAVILE, D. B. O., & J. A. en Phylogeny of — in the light of parasitism by the 
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ScHMID, B. W. Karyology and hybridization in ~ Carex flava complex in Switzerland. 
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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. 
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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. 
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SHEPHERD, G. J. Experimental arene in the genus Carex section Vesicariae. Unpubl. 
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The use of anatomical carters in the infrageneric classification of Carex 
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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- 
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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. 
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(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 
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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 
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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- 
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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- 

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Wepsser, J. M., & P. W. BALL. The taxonomy of the Carex rosea group (section Phaes- 
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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 
a0 
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 
oO 


_ 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.