Group Title: taxonomic revision of the genus Chamaesyce (Euphorbiaceae) in the Caribbean /
Title: A Taxonomic revision of the genus Chamaesyce (Euphorbiaceae) in the Caribbean
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 Material Information
Title: A Taxonomic revision of the genus Chamaesyce (Euphorbiaceae) in the Caribbean
Physical Description: x, 244 leaves : ill. ; 28 cm.
Language: English
Creator: Burch, Derek George, 1933-
Publication Date: 1965
Copyright Date: 1965
 Subjects
Subject: Botany -- Caribbean Area   ( lcsh )
Chamaesyce   ( lcsh )
Botany thesis Ph. D
Dissertations, Academic -- Botany -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis (Ph. D.)--University of Florida, 1965.
Bibliography: Includes bibliographical references (leaves 239-242).
Additional Physical Form: Also available on World Wide Web
Statement of Responsibility: by Derek George Burch.
General Note: Typescript.
General Note: Vita.
 Record Information
Bibliographic ID: UF00097892
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000439291
notis - ACJ9471
oclc - 21194359

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A TAXONOMIC REVISION OF THE GENUS

CHAMAESYCE (EUPHORBIACEAE) IN

THE CARIBBEAN


















By
DEREK GEORGE BURCH









A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARIIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY










UNIVERSITY OF FLORIDA
August, 1965













ACKNOWLEDGEMENTS


I should like to express my thanks to Dr. Daniel B. Ward

for his advice and helpful criticism in his capacity as Chairman

of the supervisory committee for this study. I should also like to

thank the other members of the committee, Dr. W. 0. Ash and Dr.

D. A. Roberts, for reading and evaluating the manuscript.

I am grateful to the Curators and staff of the herbaria listed

in the introduction for lending their material, including type-

specimens, for use in this study, and my thanks are also due to

Mr. W. T. Steam of the British Museum (Natural History) and Dr.

R. K. Rechinger of the Naturhistoriskmuseum, Vienna, for searching

their collections for additional specimens of interest.

The Department of Botany and the Graduate School of the Uni-

versity of Florida contributed to the expenses of a visit to several

important European herbaria, which added greatly to the coverage of

this work.

Finally, my thanks are due to the anonymous assistants for the

final stages of the preparation of the dissertation, and particularly

to my wife Nancy, who rounded out two years of "herbarium-widowhood"

by finding out just how I had been spending my nights during that time.

All photographic work in the dissertation was done by Mr. Jerry Uelsmann,

and I am grateful to him both for the long hours of work that went into

the plates, and for their leavening effect in the mass of text.













TABLE OF CONTENTS


ACKNOWLEDGEMENTS

LIST OF MAPS

LIST OF PLATES


INTRODUCTION

GENERAL DISCUSSION

I. HISTORY

II. THE SEGREGATION OF THE GENUS CHAMAESYCE

II MORPHOLOGY

SYSTEMATIC TREATMENT

Chamaesyce

Key to the species of Chamaesyce in the Caribbean

1. Chamaesyce vaginulata

2. Chamaesyce articulata

3. Chamaesyce buxifolia

4. Chamaesyce lecheoldes

5. Chamaesyce cayensis

6. Chamaesyce porteriana

7. Chamaesyce myrtillifolia

8. Chamaesyce nutans

9. Chamaesyce hypericifolia


Page

ii

vi

viii







TABLE OF CONTENTS, Continued

SYSTEMATIC TREATMENT, Continued Page

Chamaesyce

10. Chamaesyce hyssopifolia 76.

11. Chamaesyce lasiocarpa 86.

12. Chamaesyce parciflora 88.

13. Chamaesyce brittonii 90.

14. Chamaesyce cowellii 93.

15. Chamaesyce blodqettii 97.

16. Chamaesyce serpens 107.

17. Chamaesyce centunculoides 111.

18. Chamaesyce torralbasii 117.

19. Chamaesyce camaguayensis 121.

20. Chamaesyce cordifolia 124.

21. Chamaesyce polygonifolia 128.

22. Chamaesyce ammannioides 130.

23. Chamaesyce cumulicola 134.

24. Chamaesyce prostrata 138.

25. Chamaesyce mendezii 144.

26. Chamaesyce hepatica 148.

27. Chamaesyce helwiqii 150.

28. Chamaesyce hirta 152.

29. Chamaesyce opthalmica 160.

30. Chamaesyce berteriana 165.

31. Chamaesyce maculata 168.






TABLE OF CONTENTS, Continued


SYSTEMATIC TREATMENT, Continued Page

Chamaesyce

32. Chamaesyce thymifolia 175.

33. Chamaesyce adenoptera 180.

34. Chamaesyce conferta 190.

35. Chamaesyce garberi 193.

36. Chamaesyce multinodis 203.

37. Chamaesyce minutula 205.

38. Chamaesyce turpinii 207.

39. Chamaesyce paredonensis 210.

40. Chamaesyce pinetorum 213.

41. Chamaesyce deltoidea 216.

42. Chamaesyce tumistyla 224.

43. Chamaesyce leonardii 226.

DOUBTFUL SPECIES 229.

PHYLOGENY 231.

BIBLIOGRAPHY 239.

APPENDIX 243.

BIOGRAPHICAL SKETCH 244.












LIST OF MAPS


Map 1.

Map 2.

Map 3.

Map 4.

Map 5.

Map 6.

Map 7.

Map 8.

Map 9.

Map 10.

Map II.

Map 12.

Map 13.

Map 14.

Map 15.


Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution


Distribution of C.

Distribution of C.

Distribution of C.

Distribution of C.

Distribution of C.

Distribution of C.

Distribution of C.


of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.

of C.


vaqinulata

buxifol ia

lecheoides var. wilsonii

lecheoides var. lecheoides

lecheoides var. exumensis

cayensis

porteriana var. keyensis

porteriana var. porteriana

porteriana var. scoparia

articulata and C. myrtillifolia

nutans

hypericifolia

hyssopifolia

lasiocarpa

parciflora

brittonii

cowel 1 ii

blodgettii

serpens

centunculoides

torralbasii

camaquayensis










LIST OF MAPS, Continued


Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution

Distribution


of C. cordifolia

of C. polygonifolia

of C. ammannioides

of C. cumulicola

of C. prostrata

of C. mendezii

of C. hirta

of C. opthalmica

of C. berteriana

of C. maculata

of C. helwiqii and C. paredonensis

of C. thymifolia

of C. adenoptera ssp. adenoptera

of C. adenoptera ssp. perqamena

of C. adenoptera ssp. qundlachii

of C. conferta

of C. garberi

of C. turpinii

of C. pinetorum

of C. deltoidea ssp. serpyllum

of C. deltoidea ssp. deltoidea
var. deltoidea
of C. deltoidea ssp. deltoidea
var. adhaerens


Page

127.

127.

135.

135.

143.

143.

159.

159.

167.

167.

179.

179.

188.

188.

189.

189.

202.

202.

215.

215.












LIST OF PLATES


Page

Plate 1. CHAMAESYCE VAGINULATA (Griseb.) Millsp. 31.

Plate 2. CHAMAESYCE ARTICULATA (Aubl.) Britton 35.

Plate 3. CHAMAESYCE BUXIFOLIA (Lam.) Small 38.

Plate 4. C. LECHEOIDES (Millsp.) Millsp. var. WILSONII
(Millsp.) ined. 46.

Plate 5. CHAMAESYCE LECHEOIDES (Millsp.) Millsp. var.
LECHEOIDES 47.

Plate 6. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS
(Millsp.) ined. 48.

Plate 7. CHAMAESYCE CAYENSIS (Millsp.) Millsp. 53.

Plate 8. CHAMAESYCE PORTERIANA Small var. KEYENSIS (Small)
ined. 58.

Plate 9. CHAMAESYCE PORTERIANA Small var. PORTERIANA 59.

Plate 10. CHAMAESYCE PORTERIANA Small var. SCOPARIA (Small)
ined. 60.

Plate 11. CHAMAESYCE MYRTILLIFOLIA (L.) Millsp. 64.

Plate 12. CHAMAESYCE NUTANS (Lag.) Small 67.

Plate 13. CHAMAESYCE HYPERICIFOLIA (L.) Millsp. 73.

Plate 14. CHAMAESYCE HYSSOPIFOLIA (L.) Small 79.

Plate 15. CHAMAESYCE HYSSOPIFOLIA (L.) Small 80.

Plate 16. CHAMAESYCE HYSSOPIFOLIA (L.) Small 81.

Plate 17. CHAMAESYCE BRITTONII (Millsp.) Millsp. 92.

Plate 18. CHAMAESYCE COWELLII Millsp; in Britton 95.

Plate 19. CHAMAESYCE BLODGETTII (Engelm. ex Hitchc.) Small 99.










LIST OF PLATES, Continued


Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate

Plate


CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE

CHAMAESYCE


Plate 39. CHAMAESYCE


BLODGETTII (Engelm. ex Hitchc.) Small

BLODGETTII (Engelm. ex Hitchc.) Small

SERPENS (HBK.) Small

CENTUNCULOIDES (HBK.) Millsp.

CENTUNCULOIDES (HBK.) Millsp.

TORRALBASII (Urb.) Millsp.

CAMAGUAYENSIS Millsp.

CORDIFOLIA (Ell.) Small

POLYGONIFOLIA (L.) Small

AMMANNIOIDES (HBK.) Small

CUMULICOLA Small

PROSTRATA (Ait.) Small

MENDEZII (Boiss.) Millsp.

HEPATICA (Urb. & Ekm.) ined.

HELWIGII (Urb. & Ekm.) ined.

HIRTA (L.) Millsp.

OPTHALMICA (Pers.) ined.

MACULATA (L.) Small

MACULATA (L.) Small

THYMIFOLIA (L.) Millsp.


Plate 40. C. ADENOPTERA (Bertol.) Small var. PERGAMENA
(Small) ined.

Plate 41. C. ADENOPTERA (Bertol.) Small var. PERGAMENA
(Small) ined.









LIST OF PLATES, Continued

Page

Plate 42. C. ADENOPTERA (Bertol.) Small var. GUNDLACHII
(Urban) ined. 187.

Plate 43. CHAMAESYCE CONFERTA Small 192.

Plate 44. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 196.

Plate 45. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 197.

Plate 46. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 198.

Plate 47. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 199.

Plate 48. CHAMAESYCE MULTINODIS (Urb.) Millsp. 204.

Plate 49. CHAMAESYCE TURPINII (Boiss.) Millsp. 209.

Plate 50. CHAMAESYCE PAREDONENSIS Millsp. 212.

Plate 51. CHAMAESYCE PINETORUM Small 214.

Plate 52. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small
ssp. SERPYLLUM (Small) ined. 219.

Plate 53. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small
ssp. DELTOIDEA var. DELTOIDEA 221.

Plate 54. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small
ssp. DELTOIDEA var. ADHAERENS (Small) ined. 222.

Plate 55. EVOLUTION OF BRANCHING PATTERN IN CHAMAESYCE 234.













INTRODUCTION


The genus Chamaesyce (Euphorbiaceae) is represented on all

continents, and, while many species are widespread in temperate areas,

the genus reaches its greatest diversity in tropical and subtropical

regions.

In 1941 Wheeler revised the group in North America exclusive of

Mexico and southern Florida, treating it as a subgenus of the Linnaean

genus Euphorbia. He found areas with high concentrations of species

centered in Arizona and in Texas, and suggested that a similar concen-

tration might exist in south Florida involving species with a Caribbean

affinity. The most recent treatment of the genus in Florida by Small

(1933) indicates the same thing, since eighteen species confined to

Florida are included in the twenty-nine that he lists for the state.

Many of these are from the pinelands around Miami or on the lower Keys,

areas with very distinctive plant associations, but not generally rich

in endemic species. The plants of south Florida are known to have strong

floristic ties both with Cuba and the Yucatan peninsula, and with the

Bahama Islands, and it was clear that the Florida species could not be

dealt with effectively without considering those of the surrounding area.

The Caribbean as a whole here considered to include southern Florida

as well as the Bahamas and the Greater and Lesser Antilles, but excluding

Trinidad and the islands of the Dutch West Indies off the coast of

Venezuela does, however, make up a contained unit floristically, and

is a convenient area for study. It has the added advantage that no









synthesis of the genus for this area has been attempted since Boissier's

treatment of Euphorbia sensu lato on a worldwide basis in 1862, although

certain regions have been extensively collected since that time, and the

genus revised in these localities.

Regional floras with relevant material include The Bahama Flora

by Britton and Millspaugh (1920), which draws on Millspaugh's strong

interest in Chamaesyce, Fawcett and Rendle's Flora of Jamaica (1920),

Scientific Survey of Porto Rico and the Virgin Islands by Britton and

Wilson (1924), and Flora de Cuba by Alain (1953). There is also material

relating to Hispaniola and to Cuba in Urban's Symbolae Antillanae (1898-

1928), and in the reports of Ekman's collecting trips in Haiti (Urban

and Ekman, 1929).

The consequence of this publication of works covering only small

geographic areas has been the description of the same plant under

different names from different parts of its range. It is fortunate

that much of the type material for these names is still accessible, and

it has proved possible in the course of the study to reduce many of them

to synonymy. Material from twenty-two herbaria has been examined, and

the large number of specimens available of many species has given

valuable indications of the extent of morphological variation to be

expected and taken into account when considering the validity of names

about which there is doubt.

Some 4500 sheets have been studied and additional material, as

well as valuable type specimens, examined during visits to other insti-

tutions. The herbaria from which material has been seen are listed below,

together with the abbreviations (most from Lanjouw and Stafleu, 1964),










by which they will be referred to in the text.


A Arnold Arboretum, Cambridge, Massachusetts

BM British Museum (Natural History), London

BRASS Herbarium of Archbold Biological Station, Lake Placid, Florida

BUS University of Miami, Coral Gables, Florida

DUKE Duke University, Durham, North Carolina

E Royal Botanic Garden, Edinburgh, Great Britain

F Chicago Natural History Museum, Chicago, Illinois

FLAS University of Florida, Gainesville, Florida

FSU Florida State University, Tallahassee, Florida

GH Gray Herbarium of Harvard University, Cambridge, Massachusetts

IJ Institute of Jamaica, Kingston, Jamaica

K Royal Botanic Gardens, Kew, Surrey, Great Britain

LINN Linnaean Society Herbarium, London

MO Missouri Botanical Garden, St. Louis, Missouri

NCU University of North Carolina, Chapel Hill, North Carolina

NSC North Carolina State University, Raleigh, North Carolina

NY New York Botanical Garden, New York

PUR Purdue University, Lafayette, Indiana

RUS Louisiana Technical College, Ruston, Louisiana

UCWI University of the West Indies, St. Andrews, Jamaica

US U. S. National Museum, Washington, D. C.

USF University of South Florida, Tampa, Florida












GENERAL DISCUSSION


I History


Many taxonomists have treated Chamaesyce as no more than a

subdivision of the large, diverse genus Euphorbia L. To a certain

extent the choice of which of these generic names to use is an

academic matter, but, insofar as the schemes of biological classifi-

cation are intended to have a natural basis, the collection of species

into a genus should reflect a close degree of common ancestry, and

the thesis of workers who would split Euphorbia into several genera

is that such a division represents this close relationship better than

would the maintenance of the broad genus.

The genus Euphorbia was described by Linnaeus in Genera Plantarum

ed. 5 (1754), and was delimited on the basis of its floral characters.

Reference was made to the earlier names Euphorbium proposed by D'Isnard

in 1720 for much the same group, and to Tithymalus and Tithymaloides,

both of which were used by Tournefort (1700), one of the first botanists

to make such formal definitions of groups rather than simply listing

species.

The "sexual system" of classification followed by Linnaeus

divided plants into Classes according to the number of stamens present

in the flower, with these Classes then divided into Orders, usually

formed by reference to the number of styles. The genus Euphorbia was

placed in the third Order of the eleventh Class, DODECANDRIA TRIGYNIA,









since Linnaeus had found that twelve or more stamens were usually present,

and that there were three styles. The description in Genera Plantarum

stresses the characteristic calyx of the genus, with its segments joined

to form a four-or five-toothed cup and petals set on the margin alter-

nating with the teeth, and also the tricoccate capsule with a single

seed in each locule. Some of the variety to be found in the shape of

the petals and in the form of the stem is also mentioned. Linnaeus was

familiar with some fifty-six species at this time, and his descriptions

in Species Plantarum (1753) indicate that he felt the characteristic

flower to be a strong enough factor to hold together in one genus such

diverse morphological forms as the cactus-like E. mammillaris, the small,

prostrate E. maculata, and the erect E. lathyrus with its umbellate

branches and crescent-shaped petals.

The genus, characterized by the flower type, was accepted by most

authors of the period, although Wheeler (1943) points out that as early

as 1754 Miller had proposed that part of the genus, homogeneous for cer-

tain morphological characters, be split off under another name.

In 1818 Robert Brown, an Edinburgh surgeon and keen botanist,

published the results of anatomical work which showed that the flower

which Linnaeus described was, in fact, an inflorescence. The tricoccate

ovary was actually a terminal female flower with the perianth parts re-

duced to a vestigial calyx, and below this in the inflorescence were

groups of male flowers, each reduced to a single stamen. The cup-like

calyx which Linnaeus described was an involucre of fused bracts, while

the petals were glands and associated appendages set on the upper rim

of the involucre. The name cyathium was given to this type of inflorescence.








This work of Brown weakened the case for maintaining the diverse

forms under one genus because it is inconsistent with usage in the rest

of the plant kingdom, and because Linnaeus would presumably not have

defined a genus on this basis of a shared inflorescence type. The

parallel has been drawn with the situation in the Compositae, which

are defined as a family by the capitate inflorescence and by reduction

in some flower parts, and which would form a genus of some 20,000 species

if these were considered generic rather than family characters. This

obviously represents neither a realistic appraisal of the close degree

of common ancestry which is usually sought for a genus, nor an efficient

means of data retrieval which is one of the tacit aims in setting up

such a system of classification.

In spite of this or perhaps because Brown's results were not

universally accepted at first (Haber, 1925) many of the diverse species

described in the hundred years after the publication of Species Plantarum

were named using Euphorbia in its broad sense, and the stamp of approval

was given to the practice by Boissier in a series of publications cul-

minating in his monumental treatment of the family with J. Muller in

de Candolle's Prodromus (1862). The characters of the inflorescence

were still used to define the genus, but by this time the number of species

known had swollen to the point at which internal subdivision of the genus

was standard practice. Persoon (1806), Rafinesque (1817), and Reichen-

bach (1841) had all set up subgenera, while Haworth (1827) and Roeper

(1828) had divided the genus into sections. Boissier himself used sec-

tions and subsections, but was apparently dissatisfied with the bases for

his division, since he comments in his notes on the genus (1862) "The










characters of the sections are often trifling and open to exception

but better ones cannot be found" (translation by Croizat, 1936a). We

may assume from this that Boissier maintained the genus Euphorbia in

its widest sense not only because of the common inflorescence type, but

also because he could find no consistent characters on which to divide

it.

Many more species have been added to the genus since the time

of Boissier, until today Scholz (1964) estimates that it includes some

1600 entities.

The retention of the genus in its broad sense has been opposed

on a number of grounds, and attempts have been made from time to time

to justify the segregation of certain groups. Croizat (1936a) has

pointed out that the possession of a cyathium is not restricted to this

genus, but occurs in several others recognized by even the most con-

servative taxonomists. Among these Pax (who had written the account

of the family for Engler and Prantl's Pflanzenfamilien in 1890) is

quoted as saying in 1924 that the cyathium was the main character of

the Tribe Euphorbieae rather than the genus Euphorbia. Croizat also

suggests that the species in Euphorbia are more variable in their degree

of evolution than would be expected in a single genus, and that the

cyathium is not an anatomic constant, and concludes that the use of the

single character of the presence of a broadly similar inflorescence,

cutting across other indications of relationship as it does, is unsound

from a taxonomic point of view.

Evidence from new areas of taxonomic research confirms the lack

of homogeneity in the genus. Perry (1943) reports that a survey of








somatic chromosomes in the Euphorbiaceae showed a considerable degree

of uniformity in size, shape, and basic number within all genera except

Euphorbia, which seemed to include three distinct groups.

Studies on the electrophoresis of latex by Moyer (1934) gave

groupings within the genus corresponding roughly to the sections of

Boissier. The work on pollen morphology by Erdtman (1952) and Punt

(1962), however, gave no clear divisions, although some of the patterns

of exine surface showed differences through the genus. A difficulty

with interpreting these studies lies in the fact that no voucher speci-

mens are available, so that any conclusions drawn must be tempered by

the uncertainty as to the exact taxonomic entities represented by the

names used.

Haber (1925), while not favoring a division of the genus, found

that the sections described by Pax in Engler and Prantl's Pflanzen-

familien (1890) showed little affinity in terms of anatomy and morphology

of the cyathium.

For these reasons, together with the diversity of habit and form

in the group, it is hard to accept that Euphorbia fulfills the condition

of relatively close ancestry which is usually imposed when judging the

soundness of a genus.

It is unfortunate that no comprehensive evaluation has been made

since Boissier's work more than a century ago, since many of the proposals

for segregating groups which have seemed inconsistent with usage in the

rest of the unchanged genus, might well have been perfectly acceptable in

the light of an overall revision. One such segregation to which this

criticism of inconsistency might apply is the separation of Chamaesyce






9.


as a genus without considering the equally valid genera Tithymalus,

Agaloma and Poinsettia. It is beyond the scope of the present work

to examine all these genera, but Chamaesyce is such a discrete group

in the area under consideration, and so easily distinguishable from

other members of Euphorbia sensu lato, that it will be accepted here

as a distinct genus. A case will be presented for this division below,

but final judgment of its validity must wait until a revision of the

complete Tribe Euphorbieae of the Family Euphorbiaceae has been under-

taken.












II. The Seqregation of the Genus Chamaesyce


Chamaesyce was first proposed as a segregate genus by S. F. Gray

in 1821 with a single species which would appear automatically to become

the type species. Wheeler (1943), however, supports Millspaugh (1909)

in designating Euphorbia peplis L. as type since Gray was clearly describ-

ing this species under his name C. maritima and, in fact, gave E. peplis

in synonymy, thus making the later name illegitimate. The type specimen

of E. peplis L. in the Linnaean Herbarium becomes the type of C. maritime

S. F. Gray and of the genus Chamaesyce S. F. Gray.

Wheeler (1943) points out that the name had been validly pub-

lished as a subgenus, and suggested provisionally as a genus, by Rafinesque

in 1817, but such provisional publication is not acceptable. Gray was

justified in selecting this generic name even though Anisophyllum had

been proposed for the same group at this taxonomic level by Haworth in

1812, since that name was preoccupied by Jacquin's use of it in another

sense in 1763.

Gray took the name Chamaesyce from Dioscorides, a Greek herbalist

of the first century A.D., who compiled lists of plant names and uses

from medical works and from herb-gatherers and peasants (Gunther, 1959).

The word is a transliteration of two Greek components, chamae, of the

ground or low-growing, and Syce which was one of the names used for the

fig, and the association is probably due to the presence in members of

the genus Euphorbia of a milky latex similar to that found in fig trees.

The name was listed as a synonym for the genus Euphorbia by Wiman (1752)









with a similar comment that it was a type of low-growing fig (quasi

humilis ficus).

Gray's name was not widely adopted for a number of years after

publication, but the cohesion of the species which belong in the genus

is indicated by their treatment by subsequent authors. Roeper pub-

lished the group as Euphorbia section Anisophyllum in 1828, while

Rafinesque changed his mind after his publication of the subgenus

Chamaesyce in 1817 to treat them as a set of related genera in 1838.

They were put back into a single category as Euphorbia subgenus

Anisophyllum Caesalp. ex Reichenbach in 1841, treated as the genus

Anisophyllum by Klotzsch and Garcke (1859), and finally the definitive

work by Boissier in 1862 followed Roeper by using section Anisophyllum

for the group.

Croizat (1936a, 1936b, 1937) has examined these species which

have been combined so consistently by authorities on the family. They

share a number of morphological characters, which have been used by the

various authors in an attempt to delimit the group at the particular

taxonomic level which they favored. Boissier (1862), for example, used

leaf arrangement and shape, stipule shape, inflorescence type, number

and form of glands and the ecarunculate nature of the seed in his com-

parisons with the other twenty-six sections which he recognized, while

Wheeler (1941). considering only New World species, separated four sub-

genera on the basis of number and form of glands and their appendages,

and on leaf arrangement and stipule development. Boissier's dissatis-

faction with his separation has already been mentioned, and Wheeler is

equally pessimistic about his success in defining his divisions. There









is general agreement in the literature, however, about the characters

shared by this group of species. They all have opposite, petiolate

leaves with oblique bases and interpetiolar stipules. All have an

interesting distribution of chlorophyll-bearing tissue in the leaves,

where it surrounds the veinlets leaving clear areas between them, and

all have cyathia with four glands whose appendages, if present, are

petalloid, never lunate. The fifth gland is either obsolete or much

reduced and set in a sinus through which the capsule stalk often droops

at maturity, and, finally, the seeds are ecarunculate.

None of these characters is restricted to this group of species,

but Croizat (1936a, 1936b) has demonstrated that a particular type of

branching, in which the main axis aborts early and is replaced by one

or more secondary axes, is peculiar to the group, and serves as a

diagnostic feature. Here, then, is a discrete character of a fundamental

nature the presence or absence of a type of branching which gives

an absolute separation of a group of species from the heterogeneous

Euphorbia sensu lato. What is more, the species showing this type of

branching share many other morphological features, and have been accepted

as a cohesive group by most authorities on the family. On these grounds

their segregation as the distinct genus Chamaesyce appears well justified.

This separation has been adopted in the present work, and is

particularly appropriate in the New World where the species of Chamaesyce

are easily distinguishable from other members of the family on even a

casual examination. The usage is by no means novel in this area in that,

of the floras mentioned in the Introduction, only those by Urban and by

Fawcett and Rendle maintain Euphorbia sensu lato. The split has not been






13.



as widely adopted in other regions, however, and an assessment of its

value on a worldwide basis must await a more comprehensive study.













Ill. Morphology


The largest members of the genus Chamaesyce are found in the

Hawaiian Islands where some species have become arborescent. In the

Caribbean no such development is found, although one species, C.

vaqinulata (Griseb.) Millsp., has strong affinities with species of

the Galapagos Islands, and exhibits some of the peculiarities of

structure otherwise found only in endemics of that group. The remarks

in the following sections are intended to be limited to species grow-

ing under natural conditions in the Caribbean, which include annual and

perennial herbs of prostrate, ascending or erect habit, and shrubs and

subshrubs of open form reaching a height of one to six meters.

Roots

The root system is not extensive, and the laterals are slender

and branching. The main root in annual species is substantial and

little branched, and may become quite thick in plants living beyond

their normal season. Some of the perennials develop a short, thick

taproot, which develops buds at or below the surface of the ground

throughout the life of the plant, and often regardless of whether the

above-ground portions have died back or maintained their growth.

Stems

The main axis is interrupted by the abortion of the growing point

after the production of one pair of true leaves. Lateral branches

(secondary axes) develop from a point below the apex with no apparent

relation to the position of the pair of leaves (Croizat, 1936b). If

14..









only one secondary axis develops its thickening may obscure and eventually

conceal its origin, but in species in which a number of branches grow

(particularly those in which they lie prostrate) this apex is well marked.

On these secondary axes laterals (tertiary axes) may develop,

either throughout the length, or only near the lower or upper end. A

common form is the production of short, rather congested laterals from

the upper nodes on which most of the cyathia develop. Branching appears

to be alternate, with occasional production of other weaker branches at

the node, often becoming dichotomous near the apex, and may be at an

obtuse or acute angle with the stem. In the herbaceous species, stems

are wiry and brittle, or wider and of softer texture, often becoming

woody at the base. Rooting at the nodes is common in some species.

Prostrate species form tight or loose mats to 5 dm. diameter, the erect

herbaceous species may reach 6-8 dm.

Croizat (1936b) describes the growth of the secondary axes as

sympodial with each node terminated by a bud, a cyathium or a capitulum,

and the tertiary axes developed lateral to these. This is in agreement

with Roeper (1824) who explains the apparently normal lateral branching

as actually a reduction from the potential dichotomy which is expressed

at times at the tips of branches (Wheeler, 1941). The strongly alternate

branching which many species exhibit merely requires that the reduction

of the branches of the dichotomy alternates at the nodes. The same

pattern of branching may develop on tertiary axes, and rarely on those

of higher orders.

In the systematic part of this study the word "stem" will be used

to indicate the secondary axes, and lateralss" or "side branches" will









be used for tertiary and higher order axes. The aborted apex of the

stem, from which the secondary axes develop, will be called the crown

of the root for convenience in making comparisons with descriptions by

other authors.

Leaves

Leaves are opposite on the stem, and more or less two-ranked.

The blades are simple, membranous or sometimes coriaceous or sometimes

somewhat fleshy, and broad or rarely almost linear, 1 x I to 35 x 15 mm.

The base is inequilateral, usually markedly so, the margin entire or

variously toothed and plane or somewhat revolute, and the apex round

to acute or sometimes minutely apiculate. The veins are rarely promi-

nent, but in thin leaves the concentration of chlorophyll-bearing cells

around the conducting tissue with clear spaces between marks out the

reticulation of the veinlets.

Petioles are short, and the interpetiolar stipules may be

separate, or those on either side of the stem may be joined to the

corresponding one from the paired leaf. Their texture is usually mem-

branous, and the margins may be entire or variously toothed or parted.

In some species the pairs of leaflets droop at night in well-marked

nastic movement.

Cyathia

These are solitary and terminal, but often appear axillary when

the growth of a lateral continues the axis. They are often borne on

congested leafy laterals or in short-stalked lateral and terminal

dichasia, and sometimes, by reduction, in leafless dichasial glomerules.

Each cyathium is itself a compound inflorescence, but strongly










simulates a single flower. A single terminal female flower with, at

most, a vestigial calyx to represent the perianth, is surrounded by

groups of male flowers, each flower reduced to a single stamen and

interspersed by bracteoles. These are enclosed in a cup-like involucre

made up of five foliaceous structures, united at their margins but with

free tips. Between these, glands are borne on the rim of the cup,

supplied by a trace from each of the adjacent foliar structures (Haber,

1925) and often with a petalloid appendage, which may be as long as,

and up to four times as wide as, the subcircular or transversely ellip-

tic gland. The fifth gland is obsolete or reduced to a linear structure,

and the space between the two lobes which it would occupy becomes a

sinus into which the stalk of the capsule often droops at maturity. The

cyathia are short-peduncled or rarely sessile. The term lobe is used

here and in the systematic section to indicate the free portion of the

foliar structures which are fused to make up the involucre.

The glands appear moist under magnification, and may produce a

substance attractive to insects. A large number of ants are often

found on and around the plants, and may be the agent for cross-pollination.

Male Flower

An articulation between the pedicel and stamen indicates that

these monandrous structures are each a reduced flower rather than simply

floral parts. Each is subtended by a bracteole which may grow up between

the flowers. The staminate flowers form in five fascicles, arranged

opposite the lobes of the involucre, and in each fascicle the flowers

develop centrifugally with respect to the cyathium axis. In our species

five, or rarely fewer, to about thirty is the usual number of staminate









flowers to each cyathium. The number is relatively constant in a

given species. The stamens are exserted at maturity, carried on

elongated glabrous pedicels (Wheeler's term "androphore" is adopted

for this pedicel in the systematic section). Dehiscence of the

anthers is transverse, and the serial maturation of the flowers spans

the time during which the female flower is developing.

Female Flowers

The single terminal flower is pedicillate (Wheeler's term

"gynophore' will be used for this pedicel), with the perianth reduced

to a vestigial three-lobed calyx or a mere disc below the ovary. The

pistil has three united carpels, each with a single pendulous,

anatropous ovule. The three styles are joined at the base or rarely

free or connate for a greater distance, and are bifid for part of their

length. The gynophore is glabrous or variously pubescent, and in most

cases elongates with age and bends to lie in the sinus that marks the

position of the fifth gland.

Fruit

This is a trilocular capsule with one seed in each chamber,

usually roundly lobed or triangular but sometimes subspherical.

Dehiscence is by longitudinal splitting at the sutures, and to a

smaller extent at the angles. The base of the capsule is released

first, and the seed may be thrown a short distance from the flower by

the parting of the capsule wall.

Seed

The caruncle, which is so well developed in other species of

Eunhorbia sensu lato, is completely absent in Chamaesyce. The embryo









and endosperm are surrounded by a membrane and the testa, which may in

turn be covered by a persistent cellulose layer. This layer and the

testa may be patterned or sculpted.

The shape of the seed follows that of the trilocular capsule in

which it develops. In the systematic section the faces to the outside

of the capsule will be referred to as the dorsal faces, subtending the

dorsal angle, and the other two as the ventral faces subtending the

ventral angle. The seed is irregularly ovoid in longitudinal section

with the ventral side flattened. In cross section it is four-angled,

often obscurely so, but with the dorsal angle usually more acute and

pronounced than the ventral, and the lateral angles intermediate

between the two.

Vesture

The stems, leaves, cyathia, ovary and capsules may be glabrous

or variously long- or short-hairy. The hairs are all simple, and may

be single-or several-celled. In some species the surface of the leaves

and young stems may be entirely minutely white-papillose, appearing

almost granular under magnification.

Latex

All species have a milky latex to which various caustic properties

have been ascribed.

Morphological characters have proved to be very plastic within

many of the species examined. The habit of the herbaceous species

often varies widely, with occasional erect specimens in species which

are usually prostrate to decumbent, and with sprawling plants from species

which are normally completely upright to ascending. The number of stems









growing from the root is a more constant feature, with prostrate plants

more likely to have multiple stems than erect species, but the number

often changes with the change in habit in the case of erect specimens

of prostrate species.

The pubescence of the plant is usually reliable as a character

when contrasted with the absence of hairs (there are exceptions to this

among the south Florida species), but the degree of pubescence and even

the form of the hairs present is likely to vary over a wide range within

a species.

Leaf shape is variable, as might be expected, but in most cases

there is also a strong likelihood of toothed margins on occasional

leaves (or whole plants) of species which normally have entire leaves.

Cyathial characters in general show less plasticity, but the size

and form of appendages is rarely reliable, and vestiture may differ

widely from plant to plant or branch to branch. The same is true of

capsular characters, but the seed has proved to be stable enough that

characters relating to its morphology are often sufficient to key plants

out, and certainly to confirm doubtful identifications.













SYSTEMATIC TREATMENT


CHAMAESYCE S. F. Gray, Nat. Arr. Brit. PI., 2: 260. 1821.

Type: Chamaesyce maritima S. F. Gray. This is a superfluous name for

Euphorbia pepiis L., and the type specimen of this species in the

Linnaean Herbarium (sheet 630-18) is thus the type for C. maritima and

for the genus. Euphorbia subgenus Chamaesyce (S. F. Gray) House, Bull.

N. Y. State Mus. 254: 470. 1924.

Anisophyllum Haworth, Syn. PI. Succ., 159. 1812. Type:

Anisophyllum peplis (L.) Haw., designated by Wheeler, Rhodora 43: 110.

1941. Euphorbia section Anisophyllum (Haw.) Pax in Engler & Prantl,

Nat. Pflanzenfam., 3(5): 104. 1891.

Euphorbia subgenus Chamaesyce Raf., Amer. Monthly Mag. 2: 119.

1817. Type: Euphorbia supina Raf.

Euphorbia section Anisophyllum Roeper in Duby, A. P. de Candolle,

Bot. Gall. ed 2, 1: 412. 1828. Type: Euphorbia peplis L., designated

by Wheeler, Rhodora 43: 111. 1941.

Euphorbia subgenus Chamaesyce Caesalp. ex Reichenbach, Fl. Germ.

Excurs., 755. 1832. Type: Euphorbia chamaesyce L., designated by

Wheeler, Rhodora 43: 111. 1941.

Ditritra Raf., Fl. Tell., 4: 114. 1838. Type: Ditritra hirta

Raf., designated by Wheeler, Am. Mid. Nat. 30: 465. 1943.

Endoisila Raf., Fl. Tell., 4: 114. 1838. Type: Endoisila

myrsinites Raf.









Aplarina Raf., New Fl. N. Amer., 99. 1838. Type: Aplarina

prostrata Raf., designated by Wheeler, Rhodora 43: 111. 1941.

Annual or perennial herbs, often suffrutescent, or shrubs;

prostrate to ascending or erect; glabrous or variously pubescent; milky

latex in all parts. Main axis aborting, secondary axes few to many,

wiry or thicker, sometimes rooting at nodes. Leaves opposite, petiolate

with inter-petiolar stipules; blades expanded, bases inequilateral.

Cyathia terminal but appearing axillary, solitary at nodes or clustered

in cymules; lobes five, approximating glands; glands four or with fifth

vestigial, alternating with lobes, with or without petalloid appendages.

Staminate flowers monandrous, naked, few to many, maturing serially,-

exserted when mature, anther dehiscence transverse. Pistillate flower

solitary, terminal; perianth reduced to pad of tissue or vestigial

calyx; very three-celled, each with a single ovule; styles three, free

or joined at base, usually bifid for part of length. Capsule dehiscent

at sutures. Seed ecarunculate, coat smooth or variously sculptured.

Plants of disturbed habitats, probably poor competitors in

many cases. Many species favor sandy soils, either coastal or inland

and often unstable, while others are restricted to pinelands whose

characteristic flora is maintained by periodic burning. The most common

species, C. maculata in Florida and C. hyssopifolia in the New World

tropics as a whole, follow man, and are frequent as weeds in paved areas

and cultivated or abandoned ground. Many species have a wide geographic

range, passing unfavorable periods of the year as seed or with growth

almost quiescent and quickly resumed when conditions improve. Seed pro-

duction is abundant in most species.









The use of the taxonomic categories "species", subspecies", and

"variety" in this work follows conventional lines (see, for example,

Davis and Heywood, 1963). They are considered to constitute an hier-

archy, and the increasing degree of morphological homogeneity as the

rank of the group decreases is taken to imply a greater freedom of

gene interchange within the group. The use of these three ranks is

a compromise between the naming as a distinct taxon of any population

which is prevented by any means from exchanging genes with other popu-

lations, whether or not morphological differences have become established,

and the more classical approach of applying names only to taxa which are

discrete by a set number of morphological characteristics.




Key to the Species of Chamaesyce in the Caribbean


1. Much-branched shrub, low-growing or 2-6 m. tall ..... 2.

I. Annual or perennial herbs, often woody at the base
and suffrutescent, but rarely reaching 1 m. ..... 3.

2. Leaves fleshy, subglobose or linear; closely branched
with stems often almost prostrate .... C. vaqinulata.

2. Leaves membranous, ovate to ligulate; branching
divaricate, stems upright ..... 2. C. articulata.

3. Capsule glabrous ..... 4.

3. Capsule pubescent ..... 24.

4. Plants erect or ascending ..... 5.

4. Plants prostrate to decumbent ..... 12.

5. Leaf margins manifestly toothed, or, if entire,
leaves ligulate ..... 6.

5. Leaf margins entire or obscurely toothed below apex,
leaves never ligulate ..... 10.









6. Stems several to many from the crown of a heavy root-
stock, wiry, rarely more than 1 mm. diameter ..... 13. C. brittonii.

6. Stems few to several, thick, sometimes
reaching 4 mm. diameter ..... 7.

7. Cyathia few, borne singly or rarely in groups of two
or three; plant perennial ..... 12. C. parciflora.


7. Cyathia many, grouped on short leafy laterals or in
glomerules; annual but sometimes perennating


..... 8.


8. Capsule less than 1.4 mm. long; cyathia usually in
peduncled leafless glomerules ..... 9

8. Capsule more than 1.6 mm. long; cyathia on
leafy laterals


9. Tips of shoots pubescent, at least in lines on
the side of the stem; leaves pilose beneath; seed
with a rippled surface

9. Tips glabrous; leaves glabrous or sparsely pilose
beneath; seed with 2-4 lateral ridges
on each face .....


. C. hypericifolia.


..... 9.


..... 8. C. nutans.



10. C. hyssopifolia.


10. (5) Leaves and young stems fleshy; stipules conspicuous,
membranous, ca. 1 mm. long, white ..... 3. C. buxifolia.


10. (5) Leaves membranous or somewhat coriaceous, young
stems not fleshy; stipules not conspicuous unless
because of ciliate inner surface, coriaceous,
ca. 0.5 mm. long, brown


..... 11.


11. Stipules densely white ciliate on surface nearest stem;
leaf and young stem surface minutely white-papillose;
glands dark, appendages obsolete .... 4. C. lecheoides.

11. Stipules not ciliate; stem surface smooth, somewhat waxy,
leaf surface with raised markings but not papillose;
glands green or red, appendages small or to twice
width of gland ..... 6. C. porteriana.


12. (4) Stems wiry and stiff, rarely exceeding 0.5 mm.
diameter, usually many from root stock

12. (4) Stems flexible rather than wiry, few to several
from rootstock, either reaching 2-3 mm. diameter or
minute, threadlike


13. Leaf margins entire


..... 13.



..... 16.

..... 14.










Leaf margins serrate, at least in basal leaves


14. Leaves deltoid, margin somewhat revolute; glands fleshy,
appendages obsolete or minute ..... 41. C. deltoidea ssp. deltoidea
var. deltoidea

14. Leaves orbicular to elliptic, plane; glands cupped,
appendages prominent, to three times
as wide as gland ..... 19. C. camaguayensis.

15. All leaves serrate, most 4-8 mm. long; plants usually
decumbent ..... 18. C. torralbasii.


15. Only basal leaves serrate, most 2-3 mm. (rarely
to 5 mm.) long; plants usually forming a
lax, prostrate mat ..... 17. C.

16. (12) Leaves and stems pubescent ..... 39. C

16. (12) Leaves and stems glabrous

17. Stems minute, threadlike, tomentose on upper surface;
cyathia subsessile; styles undivided at apex; capsule
less than 0.7 mm. long ..... 42

17. Stems often reaching 2-3 mm. diameter, glabrous;
cyathia with an obvious peduncle, styles bifid for
part of length; capsule more than 1 mm. long

18. Stipules not united, or minutely so at base, deeply
parted or laciniate

18. Stipules joined, at least on upper or lower surface of
tips of branches, fringed or entire, never laciniate

19. Seeds strongly angled; appendages conspicuous; leaves
ovate to ovate-oblong, almost as wide as long..... 20.

19. Seeds terete to obscurely angled; appendages incon-
spicuous or absent; leaves narrowly elliptic
to oblong, much longer than wide


centunculoides.

. paredonensis.

..... 17.



. C. tumistyla.


..... 18.


..... 19.


..... 22.


C. cordifolia.



..... 20.


20. Capsule more than 2.5 mm. long; seed cuneiform,
more than 2 mm. long ..... 21. C. polygonifolia.


20. Capsule less than 2.5 mm. long; seed ovoid, less
than 2 mm. long

21. Seed 1.4-1.9 mm. long; leaves usually fleshy, size
often widely different on main stem
and laterals ..... 2:


..... 21.


2. C. ammannioides.


..... 15.











21. Seed 1-1.4 mm. long; leaves not fleshy, all
similar in size ..... 23. C. cumulicola.

22. (18) Stipules toothed at apex, white, prominent
although only ca. 0.5 mm. long; faces of seed smooth;
glands brown or deep purple, appendages white ..... 16. C. serpens.

22. (18) Stipules toothed or bifid almost to base, green or
tan colored, not conspicuous although often more than
1 mm. long; faces of seed obscurely transversely
ridged; glands greenish, appendages white, yellor or pink ..... 23.

23. Stipules short, to 0.4 mm.; texture of all parts of
plant subcoriaceous; leaf and stem surface
somewhat papillose ..... 14. C. cowellii.

23. Stipules long, often more than 1 mm.; leaf texture only
rarely firmer than membranous; leaf and stem surface
smooth ..... 15. C. blodqettii.

24. (3) Leaf margins manifestly serrate ..... 25.

24. (3) Leaf margins entire or obscurely toothed ..... 36.

25. Plants erect with one or few thick stems ..... 11. C. lasiocarpa.

25. Plants prostrate to decumbent with several to many
stems which rarely exceed 2 mm. diameter ..... 26.

26. Cyathia solitary at leafy nodes, appearing clustered
if on congested laterals, but not in glomerules ..... 27.

26. Cyathia in peduncled glomerules ..... 34.

27. Appendages of glands subequal in size, often much
reduced, or if two appendages much longer than other
two, capsule not fully exserted, splitting one side
of cyathium at maturity, glands subcircular or some-
what transversely elongated ..... 28.

27. Appendages of glands greatly unequal in size, one pair
longer than the other pair; capsule fully exserted and
nutant at maturity, often concealed by long appendages;
glands enlarged in length, to extend much of the
distance around rim of cyathium ..... 33.

28. Capsule pubescent only along the angles ..... 29.

28. Capsule pubescent all over ..... 30.










29. Stem short-pubescent in lines at sides and so
upper surface; faces of seed with deep,
transverse furrows


29. Stem long-hirsute, at least in lines at sides;
surface of seed rippled

30. Capsule not completely exserted, splitting one
side of cyathium at maturity


Capsule completely exserted at maturity


imetimes on


..... 24. C. prostrata.


..... 25. C. mendezii.


.... 32. C. thymifolia.


..... 31.


Leaf margin and coarsely serrate, often incised
1/2-1/3 distance to midrib; stems wiry, scarcely reaching
1 mm, diameter, many from heavy root stock ..... 26. C. hepatica.


31. Leaf margin serrate but not deeply incised; stems not
wiry, usually 1 mm. or more in diameter, few to
several from annual or perennating root stock

32. Stem deliquescent through dichotomous branching in upper
part; leaf texture heavy; seeds with two broad faces
and one angle acute, almost triangular in
cross-section ..... 27. C


.... 32.


. helwiqii.


32. Stem excurrent, forming only congested leafy laterals in
upper part; leaf texture membranous; seeds with all
faces subequal, angles obtuse, almost square
in cross-section ..... 31. C. maculata.

33. (27) Stems long-pilose on upper surface; leaf apex acute;
cyathia congested on short laterals ..... 34. C. conferta.

33. (27) Stems short-tomentose or strigose on upper surface;
leaf apex obtuse or rounded; cyathia borne singly or
in small groups at upper nodes, if on laterals
not congested ..... 33. C. adenoptera.


34. (26) Leaf margins crenate-dentate or roundly
serrate, apex obtuse

34. (26) Leaf margins sharply serrate, apex acute


.... 30. C. berteriana.

..... 35.


35. Cymules terminal and lateral on leafless peduncles; stem
branching at base but only infrequently near tips (when
plant may appear to have cymules on laterals with a pair
of subtending leaves); mostly robust, ascending,
large-leaved plants ..... 28. C. hirta.

35. Cymules terminal and on leafy laterals; stem branching
freely; mostly low, small-leaved plants ..... 29. C. opthalmica.









36. (24) Plants suffrutescent; branches to 5 dm. long ..... 37.

36. (24) Plants herbaceous, or, if woody at the base, of small
stature, branches rarely exceeding 1 dm. long ..... 39.

37. Appendages obsolete; glands deep purple ..... 5. C. cayensis.

37. Appendages prominent; glands yellow or green ..... 38.

38. Open shrubs, branched from base; capsules more
than 2 mm. long ..... 7. C. myrtillifolia.

38. Suffrutescent, base mostly unbranched; capsules
1.3-1.5 mm. long ..... 6. C. porteriana var. keyensis.

39. (36) Stems pubescent on only one surface ..... 40.

39. (36) Stems pubescent on all surfaces ..... 41.

40. Stems tufted, upright, usually less than 0.5 mm.
diameter; leaf surface glabrous ..... 37. C. minutula.

40. Stems prostrate, usually ca. I mm. diameter;
scattered white hairs on
surface of leaf ..... 36. C. multinodis.

41. Cyathia in groups of 2-5 at nodes ..... 43. C. leonardii.

41. Cyathia solitary at nodes ..... 42.

42. Plants robust, stems 1-3 mm. diameter, to 3 dm.
long, leaves 4-9 mm. long ..... 35. C. garberi.

42. Plants delicate, stems wiry, scarcely reaching 1 mmn.
diameter even in old plants, to 2 dm. long,
leaves 2-5 mm. long ..... 43.

43. Plants closely appressed forming a dense mat, sometimes
becoming diffuse with age ..... 41. C. deltoidea.

43. Plants decumbent at tips or erect ..... 44.

44. Stems almost unbranched, villous-hirsute, tips canescent;
capsule reniform, sharply three-lobed,
angles acute ..... 40. C. pinetorum.

44. Stems branched freely, particularly in upper part of stem,
sparsely or densely short-pilose, tips only occasionally
canescent; capsule ovoid, roundly lobed,
angles obtuse ..... 38. C. turpinii.










1. CHAMAESYCE VAGINULATA (Griseb.) Millsp., Field Mus. Bot. 2:

302. 1909. Euphorbia vaqinulata Griseb., Fl. Brit. W. I. Islands, 52.

1859. Type: Turk Islands, Hjalm. (Lectotype: K !, fragment NY !).

Designation of the lectotype is made here. The specimen at Kew was

chosen because there is a strong probability that it was examined by

Grisebach. Many of Hjalmarsson's collections were deposited at Systema-

tisch-Geobotanisches Institute, Goettingen, and an isotype may exist

there. The photograph of a Hjalm. sheet from Kew at IJ can have no

standing as type material since it is not of this species but of C. ?

lecheoides (Millsp.) Millsp.

Shrub; prostrate or ascending to 2 m. Stem to 1.5 cm. diameter,

internodes to 3 cm. or congested; branching from base and throughout

length, laterals often lying in one plane, not rooting at nodes, glabrous,

becoming dark brown with age. Leaves fleshy; blades subglobular or

linear-elliptic, 1.5 mm. diameter or 5 x 1-1.5 mm., base subequal, sub-

cordate or rounded, margin entire, apex rounded, subarticulate from

petiole, glabrous, yellow-green to purple-brown; petiole 0.3-1 mm. long;

stipules sheathing stem and persistent, short, ciliate, white. Cyathia

solitary at nodes, often on vestigial branches; peduncle short, somewhat

expanded into obconical involucre, to 1 mm. diameter at mouth, lobes

fleshy, equalling glands, triangular, glabrous on outside, ciliate within,

glands transversely elliptic, 0.5 x 0.3 mm., fleshy, deep purple, append-

ages absent, fifth gland triangular, large, equalling lobes or rarely of

same form as other glands, sinus shallow. Staminate flowers 3-5 per

cyathium or absent, androphores glabrous, scarcely exserted. Gynophore

glabrous, to 1.5 mm. long at maturity, exserted, usually upright;









calyx a triangular pad of tissue; ovary glabrous; styles spreading,

o.4 mm. long, joined at base, bifid for 1/2 length. Capsule glabrous,

globose, 1.5 mm. long, 1.8-2 mm. at equator, roundly three-lobed,

angles rounded. Seed ovoid, 1.5 x 1.2 mm., scarcely angled, usually

white.

Restricted to a few islands of the southern Bahamas, but

locally frequent on these in rocky and sandy areas behind the shore

(Map 1).

Representative specimens seen: BAHAMAS. Inagua, 6 Mar. 1962,

Dunbar 107 (GH); Turtle Cove, Inagua, 19 Oct. 1904, Nash & Taylor 1170,

(NY); Caicos Islands: North Caicos, 24 Apr. 1954, Lewis s.n. (GH, IJ);

Bambarra, Middle Caicos, 2 July 1954, Proctor 9096 (IJ); South Caicos,

14-16 Dec. 1907, Wilson 7681 (GH, NY). Turks Islands: Waterloo, Grand

Turk, 20 Feb.-24 Mar. 1911, Millspaugh & Millspaugh 9055 (GH, NY);

Grand Turk, 27 Aug.-l Sept. 1905, Nash & Taylor 3764 (NY); Long Cay,

12 June 1954, Proctor 8793 (GH, IJ). Watling's Island: Southeast End,

27, 28 Nov. 1907, Wilson 7259 (NY). Castle Island, 22 Dec. 1907, Wilson

7790 (NY).

The fleshy leaves and involucres, and the shrubby habit with the

closely branched stems often held almost prostrate, make this quite

different from any other species in the area. Boissier (1862) grouped

it with endemics of the Galapagos Islands on the basis of its appearance,

fleshiness, and the persistent, ciliate stipules that ring the stem,

but this morphological similarity is probably fortuitous and not an indi-

cation of close common ancestry. Many species of other families grow in














































I NEW YORK OTif (l .Iiln

S9- ? TH-E C AICO



plate 1. CHAMAESYCE VAGINULATA (Griseb.) Millsp.

C. va inulata (Griseb.) Millsp., Long Cay, Caicos Group, Bahamas, 7 Mar.
1911, Millspaugh & Millspaugh 9237 (NY).


.~

















'"> --- ------











Map 1. Distribution of C. vaqinulata





\ '


----- -
'.-- --. -








Map 2. Distribution of buxif




Map 2. Distribution of C. buxifolia









both the Bahamas and the Galapagos, but most of these are common through-

out the intervening tropics, and give no hint of a disjunction of range

such as this species would show from those which it resembles.

2. CHAMAESYCE ARTICULATA (Aubl.) Britton, Mem. N.Y. Bot. Card. 6:

574. 1916. Euphorbia articulata Aubl., P1. Guian., 1: 480. 1775. Type

in Herb. Brit. Mus. if extant.

Euphorbia linearis Retz., Observationes Botanicae, 3: 32. 1783.

Type: Bot. Mus., Lund, Sweden, if extant. E. linearls heterophylla

Kuntze, Rev.Gen. Pl., 605. 1891. C. linearis (Retz.) Millsp., Field

Mus. Bot. 2: 410. 1916.

Anisophyllum vahlii K1. & Gke., Monats. Akad. Berlin 1859: 36.

1859. Type: Specimen by Willdenow, in Bot. Mus. Berlin, if extant.

Euphorbia vahlii Willd. ex K1. & Gke., loc. cit. 37. 1859. Chamaesyce

vahlil (K1. & Gke.) P. Wilson, Bull. N.Y. Bot. Gard. 8: 395. 1917.

Shrub; to 6 m., erect. Stems to 2 cm. at base, internodes to

5 cm. long; divaricate-branched throughout length, glabrous or canescent

at tips, light brown becoming black with age. Leaves membranous; blades

elliptic, broadly so to linear, 20-60 x 2-12 mm., base subequal, rounded,

margin entire, slightly thickened, apex obtuse, mucronate, midrib promi-

nent on underside, glabrous or short pubescent, light green; petiole

1 to 1.5 mm. long; stipules joined, deltoid, to 0.5 mm. long, densely

ciliate on inner surface, somewhat persistent. Cyathia solitary; peduncles

5 to 7 mm. long, involucre obconical, to 2 mm. diameter at mouth, lobes

scarcely equalling glands, deltoid, glabrous on outside, densely ciliate

within, glands broadly transversely elliptic, to 1 mm. long, broadly

cupped, appendages absent, fifth gland deltoid, short, sinus broad, shallow.






Staminate flowers 8 20 per cyathium, androphores glabrous, to 2.5 mm.

long at maturity. Gynophore glabrous, to 3 mm. long at maturity, ex-

serted, nutant; calyx vestigial; ovary glabrous; styles erect, to 3 mm.

long, joined for half length, bifid in apical 1/3. Capsule glabrous or

minutely pubescent, ovoid or conical, 2.3 mm. long, 2.3 mm. wide above

base, roundly three-lobed, angles rounded. Seed elliptic-ovoid, 1.8 2

mm. long, 1.2 1.4 mm. radial width at equator, four-angled, the dorsal

faces longer and dorsal angle more acute than others, surface pitted and

wrinkled, brown or ashen.

Frequent from Puerto Rico south through the Lesser Antilles to the

northern coast of South America. No collections have been seen from the

Bahamas since those of Millspaugh in 1911 from East Caicos, but these

specimens are undoubtedly of the species (Map 10).

Representative specimens seen: BAHAMAS. East Caicos: Rocky plain,

Jacksonville and vicinity, 26-27 Feb. 1911, Millspaugh & Millspaugh

9086 (F, NY); Low scrub, Jacksonville and vicinity, 26-27 Feb. 1911,

Millspaugh & Millspaugh 9102 (F, NY). PUERTO RICO. Susua State Forest,

28 June 1961, Alain 9265 (IJ); Yauco, 2 Apr. 1880, Garber s.n. (GH);

Seashore, Fajardo, 17 Feb. 1900, Heller 4608 (F, GH, MO, NY); Santurce,

21 Feb. 1900, Heller 4678 (F, MO, NY); Rio Piedras, Fajardo, 4 July 1913,

Johnston 792 (NY); Guayanilla to Tallahoa, 13 Mar. 1913, Shafer 1993 (F,

NY); Fajardo, 19 May 1885, Sintenis 1612 (MO, NY). LESSER ANTILLES.

Culebra, 3-12 Mar. 1906, Britton & Wheeler 4 (F, NY); Culebras Island,

19-20 Jan. 1899, Millspaugh 1781 (F); Racoon Bay, St. Thomas, 31 Jan.-

4 Feb. 1913, Britton, Britton & Shafer 167 (F, NY); St. Thomas, Jan. 1887,

Eggers s.n. (F); Tortola, 16 Nov. 1918, Fishlock 263 (F, GH, NY); Anagada,

19-20 Feb. 1913, Britton & Fishlock 978 (NY); Barbuda, 6 Apr. 1956, Smith

10466 (GH, IJ). St. Kitts: Canada Estate, 8 Sept.-5 Oct. 1901, Britton &























































E. linearis Retz., St. Thomas, ? Bertero (MO).
E. linearis Retz., St. Thomas, ? Bertero (MO).







Cowell 263 (NY); Sir Timothy's Hill, 24 Dec. 1958, Proctor 18418 (IJ);

Nag's Head, 27 Dec. 1958, Proctor 18514 (IJ). Antigua: Santa Maria Hill,

25 Apr. 1937, Box 756 (MO); Nook Hill, 3 Oct. 1937, Box 1134 (MO).

Montserrat, 8 Feb. 1959, Proctor 19079 (IJ); Guadeloupe, 1892, Duss

2451 (NY); Martinique, Sieber 392 (F); St. Lucia, 22 Apr.-18 May 1950,

Howard 11417 (IJ). St. Vincent: Mt. Pleasant, 22 Mar. 1962, Cooley

8534 (GH); Bequia, 26-31 Mar. 1950, Howard 11269 (GH).

The affinities of this species are with C. myrtillifolia (L.)

Millsp. and with coastal species of the Bahamas and Florida such as

C. lecheoides (Millsp.) Millsp. and C. porteriana Small. These, however,

differ in size, habit, cyathial and stipular characters.

A number of pubescent collections of C. articulata have been

compared with the more common glabrous plants, but these prove to be

only the extremes of a range, with many intermediates showing occasional

shoots with minute velvety pubescence.

The name is accepted although no type material has been seen.

There is general agreement in the literature with this application, and

that E. linearis is a synonym.

3. CHAMAESYCE BUXIFOLIA (Lam.) Small, Fl. SE. US., 712. 1903.

Euphorbia buxifolia Lam., Encyc. 2: 421. 1786. Type: plate in manuscript

of Botanicum Americanum t.4, f.2 by Plumier, in the Bibliotheque General,

Museum D'Histoire Naturelle, Paris.

E. qlabrata Sw., Prodr., 76. 1788. Type: unknown. A specimen

from Swartz with this name is reported from Herb. Stockholm, Fawcett &

Rendle, Fl. Jam., 4(2): 336. 1920.

E. litoralis HBK., Nov. Gen. et Sp., 2: 44. 1817. Type: Cumana,

prope Bordones (Herb. Mus. Paris, not seen).

E. flexuosa HBK., Nov. Gen. et Sp., 2: 44. 1817. Type: Cumana,









Bonpland (Herb. Mus. Paris, not seen). E. buxifolia Lam. var. flexuosa

(HBK.) Soiss. in DC., Prod., 15(2): 15. 1862.

Perennial shrub or subshrub; erect to ascending, or nearly

decumbent in shifting sand, to 6 dm. Stems to 8 mm. diameter at naked

woody base, internodes to 3 cm.; branching sparse at base, more frequent

near shoot apex, glabrous, green becoming darker. Leaves somewhat

fleshy; blades ovate to elliptic, 5-12 x 3-6 mm., base slightly unequal,

truncate to cordate and obscuring stem, margin entire, apex obtuse, mid-

rib sometimes prominent on underside of folded leaf, glabrous, yellowish

to dark green and glaucous; petiole 0.5 to 1 mm. long; stipules united,

to 1 mm. long, ligulate or deltoid, short cleft or fringed, white.

Cyathia solitary at upper nodes; peduncles 0.5-1 mm. long, cyathia campanu-

late, to 0.7 mm. diameter at mouth, lobes equalling glands, triangular

glabrous outside, densely ciliate in throat, glands transversely elliptic,

to 0.5 mm. long, brown, fleshy, appendages slightly longer than, and as

wide as gland, white, margin entire or undulate, fifth gland linear,

shorter than lobes, sinus shallow, broad. Staminate flowers 12-20 per

cyathium, androphores glabrous, to 1 mm. long at maturity. Gynophore

glabrous, to 3 mm. long at maturity, exserted, nutant; calyx a triangular

pad of tissue; ovary glabrous; styles spreading, 0.3-0.4 mm. long,

joined at base, bifid for 1/2 length. Capsule glabrous, subspherical

1.5-2 mm. long, 2.2-5 mm. wide just below equator, deeply three-lobed,

angles rounded. Seed broadly ovoid, 1.3 mm. long, 1.2 mm. wide just below

equator, angles mere ridges except near apex, faces plump, convex, surface

obscurely pitted, ashen.

A frequent component of the vegetation of sandy and rocky shores,





























































.I I I II I. I


Plate 3. CHAMAESYCE BUXIFOLIA (Lam.) Small

C. buxifolia (Lam.) Small, Grassy Key, Monroe County, Florida, 9 Aug.
1963, Burch 153 (FLAS).








and of their associated beach-scrub; Bermuda, Florida in all coastal

areas south of Pinellas and Volusia counties, Bahamas, Yucatan, Cuba,

Jamaica, Hispaniola, Puerto Rico, Lesser Antilles and the northern

coast of Venezuela (Map 2).

Representative specimens seen: FLORIDA. Brevard: Cape Canaveral,

2-5 Apr. 1904, Burgess 668 (F, NY); Merritt Island, 18 Apr. 1946,

Foster & Smith 1364 (DUKE, F, FLAS, IJ, MO, NSC, NY); Malabar Cape, 17

Oct. 1964, Lakela (FLAS, USF). Broward: Port Everglades, 3 Aug. 1955,

Godfrey & Kral 53854 (FSU, NSC); Lauderdale Beach, 17 Dec. 1929, Moldenke

263a (DUKE, MO, NY). Collier: Vanderbilt Beach near Naples, 13 Aug. 1963,

Burch 182 (FLAS); Marco Island, 19 Mar. 1938, Standley 57622 (F). Dade:

Soldier Key, 30 Mar. 1904, Britton 337 (F, NY); Miami, May 1877, Garber

s.n., (FLAS, NY); Key Biscayne, 17 Jan. 1937, McAllister 210a (DUKE);

Lemon City, 6 Nov. 1903, Small & Carter 1116 (NY). Flagler: Beach near

Marineland, 18 Nov. 1961, Godfrey 61675 (FSU). Indian River: Vero Beach,

7 Aug. 1963, Burch 112 (FLAS). Lee: Sanibel Island, 19 Dec. 1953,

Cooley 2401 (FLAS, NY); Punta Rassa, July-Aug. 1900, Hitchcock 325 (F,

MO); Captiva Island, 21 May 1901, Tracy 7226 (F, MO, NY). Manatee:

Anna Maria Key, 12 July 1917, Cuthbert 1578 (FLAS, NY); Sneed's Island,

9 Oct. 1899, Tracy 6377 (F, MO, NY). Martin: Atlantic side of Jupiter

Island, 7 Aug. 1963, Burch 125 (FLAS). Monroe: Cape Sable, Middle Cape,

18 Apr. 1964, Burch & Ward 304 (BM, BUS, E, F, FLAS, GH, IJ, MO, NSC, NY);

New Found Harbor Key, 13-14 Feb. 1935, Killip 31395 (NY); Marquesas Keys,

10-12 Mar. 1904, Lansing, Jr. 2225 (F, NY); Key Largo, 12 Jan. 1930,

Moldenke 402 (DUKE, MO); Key West, 1874, Palmer 499 (MO, NY); Big Pine

Key, 27 June 1960, Webster & Samuel 10210 (DUKE, IJ, PUR). Palm Beach:








Beach between Palm Beach and Lake Worth, 7 Aug. 1963, Burch 118 (FLAS);

Palm Beach, 24 May 1895, Curtiss 5404 (NY); Juno Beach, 20 Apr. 1962,

Lakela 24955 (FLAS, USF); Lake Worth Inlet, 28 Dec. 1895, Webber 236

(F, MO). Pinellas: St. Petersburg, Williamson s.n. (NY). St. Lucie:

North of Ft. Pierce, 7 Aug. 1963, Burch 120 (FLAS). Sarasota: Longboat

Key, 20 Nov. 1964, Godfrey 65236 (FSU); South end of Longboat Key,

6 Oct. 1964, Long & Lakela 27547 (FLAS, USF); Casey Keys, Spring 1887,

Rothrock s.n. (F, NY). Volusia: Turtle Mound, 27 Apr. 1961, Ray, Wood,

Smith & Eaton 10821 (FSU, NCU). BAHAMAS. Great Bahama, 5-13 Feb. 1905,

Britton & Millspaugh 2365 (F, NY); Bimini, 27 July 1964, Stimson 776

(DUKE, FLAS); Riding Rocks, Great Bahama Bank, 13 May 1909, Wilson 7914

(F, MO, NY); New Providence, Nov. 1890, Hitchcock 26 (F, MO); Salt Key,

21 May 1909, Wilson 8089 (F, MO, NY); Watling's Island, Graham's Harbor,

26 Nov. 1907, Wilson 7228 (F, NY); Hog Island, Jan. 1905, Wight 55a

(F, NY); Inagua, 15-25 Aug. 1960, Williams 7 (DUKE, PUR); South Caicos,

27 June 1954, Proctor 8981 (IJ); Grand Turk, 22-27 Apr. 1954, Lewis s.n.

(IJ). CUBA. Isles of Pines: Caleta Cocodrilos, 8 Mar. 1916, Britton,

Wilson & Leon 15295 (F, NY). Havana: Havana, 6 May 1905, Curtiss 752

(F, MO, NY). Oriente: Santiago, Siboney, 1917, Ekman 8719 (F). Camaguey:

Cayo Sabinal, 9 Mar. 1909, Shafer 699 (F, NY); Cayo Palo, 24 Oct. 1909,

Shafer 2733 (F, NY). Pinar del Rio: Las Martinas to the coast, 19 Dec.

1911, Shafer 11105, (MO, NY). CAYMAN ISLANDS. Cayman Brac., South

Shore, 10 Feb. 1899, Millspaugh 1233 (F); Grand Cayman, Grape Tree Point,

14 Apr. 1956, Proctor 11973 (IJ). JAMAICA. Portland: Port Antonio, 21

June 1897, Fredholm 3038 (NY). St. Catherine: Healthshire Hills, 3 May

1913, Harris 11627 (F, MO, NY, UCWI). St. Elizabeth: Long Acre Point,








13 Sept. 1907, Harris 9967 (F, NY, UCWI). St. James: Montego Bay,

June 1951, Robbins 423 (UCWI). St. Mary: Galina Point, 7 Dec. 1952,

Proctor 7452 (IJ). St. Thomas: Morant Point, 1-13 Mar. 1909, Britton

4098 (F, NY). Westmoreland: Negril area, Homer's Cove, 23 June 1959,

Webster, Ellis & Miller 8501 (PUR). HAITI. Bayeux, 28 Nov. 1924,

Ekman H2676 (IJ); Port de Paix, 28 Apr. 1925, Ekman H3957 (IJ); Gonave

Island, Aug. 1927, Eyerdam 213 (F, NY); Tortue Island, 3-10 May 1929,

Leonard & Leonard 15328 (MO). DOMINICAN REPUBLIC. Monte Cristi: Monte

Grande, 12 July 1929, Ekman H13165 (F, PUR). Barahona: San Domingo,

May 1911, Fuertes 1111 (F, NY); Trail between Pedernales and Aceitial,

8-12 Aug. 1946, Howard 8106 (NY). PUERTO RICO. Condado, San Juan,

16 Sept. 1941, Blomquist 11789 (DUKE); Fajardo, 4 July 1913, Johnston

791 (NY); Ratones Island, 21 Jan. 1899, Millspaugh 658 (F); Salinas de

CaboRojo, 6 Feb. 1885, Sintenis 559 (UCWI); Beach at Santurce, 21 Feb.

1900, Heller 4681 (F, MO, NY). LESSER ANTILLES. Soldier Bay, St.

Thomas, Dec. 1880, Egger s.n. (F); St. Croix, I Nov. 1895, Ricksecker

36 (F, MO, NY); Anguilla, 1908, Boldingh 3561 (NY); Barbuda, 27 Apr.

1959, Cowan 1656 (IJ, NY); Guadeloupe, 1894, Duss 2741 (MO, NY);

Martinique, 25 June 1939, Egler 173 (NY); St. Lucia, I Aug. 1958, Sauer

2174 (F); St. Vincent, Smith & Smith, 208 (NY); Grenada, Pointe Saline,

23 Sept. 1945, Beard 1222 (MO, NY). Extra-territorial specimens: BERMUDA.

Ducking Stool, 31 Aug.-20 Sept. 1905, Brown & Britton 6 (F, NY); Agar's

Island, 22 July 1913, Collins 219 (F, NY); Rocky shore, 18 Mar. 1910,

Flynn 101 (MO); Hamilton Bay Islets,. 29 Dec. 1898, Millspaugh 10 (F).

The species named by Kunth are no more than growth forms of this

taxon, which is just as plastic in its response to soil and wind conditions









as the completely herbaceous species.

Euphorbia mesembrianthemifolia Jacq., Enum. PI. Carib., 22.

1760, and Sel. Stirp. Amer., 150. 1763, was put into synonymy for

this species by Boissier (1862) on the basis of Jacquin's description.

This is unacceptable, since, if the name can be linked to the species

beyond reasonable doubt, it is much earlier than Lamarck's and would

have to replace it. Jacquin's description suggests this species in

many respects, but has no character which is diagnostic. However, if

it could be shown that specimens of this entity were in Jacquin's

possession at the time that his description was written, this would

support the acceptance of his name. Mr. W. T. Steam of the British

Museum (Natural History) was kind enough to search their herbarium, and

Dr. K. H. Rechinger checked that of the Naturhistorisches Museum, Vienna,

but in neither case were any specimens found which were known to have

passed through Jacquin's hands at this time. Many of Jacquin's specimens

were lodged in these two herbaria, and the absence of any relevant to

this problem, together with the fact that Boissier did not see any

specimens to support the name, suggest that none have survived. With

nothing more than Jacquin's description to typify the name E. mesembri-

anthemifolia, it would be irresponsible to accept it and upset the well-

understood use of C. buxifolia, which will be maintained here until some

firm evidence supports a change.

4. CHAMAESYCE LECHEOIDES (Millsp.) Millsp., Field Mus. Bot. 2:

301. 1909.

Shrub; erect or ascending to 4 dm. Stem to 8 mm. diameter, inter-

nodes to 2 cm.; branched either from crown of root and throughout length









or from only the upper half of the single or few stems from base,

branching open or strict, old stem ringed by stipule scars, glabrous,

brown. Leaves membranous, somewhat coriaceous; blades ovate to linear

elliptic, 3-7 x 1.5 5 mm., midrib sometimes prominent on under side

of leaf, marked by depression on upper surface, glabrous, the surface

minutely white-papillose, base inequilateral, rounded to subcordate,

margin entire, somewhat revolute, rarely crenulate or serrate toward

apex, apex obtuse or rounded, gray-green or pink; petiole 0.5 I mm.

long; stipules distinct or joined, deltoid to 1 mm., densely ciliate

on inner surface, brown, cilia white. Cyathia in upper nodes; peduncles

to 1 mm. long, involucre campanulate, 1-1.6 mm. diameter at mouth, lobes

equalling glands, triangular, glabrous on outside, densely pilose within,

glands broadly transversely elliptic, 0.4 0.7 by 0.2 mm., often dark

purple, rarely much reduced, appendages obsolete, fifth gland vestigial

or absent, sinus V-shaped, to 1/4 depth of cyathia. Staminate flowers

to 30 per cyathium, androphores glabrous, to 0.7 mm. at maturity.

Gynophores glabrous, to 2 mm. long at maturity, exserted, nutant or

upright; calyx a triangular pad of tissue; ovary glabrous; styles spread-

ing, 0.3 mm. long, joined at base, subentire to 1/3 bifid, somewhat

clavate. Capsule glabrous, ovoid, 1.5 mm. long, 2 mm. wide between equator

and base, three-lobed, angles obtuse. Aberrant inflorescences are quite

frequent, producing a thick (to 0.6 mm.) gynophore and sterile capsule to

2.5 mm. long. Seed angular-ovoid, 1 mm. long, to 0.7 mm. diameter at

equator, ventral angle somewhat obscure, others well marked, faces flat,

somewhat pitted, gray-brown.









Key to varieties of C. lecheoides


1. Plant branching from crown of root and throughout length of

stem; branches decumbent ..... a. var. wilsonii.

1. Plant with few stems from ground level, branching mostly in

upper part of stem; branches erect, strict ..... 2.

2. Leaves linear-ovate, much longer than wide, apex

acute ..... b. var. lecheoides.

2. Leaves ovate-elliptic, scarcely longer than

wide, apex obtuse ..... c. var. exumensis.

4a. C. LECHEOIDES (Millsp.) Millsp. var. WILSONII (Millsp.) comb.

nov. C. wilsonii Millsp., Field Mus. Bot. 2: 301. 1909. Type: Castle

Island, Bahamas, 22 Dec. 1907, P. Wilson 7795 (Holotype: F !; Isotype:

NY 1).

Restricted to the whitelands of islands of the Inagua, Turks

and Caicos groups at the southern end of the Bahamas (Map 3 ).

Representative specimens seen: BAHAMAS. Great Inagua, near

lighthouse, 20 Feb. 1963, Dunbar 239 (GH); Inagua, 14 Oct. 1904, Nash

& Taylor 1034 (NY); South Caicos, near East Bay, 17 June 1954, Proctor

8828 (IJ); Turks Islands, Salt Cay, 10 Mar. 1911, Millspaugh & Millspaugh

9253 (NY).

Much branched from a heavy rootstock, with the branches lying

almost prostrate. Distinguished from var. lecheoides only on the basis

of its many wiry stems, much-branched throughout length.

4b. C. LECHEOIDES (Millsp.) Millsp. var. LECHEOIDES. Euphorbia

lecheoides Millsp., Field Mus. Bot. 2: 163. 1906. Type: Grand Turk,

Turks Islands, Bahamas, 27 Aug.-l Sept. 1905, G. V. Nash & N. Taylor 3888









(Lectotype: F !; Isotype: NY !). Selection of type made here.

Restricted to scrub areas and rocky shores on islands in the

southeast of the Bahamas chain from Watling's Island to the Turks group

(Map 4).

Representative specimens seen: BAHAMAS. Watling's Island,

S. W. End, 30 Nov. 1907, Wilson 7347 (NY); Long Cay, 7-17 Dec. 1905,

Brace 4179 (NY); Crooked Island, 9-23 Jan. 1906, Brace 4588 (NY);

Mariguana, 6-8 Dec. 1907, Wilson 7534 (NY); Inagua, 21 Jan. 1962, Dunbar

53 (GH). South Caicos: Northern Hills, 19 June 1954, Proctor 8887 (GH,

IJ); Cockburn Harbor, 28 Mar. 1958, Proctor 17497 (IJ). Between

Commissioner's House and airfield, Grand Turk, 13 June 1954, Proctor

8799 (GH, IJ).

Usually rather few, stout stems from ground level which remain

simple for about half their length. Branching is often, but not invariably,

rather strict. The leaves are smaller and narrower than those of var.

exumensis.

4c. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS (Millsp.) comb.

nov. C. exumensis Millsp., Field Mus. Bot. 2: 301. 1909. Type: Stocking

Island, Great Exuma, Bahamas, N. L. Britton & C. F. Millspaugh 3071

(Holotype: F 1; Isotype: NY !).

Whitelands and coastal sands of islands in the Bahama group from

Eleuthera south to Great Ragged Island (Map 5).

Representative specimens seen: BAHAMAS. Eleuthera, S. E. Point,

27 July 1960, Webster & Williams 10659 (DUKE, IJ, PUR); Little San Salva-

dor, 25-26 Feb. 1907, Britton & Millspaugh 5666 (NY); Cat Island, Bight

and vicinity, 1-6 Mar. 1907, Britton & Millspaugh 5804 (NY); Watling's




















































1 $ 0 0 f11 12 II 14 .



I: r T


Plate 4. C. LECHEOIDES (Millsp.) Millsp. var. wILSONII (Millsp.) ined.

Isotype: C. wilsonil Millsp., Castle Island, Bahamas, 22 Dec. 1907,
WiIson 7795 (NY).


";246





p4







v .


t. I


1905, Nash & Taylor 3888 (NY).




48.

*



























7 t



















Plate 6. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS (Millsp.) ined.

Isotype: E. exumensis Millsp., Stocking Island, Bahamas, 22-28 Feb. 1905,
Millspaugh 3071 (NY).

































Map 3. Distribution of C. lecheoides var. wilsonii


Map 4. Distribution of C. lecheoides var. lecheoides














O

;o
'-^ 0 Y -r.


Map 5. Distribution of C. lecheoides var. exumensis


Map 6. Distribution of C. cayensis


~B









Island, 12-13 Mar. 1907, Britton & Millspeugh 6143 (NY); Conception

Island, 5 Jan. 1932, Fairchild 16 (GH); Rum Cay, 4 Dec. 1905, Brace

3927 (NY); Long Island, Clarence Town, 16-19 Mar. 1907, Britton &

Millspaugh 6323 (NY); Great Ragged Island, 24-25 Dec. 1907, Wilson

7807 (NY).

Stems usually rather few and stout from ground level, branching

freely only in upper half of stem. Branching often obtuse. The leaves

are usually rather wider than those of the other two varieties.

These varieties are not well marked, but the key characters

serve to separate most specimens seen. The slight difference in range may

prove to be significant in allowing speciation to proceed if the vegeta-

tion of the islands remain undisturbed, and communication between them

remains restricted.

The ciliate stipules and glabrous, markedly papillose surface of

stems and young shoots distinguishes these plants from all others of

similar size in the area. A similar surface marking, often much less

pronounced, is found in a number of species from islands to the south,

and is usually associated with plants from a saline habitat.

5. CHAMAESYCE CAYENSIS (Millsp.) Millsp., Field Mus. Bot. 2:

302. 1909.

Subshrub; ascending to 3 dm. Stems to 3 mm. wide, internodes to

2 cm.; branching sparingly throughout, tomentose, sometimes canescent at

tips, somewhat glabrate, brown or gray. Leaves membranous or sub-

coriaceous; blades ovate, sometimes orbicular or lanceolate, 5-12 x 3-9

mm., base inequilateral, rounded or cuneate, margin entire, slightly

thickened, apex obtuse or rounded, midrib sunken on underside of leaf,








both surfaces sparsely to densely tomentose, or woolly, the surface

minutely white-papillose, yellowish green or pink, often ashen; petiole

to 1 mm. long; stipules united, triangular, to 1 mm. Cyathia solitary

at uppermost nodes; peduncle to I mm. long, involucre broadly obconical,

1.5 2 mm. diameter at mouth, lobes scarcely equalling glands, triangular,

strigose on outside, hairy within, glands broadly transversely elliptic,

0.7 x 0.5 mm., appendages obsolete, fifth gland vestigial or absent,

sinus shallow. Staminate flowers to 30 per cyathium, androphores glab-

rous, to 1 mm. long at maturity, exserted, somewhat nutant; calyx obso-

lete; ovary pilose; styles spreading, 0.3 0.4 mm. long, joined at base,

subentire to 1/3 bifid, somewhat clavate. Capsule strigose, broadly

ovoid, 1.7 mm. long, 1.8 2.1 mm. wide between base and equator, three-

lobed, angles rounded. Seed cuneiform, to 1.2 mm. long, 0.6 mm. wide

above base, strongly four-angled with ventral angle somewhat rounder than

others, faces flat, wrinkled, brown.

Restricted to the coastal whitelands and pine and palmetto-scrub

areas of the northern and western islands of the Bahamas chain (Map 6).

Representative specimens seen: BAHAMAS. Grand Bahama: Freeport,

30 Oct. 1964, McDaniel & Schmiederer 5395 (FSU); Freeport, 8 Aug. 1960,

Webster 10896 (DUKE, PUR); Abaco, 8 Dec. 1904, Brace 1571 (NY). Berry

Islands: Whale Cay, 29 Jan. 1905, Britton & Millspaugh 2178 (NY); Great

Sturrup Cay, 1 Feb. 1905, Britton & Millspaugh 2266 (NY); Great Harbor

Cay,2-3Feb. 1905, Britton & Millspaugh 2334 (NY). Salt Key Bank: North

End, Anguilla Isles, 15-18 May 1909, Wilson 7961 (MO, NY); South End,

Anguilla Isles, 15-18 May 1909, Wilson 8027 (MO, NY); Water Key, 22 May

1909, Wilson 8139 (MO, NY).












































12 I


Plate 7. CHAMAESYCE CAYENSIS (Millsp.)

Isotype: E. cayensis Millsp., Joulter's
Millspaugh 2295 (NY).


Millsp.

Cays, Bahamas, 11 April 1904,









Recent collections have extended the geographical range, and

also the habitats, from which this species is known. The whitelands,

from which the type collections came, are sandy areas with a very limited

flora, but the collections from Grand Bahama, from the pinelands and from

pine/palm scrub, suggest that the plant is able to tolerate quite strong

competition and shading.

C. bracei Millsp. is no more than a trivial variation of this

species. Of the two specimens cited by Millspaugh with his description,

one is also cited for C. cayensis, and the other differs from the usual

appearance of that species only in the degree of pubescence.

6. CHAMAESYCE PORTERIANA Small, Fl. SE. US., 711. 1903.

Annual or perennating; erect or ascending to 5 dm. Stems to

4 mm. thick at somewhat woody base, internodes to 2 cm.; branching sparse,

mostly from upper nodes or, rarely, much branched, branches at times at

a very acute angle to stem, broom-like, more usually lax, glabrous or

short-tomentose, green, often with a purple cast, or red. Leaves

membranous or somewhat chartaceous; blades elliptic or ovate, in some

suborbicular, in others almost lanceolate, 4-10 x 1-5 mm., base inequi-

lateral, rounded or subcordate, margin entire or rarely somewhat serrate,

apex obtuse to acute, sometimes mucronate, glabrous or sparsely pubescent,

rarely tomentose, green, often suffused purple or red particularly on

underside; petiole 0.5 1 mm. long; stipules distinct or joined at base,

triangular, to 0.4 mm. long, entire or 2-3 parted. Cyathia solitary at

upper nodes; peduncle 0.5 1 mm. long, involucre campanulate, 0.8 1.2

mm. diameter, lobes scarcely equalling glands, triangular, entire or

short-cleft, glabrous or strigose on outside, pilose at mouth within,









glands fleshy, somewhat concave, transversely elliptic, 0.3 0.5 x

0.1 mm., appendages a mere rim, to twice as wide as gland, white, pink,

or dark red, margin entire or crenulate, fifth gland linear, shorter

than lobes or obsolete, sinus broad and shallow. Staminate flowers 8-14

or more per cyathium, androphores glabrous, ca. I mm. long at maturity.

Gynophore glabrous or sparsely pilose, 1-1.5 mm. long at maturity, ex-

serted, nutant; calyx a triangular pad of tissue; ovary glabrous or

strigose; styles spreading, 0.4 mm. long, minutely joined at base, bifid

for 1/2 length. Capsule glabrous or strigose, ovoid, 1.5 mm. long,

1.8 2.1 mm. wide close to base, three-lobed, angles obtuse. Seed

oblong-ellipsoid, 0.7 0.9 mm. long, 0.6 mm. diameter at equator, four-

angled, with ventral angle somewhat rounded, faces flat, obscurely

wrinkled, red-brown or gray.


Key to varieties of C. porteriana


1. Plant pubescent, sometimes glabrate; branching usually rather

lax; leaves membranous, usually deep green; appendages white, margin

entire or crenulate ..... a. var. keyensis.

1. Plant glabrous or, rarely, sparsely short-pubescent; branching

usually strict; leaves parchment-like, often flushed with red or

purple; appendages white or pink, margins entire .....2.

2. Leaves ovate-elliptic, most almost as wide as long, apex obtuse to

rounded; usually rather sparingly branched ..... b. var. porteriana.

2. Leaves linear-elliptic, all except oldest much longer than wide,

apex acute; freely branched, often strict and broom-like

..... c. var. scoparia.









6a. CHAMAESYCE PORTERIANA Small var. KEYENSIS (Small) comb. nov.

Chamaesyce kevensis Small, Torreya 28: 6. 1928. Type: No Name Key,

Monroe County, Florida, 4-5 Feb. 1916, J. K. Small 7439 (Holotype NY 9).

Restricted to coastal scrub and open hammocks of the Florida

Keys, as far north as Lignum Vitae Key (Map 7).

Representative specimens seen: FLORIDA. Monroe County: Lignum

Vitae Key, 13 Dec. 1921, Small, Small & Matthews 10260 (NY); Long Key,

25 Aug. 1949, Woodbury & Jackson s.n. (BUS); Greyhound Key, 21 Nov. 1964,

BLrch 562 (BM, BUS, DUKE, E, F, FLAS, GH, IJ, MO, NY); Ohio Key, 19 Nov.

1964, Burch 538 (FLAS, FSU, MO, NSC, PUR, US); Bahia Honda Key, 9 May

1919, Small, Cuthbert & Mattaus 9139 (FLAS, NY, US); Big Pine Key, 27

June 1960, Webster & Samuel 10216 (DUKE, PUR).

Strongly erect unless growing through dense grass, and evenly

branched, particularly in the upper third of the stem. Distinguished

from the other two varieties by its pubescence and the membranous leaves

which rarely have the suffusion of red or purple that the others show.

6b. CHAMAESYCE PORTERIANA Small var. PORTERIANA. Type: Miami,

Florida, 1877, A. P. Garber s.n. (Holotype: NY !).

Locally frequent in Dade and Monroe counties, Florida. In pine-

land on oolitic limestone from Miami south through the Everglades Keys,

and on the lower Florida Keys. (Map 8).

Representative specimens seen: FLORIDA. Dade County: Pinelands

between Cutler and Black Point Creek, 15 Nov. 1906, Small & Carter 2504

(NY); Pinelands east of Naranja, 14 Jan. 1909, Small & Carter 3066 (NY);

Camp Longview, Everglades, 11 Nov. 1912, Small 3705 (DUKE, FLAS); 6 mi.

west of entrance, Everglades National Park, 17 April 1964, Burch & Ward 298









(BM, BUS, E, F, FLAS, GH, IJ, MO, NSC, NY, PUR, US, USF). Monroe County:

Vacca Keys, 28-29 Jan. 1909, Small & Carter 3048 (NY); Boot Key, 7-12

April 1909, Britton 540 (NY); Big Pine Key, 14 Dec. 1951, Killip 41547

(NY); Key West, 19 Sept. 1954, Killip 44474 (US).

The leaves are broader than those of var. scoparia, and the plant

is usually more sparsely branched. Occasional specimens show sparse

pubescence on the underside of leaves and on the capsules, but the hairs

are shorter and stiffer than is usual in var. keyensis.

6c. CHAMAESYCE PORTERIANA Small var. SCOPARIA (Small) comb. nov.

Chamaesyce scoparia Small, Fl. Florida Keys, 81. 1913. Type: Big Pine

Key, Monroe County, Florida, 17 Nov. 1912, J. K. Small 3819 (Holotype:

NY 1).

Restricted to the lower Florida Keys, usually in pineland on

oolitic limestone, but occasionally in sand or in open hammocks (Map 9).

Representative specimens seen: FLORIDA. Monroe County: Pinelands,

Big Pine Key, 17 Nov. 1912, Small 3770 (DUKE, FLAS, NY); Pinelands, Big

Pine Key, 17 Dec. 1913, Small & Small 5030 (MO, NY); Pinelands, Big Pine

Key, 19 April 1942, Eyles & Eyles 8186 (GH); Pine woods, Big Pine Key, 10

Aug. 1963, Burch 140 (BM, E, F, FLAS, IJ, MO, NY, US); No Name Key, no

date, Curtiss s.n. (GH, NY); Edge of airport, Key West, 26 Dec. 1964,

G. Avery s.n. (FLAS, MO).

There is some intergradation of leaf form between this taxon and

var. porteriana, but the extremes are quite distinct. Plants showing

the strict branching that Small described as "broom-like" when describing

the species have so far only been found on Big Pine Key.

The differences between the type specimens of Small's species do




























































S'' 2 :I 7 1 Ii, I I II I


Plate 8. CHAMAESYCE PORTERIANA Small var. KEYENSIS (Small) ined.

Holotype: C. keyensis Small, No Name Key, Monroe County, Florida, 4-5
Feb. 1916, Small 7439 (NY).














/





Sr-




..I.i ,,
, ,, ', ,



^CBDLNya
PIDt^/f


Plate 9. CHAMAESYCE PORTERIANA Small var. PORTERIANA
Holotype: C. porteriana Small, Miami, Florida, 1877, Garber s.n. (NY).


"
11 ..,..

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/d~~







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.1


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-.^ ^



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2 1 I 7 A 9 1o 11 125 11 It 15


Plate 10. CHAMAESYCE PORTERIANA Small var. SCOPARIA (Small) ined.
Holotype: C. scoparia Small, Big Pine Key, Monroe County, Florida, 17
Nov. 1912, Small 3819 (NY).


\


































Map 7. Distribution of C. porteriana var. keyensis


Map 8. Distribution of C. porteriana var. porteriana




















9VB,


%II>


Map 9. Distribution of C. porteriana var. scoparia








-_- _- ---- ---_ _-




1< _____ ___ ..... -_ __
--. ----. -.- --




.. -





Map 10. Distribution of C. articulata and C. myrtillifolia


L










not hold up on examination of a range of collections. Pubescence

seems to be, in general, a very plastic character in the genus, and

can rarely be trusted as a major character for the differentiation of

species. The texture of the leaves and the coloring of the plant

appear to be correlated with habitat, and the branching pattern changes

radically with the accompanying vegetation and the period of time since

the area was exposed to fire. In consideration of these factors it

was deemed more realistic to reduce the species to varietal status until

some clear distinction between them can be demonstrated.

7. CHAMAESYCE MYRTILLIFOLIA (L.) Millsp., Field Mus. Bot. 2:

410. 1916. Euphorbia myrtillifolia L. Syst. Nat. ed. 10, 1048. 1759.

Shrub, erect, to 1 m. Stems to 5 mm. diameter, internodes to

4 cm. long; branching open, throughout length, short-pubescent, brown.

Leaves membranous; blades elliptic or obovate, 7-15 x 5-10 mm., base

oblique, obtuse or cuneate, margin serrate, often only in upper half,

apex emarginate, obscurely mucronate, midrib prominent on lower surface,

upper surface glabrous or sparsely pilose, lower pilose-pubescent, yellow-

green; petiole to 1 mm. long, short-pubescent; stipules joined for half

length, free part narrow, ciliate or toothed. Cyathia solitary at upper

nodes; peduncle to 2 mm. long, involucre obconical, to 1.5 mm. diameter

at mouth, lobes equalling glands, deltoid, toothed, pilose on outside and

within, glands transversely elliptic, 0.5 x 0.2 mm., dark brown, appendages

longer than glands and as wide, margin somewhat dentate, creamy-yellow,

fifth gland linear, short or obsolete, sinus shallow. Staminate flowers

18-30 per cyathium, androphores glabrous, to 1.5 mm. long at maturity.

Gynophore short-pubescent, to 3 mm. long at maturity, exserted, upright;




























































A I I II I

Plate II. CHAMAESYCE MYRTILLIFOLIA (L.) Millsp.

E. myrtillifolia L., Gordon Town, St. Andrew, Jamaica, 8 Dec. 1954,
Proctor 9595 (NY).









calyx a pad of tissue or obsolete; ovary sparsely pilose; styles spread-

ing, to I mm. long, joined at base, bifid for one-third length. Capsule

sparsely pubescent, glabrate, ovoid with truncate or cordiform base, 2.2

mm. long, 2.3 mm. wide above base, broadly three-lobed, angles rounded.

Seed long-conical, to 1.8 mm. long, triangular with ventral angle obso-

lete, others acute, dorsal faces convex, strongly wrinkled, base truncate,

dark reddish-brown.

Reported only from the Gordon Town and Guava Ridge area of the

Blue Mountains of Jamaica (Map 10).

Specimens seen: JAMAICA. St. Andrew: Near Gordon Town, 23 May

1894, Harris 5196 (NY); Valley of Hog Hole River above Gordon Town,

8 Dec. 1954, Proctor 9595 (IJ, NY); 1/4 mile northeast of Gordon Town,

20 Mar. 1956, Proctor 11874 (GH, IJ).

The affinities of this species are with the shrubby C. articulata,

but it differs from that species in.being smaller and having elliptic

leaves which are pilose on the under surface. The involucre of C. myr-

tillifolia is smaller, with less prominent glandular appendages, and the

style and other floral parts are reduced proportionally.

8. CHAMAESYCE NUTANS (Lag.) Small, Fl. SE. US. 712. 1903.

Euphorbia nutans Lag., Gen. et Spec. Pl. 17. 1816. Type: "Habit.[at]

in N.[ova] H.[ispania]. Perhaps at Madrid ." (Wheeler, 1941).

E. preslii Guss., Fl. Sic. Prod. 1: 539. 1827. Type: Palermo,

Italia, Todaro (fragment F seen by Wheeler). C. preslii (Guss.) Arthur,

Torreya 11: 260. 1912.

E. hypericifolia L. var. communis Engelm. in Emory, U.S. & Mex.

Bound. Surv. 2: 188. 1859. Type: New Mexico, 1851-52, C. Wright 1842









(fragment F seen by Wheeler).

E. hypericifolia L. sensu Michaux, Fl. Bor.-Am., 1803; Torrey,

Fl. State New York, 1843.

E. maculata L. sensu Wheeler, Contr. Gray Herb. 127: 74. 1939.

Annual; erect or ascending to 8 dm. Stem to 4 mm. thick at some-

what woody base, internodes to 3 cm. long; branched mostly from upper

nodes, young tips and a line on older stem densely tomentose, glabrate,

straw-colored or darker brown. Leaves membranous; blades ovate-

lanceolate, often somewhat oblong or falcate, 10-30 x 6-10 mm., base

inequilateral, rounded, margin serrate, apex acute to obtuse, glabrous

or sparsely long-pilose on upper surface, more generally long-pilose

below, both surfaces mid-green often with a red spot on upper surface

or the whole suffused with red; petiole 1-2 mm. long; stipules joined or

early distinct near tip of shoot, triangular to I mm. long, margin

toothed and ciliate, thin textured, brown. Cyathia borne singly in

lateral and terminal short-stalked compound dichasia; peduncle 0.5 2

mm. long, involucre campanulate, 0.7 I mm. diameter, lobes triangular

exceeding glands, subentire or deeply laciniate, glabrous outside,

slightly hairy within, glands broadly transversely elliptic, 0.1 0.3

mm. long, appendages obsolete or to three times width of gland, white

or pink, fifth gland linear, shorter than lobes, sinus shallow. Staminate

flowers 5-14 per cyathium, androphores glabrous, 1 mm. long at maturity.

Gynophore glabrous, to 2 mm. long at maturity, exserted, upright or nutant;

calyx obsolete; ovary glabrous; styles spreading, 0.4 1 mm. long, minutely

joined at base, bifid for 1/3 1/2 length. Capsule glabrous, broadly

ovoid with somewhat flattened base, 1.8 2.3 mm. long, 2 2.3 mm. diameter







9r1
! r '-"

v-


I I ., I. Il I:.


Plate 12. CHAMAESYCE NUTANS (Lag.) Small

C. nutans (Lag.) Small, between Marianna and Campbellton, Jackson County,
Florida, 7 Nov. 1964, Godfrey 64932 (FLAS).








below equator, strongly three-lobed, angles subacute. Seed oblong-ovoid,

to 1.3 mm. long, 0.8 mm. wide at equator, ventral angle rounded and

obscure, others well marked, faces convex, rippled, black, often with

light angles.

A common weed of the eastern United States reaching its southern

limit in north Florida. It is possible that it was present as a weed

of cultivated ground in Puerto Rico and many of the Caribbean islands

as reported by Britton and Wilson (1924), but almost all of the speci-

mens in NY and F on which these records might have been based prove to

be misidentifications of plants of E. hyssopifolia L. and E. hypericifolia

L. The collections by Fredholm and by Hitchcock from central Florida are

correctly named, but no more recent records have been obtained from this

area in spite of heavy collecting, and it is doubtful that the species is

present as more than a short-lived, casual introduction. (Map 11).

Specimens seen: FLORIDA. Alachua: June-July 1898, Hitchcock 1697

(F). Brevard: Okeechobee region, 17 Sept. 1903, Fredholm 6014 (GH, NY).

Citrus: June-July 1898, Hitchcock 1699 (F, MO). Gadsden: Jim Woodruff

Dam, Chattahoochee, 19 Oct. 1963, Burch 207 (FLAS, IJ, US); River Junction,

7 Aug. 1927, Wiegand & Manning 1818 (GH); Chattahoochee, 7 Aug. 1927,

Wiegand & Manning 1819 (GH); Chattahoochee, 8 Sept. 1931, West s.n. (FLAS).

Jackson: Marianna, 19 Oct. 1963, Burch 210 (FLAS); near Cottondale, 26 Oct.

1964, Burch 504 (FLAS, MO, NCU, NY); between Marianna and Campbelltown,

7 Nov. 1964, Godfrey 64932 (FLAS, FSU); Marianna State Park, 15 Sept. 1960,

Mitchell 839 (FSU); Marianna Caverns State Park, 19 Nov. 1960, Mitchell

903 (FSU). Leon: Tallahassee, 11 Sept. 1942, Henry & Baggs s.n. (FLAS);

18 Sept. 1942, Kurz s.n. (FLAS); Tallahassee, 2-3 Sept. 1895, Nash 2519








(F, GH, NY). Levy: June-July 1898, Hitchcock 1698 (F). Wakulla: Newport,

13 Sept. 1964, Godfrey 64618 (FLAS). Walton: Northwest section, 9 July

1964, Hume s.n. (FLAS). Washington: west side of Careyville, 26 Oct. 1964,

Burch 499 (FLAS). Extra-territorial specimens: BERMUDA. North Shore, 31

Aug.-20 Sept. 1905, Brown & Britton 8 (F, NY); Devonshire, 31 Aug.-20 Sept.

1905, Brown & Britton 79 (F, NY); Devonshire, 19 Aug. 1963, Manuel 265 (A).

This species bears a strong resemblance to C. hypericifolia (L.)

Millsp., C. hyssopifolia (L.) Small and C. lasiocarpa (Klotzsch) Arth.,

but may be distinguished by its stem pubescence, limited to the tips and

lines at the side of older parts; by the cyathia carried in a leafy

dichasium; by the large capsules and by the black, plump seed with a

rippled surface.

The application by Wheeler (1939) of the name E. maculata L. to

this species, rather than to the prostrate plant for which it has been

widely used, deserves comment. His case, based on the selection of a

specimen from Herb. Linn. as the type, was never proved, since there

was some doubt about the identification that Linnaeus made for this sheet.

The description of E. maculata in Species Plantarum is placed among the low-

growing rather than the erect species, suggesting that Linnaeus intended

it for a prostrate plant and that the traditional application is correct.

The use of the name E. hypericifolia L. for this species may be

traced to annotations on the Linnaean specimens made by Sir J. E. Smith

during the years when he owned the herbarium. Michaux (1803) may have

examined the material and arrived at the decision independently, but the

use in this fashion was confirmed by correspondence between Smith and

Torrey when his Flora of the State of New York (1843) was in preparation.






\ i


2v- o
> -^




^^~~; '


P-~ ~ ~


Map 1!. Distribution of C. nutans


K!^ -o

s-~-"~-J


- ...-- ------ ------ -


j 6
-~o


Map 12. Distribution of C. hypericifolia


,


cm -~91cl~110


---------I ---- ------0








The basis for Smith's actions was sound in that the description of

E. hypericifolia in Linnaeus' early works was written from this northern

plant. The specific limits were widened to include a tropical plant,

however, before the publication of Species Plantarum, and since the name

has been used most widely for this entity, the interests of stability of

nomenclature are best served by maintaining this application. Burch (1965)

deals more fully with the questions relating to the name of this species.

9. CHAMAESYCE HYPERICIFOLIA (L.) Millsp. Field Mus. Bot. 2: 302.

1909. Euphorbia hypericifolia L., Sp. Pl. ed. 1, 454. 1753. Type:

Sloane Hist. 1, t. 126. (lectotype, designated by Burch, Rhodora 67. 1965,

in press).

E. pilulifera L. sensu Wheeler, Contr. Gray Herb. 127: 76. 1939.

(But see disclaimer, loc. cit. 78).

C. qlomerifera Millsp. Field Mus. Bot. 2: 377. 1913. Type: El

Rancho, Dept. Jalapa, Guatemala, 20 Jan. 1908, W. A. Kellerman 8053 (F i).

E. glomerifera (Millsp.) L. C. Wheeler, Contr. Gray Herb. 127: 78. 1939.

Annual or perennating in tropics; erect to ascending, or sprawling

to 4 dm. Stem to 3 mm. diameter at somewhat woody base, internodes to

3 cm. long; branching throughout length particularly in upper part of stem,

glabrous, tan to dark brown. Leaves membranous, blades ovate-elliptic to

elliptic-obovate, at times somewhat falcate, 15-35 x 8-14 mm., base inequi-

lateral, rounded or cuneate, margin serrate, apex acute, glabrous, green,

sometimes purple-spotted above and lighter green on underside; petiole

to I mm. long; stipules joined, sheathing, I 1.5 mm. long, margin serrate,

inner surface pubescent at least at apex, light-brown, or, at upper nodes,

separate, to 0.5 mm. long, entire or deeply 2-4 cleft. Cyathia in









short-stalked, lateral and terminal, congested dichasia forming glomerules

with a few basal leaves; involucre short-peduncled with stipule-like

bracts at junction with pedicel, narrow campanulate or more flaring, to

0.8 mm. diameter at mouth, lobes equalling or exceeding glands, triangular,

toothed or cleft, glabrous on outside, sparsely long-pilose within, glands

subcircular to 0.1 mm. diameter, appendages obsolete or prominent, to

1 mm. wide, entire, white or pink, fifth gland obsolete, sinus shallow.

Staminate flowers 10-12 or rarely only 4 per cyathium, androphores glabrous,

to 0.6 mm. long at maturity. Gynophore glabrous, to 1 mm. long at matu-

rity, exserted, nutant; calyx a triangular pad of tissue; ovary glabrous;

styles 0.3 mm. long, spreading, minutely joined at base, bifid for 1/2

length. Capsule glabrous, subspherical, to 1.2 mm. diameter, roundly

three-lobed, angles rounded. Seed ovoid, to 0.8 mm. long, 0.5 mm. wide,

strongly four-sided, ventral angle more obtuse than others, faces flat

or convex, wrinkled, light brown.

A frequent plant of waste places in Old and New World tropics and

subtropics. Common in the southern part of Florida with isolated collec-

tions as far north as Sapelo Island, Georgia, and present throughout the

Greater and Lesser Antilles (Map 12).

Representative specimens seen: FLORIDA. Alachua: University of

Florida campus, Gainesville, 30 Oct. 1964, Burch 521 (FLAS). Broward:

Fort Lauderdale, 19-25 Nov. 1903, Small & Carter 644 (NY). Collier: road

to Marco Island, 15 Apr. 1954, West, Arnold & Cooley 147 (FLAS). Dade:

vicinity of Homestead, 22 Nov. 1964, Godfrey 65583 (FLAS, FSU); Buena

Vista, I Jan. 1930, Moldenke 330 (DUKE, NY); Miami, 14 May 1904, Tracy

9127 (F, MO, NY). Glades: Caloosahatchee River bank, 7 Nov. 1964,











a I a 3 t 6 7 p u a ,J ll II 1 II IS






a- 4


\"4











Plate 13. CHAMAESYCE HYPERICIFOLIA (L.) Millsp.

C. hypericifolia (L.) Millsp., University of Florida campus, Gainesville,
Alachua County, Florida, 30 Oct. 1964, Burch 521 (FLAS).









Hodgson 230 (FLAS). Highlands: Rd. 25, south of Sebring, 22 June 1948,

Garrett s.n. (FLAS). Lee: Sanibel Island, 13 Apr. 1954, West & Cooley

2630 (USF). Manatee: U.S. 41 & U.S. 19, 29 Nov. 1962, Ward & Burch 3163

(FLAS). Monroe: Upper Mateconbe Key, June, ?? Curtiss 2486 (F, MO);

No Name Key, 21 Mar. 1898, Pollard, Collins & Morris 126 (F, NY); Key

Vaca, 3 Sept. 1954, Thorne 15177 (IJ); Key Largo, north end, 27 June 1960,

Webster & Samuel 10239 (DUKE, PUR). Palm Beach: Jupiter Inlet, west of

DuBoise Park, 27 Aug. 1964, Christensen RC-44a (FLAS, FSU). Pinellas:

Don Caesar Hotel, St. Petersburg Beach, 24 May 1964, Taylor s.n. (FLAS).

Polk: Haines City, 25 June 1963, Conard s.n. (FSU). Sarasota: Laurel,

21 Nov. 1964, Godfrey 65273 (FLAS, FSU). Sumter: Wildwood exit, 1-75,

18 Nov. 1964, Burch 524 (FLAS). Volusia: Pierson, 17 Sept. 1943, West

& Arnold s.n. (FLAS). BAHAMAS. Great Bahama, 5-13 Feb. 1905, Britton &

Millspaugh 2476 (F, NY); North Bimini, 16 Apr. 1904, Millspaugh 2399 (F,

NY). Andros: Mastic Point, 19-28 Mar. 1907, Brace 7087 (F, NY); Nicholl's

Town, 4-5 Feb. 1910, Small & Carter 8937 (F, NY). New Providence, Nassau,

Nov. 1890, Hitchcock s.n. (F, MO). Eleuthera: Gregory Town, 6 July 1903,

Coker 371 (NY); Governor's Harbor, 14 Nov. 1890, Hitchcock s.n. (F). Cat

Island, the Bight & vicinity, 1-6 Mar. 1907, Britton & Millspaugh 5816

(F, NY); Watling's Island, Cockburn Town, 12-13 Mar. 1907, Britton &

Millspaugh 6053 (F, NY); Great Exuma, 22-28 Feb. 1905, Britton & Millspaugh

2945 (F, NY); Long Cay, 7-17 Dec. 1905, Brace 4036 (F, NY); Inagua, 3 Dec.

1890, Hitchcock s.n. (F, MO). CUBA. Camaguey: Atalaya, 20 Mar. 1909,

Shafer 983 (F, NY). Havana: Havana, 5 May 1905, Curtiss 743 (F, MO, NY).

Oriented: Monte Verdi, Jan.-July 1859, Wright 1422 (F, MO, NY). Santa Clara:

Cienequita, Cienfuegos, 2 May 1895, Combs 4 (F, MO, NY); Limones, Soledad,








Cienfuegos, 27 Sept. 1924, Jack 6334 (MO). GRAND CAYMAN. Georgetown,

2 June 1963, Crosby, Hespenheide, & Anderson 50 (DUKE, UCWI); Spot Bay,

13-14 Feb. 1899, Millspaugh 1304 (F). JAMAICA. Portland: Kildare, 21

Aug. 1962, Adams 11555 (UCWI); Port Antonio, 4 Jan. 1906, Wight 2 (F,

NY). St. Andrew: Mount Pleasant, Stony Hill, 21 July 1912, Harris

11134 (F, NY, UCWI); Long Mountain, June 1953, Robbins 434 (UCWI).

St. Ann: Mosely Hill Cave, 1 mile west of Blackstonedge, 12 Dec. 1952,

Proctor 7474 (IJ). St. James: Ironshore, May 1953, Asprey 432 (UCWI).

Trelawny: Cowie Park near Troy, 19 Oct. 1917, Harris 12646 (F, MO, NY,

UCWI). Westmoreland: Negril, 29 Mar. 1962, Adams 10980 (UCWI). HAITI.

Jeremie, 13 July 1941, Seibert 1725 (MO); Port au Prince, 21-23 Feb. 1920,

Leonard 2778 (NY); St. Michel de 1'Abalaye, 1927, Ekman 8394 (F, IJ).

DOMINICAN REPUBLIC. Barahona, June 1910, Fuertes 255 (F, MO, NY); San

Pedro de Macoris, 26 Mar. 1913, Rose, Fitch & Russell 4154 (NY); Santo

Domingo City, 9-10 Mar. 1913, Rose, Fitch & Russell 3769 (NY). PUERTO

RICO. Camuey, 14 June-22 July 1901, Underwood & Griggs 204 (NY);

Fajardo Light House, 17 Apr. 1899, Heller 1179 (F, NY); Guanica, 22

Jan. 1899, Millspaugh 753 (F); Guayama, May 1874, Kuntze 554a (NY);

Penuelas, 16 July 1886, Sintenis 4823 (MO, NY); Rio Piedras, 9 Apr. 1912,

Johnston 216 (NY); San Juan, Dec. 1913, Hiorani s.n. (NY). LESSER ANTILLES.

St. Thomas: Charlotte Amalie, 17-18 Jan. 1899, Millspaugh 382 (F); Forts

Square, Jan. 1877, Eggers s.n. (MO, NY). St. Croix: Bassin Yard, 15 Nov.

1895, Ricksecker 75 (F, MO, NY); Peter's Rest, 2 June 1924, Thompson

772 (NY). Tortola, 18 Sept. 1918, Fishlock 144 (NY). St. Kitts, 8 Sept.-

5 Oct. 1901, Britton & Cowell (NY). Guadeloupe, Pointe-a-Pitre, 19 May

1936, Stehle 1190 (NY); Dominica, Soufriere, 1903, Lloyd 490 (NY).









Martinique: Case Pilote, 1879, Duss 488 (F, MO, NY); Ste. Anne village,

29 July 1939, Egler 39-208 (NY). Barbados: Bathsheba, Apr.-June, 1895,

Waby 101 (F); Waterford, St. Michael, 15 Aug. 1906, Dash 166 (F).

Grenada, St. George's, 29 Sept. 1906, Broadway s.n. (F). Extra-territorial

specimens: BERMUDA. St. David's Isle, 22 May-6 June 1914, Brown,

Britton & Bisset 2093 (F, NY); neglected field, 7 July 1921, Degener 1003

(MO). GEORGIA. S.W. section, Sapelo Island, 13 Oct. 1956, Duncan 20624

(DUKE, NCU, NSC).

The Linnaean name E. hypericifolia has been applied to this species

and to the one known here as C. nutans. Both elements were represented

in the diagnosis and in the synonymy of the original publication in Species

Plantarum, and the choice of this species to carry the name was made on

the basis of its widespread application in this way dating from Miller's

Gardener's Dictionary edition 8 (1768). This matter, and Wheeler's appli-

cation of the name to the plant known here as C. lasiocarpa, are discussed

in more detail by Burch (1965).

The use of the name E. pilulifera L. for this species cannot be

accepted if the sheet (630-8) in the Linnaean Herbarium bearing this name

is allowed any status as type material. The identity of the specimen is

not clear, but the small part of a seed visible in one capsule showed none

of the surface markings characteristic of seed of C. hypericifolia, and

the general aspect of the plant was closer to that of C. hirta.

10. CHAMAESYCE HYSSOPIFOLIA (L.) Small,J. N.Y. Bot. Gard. 3: 429.

1905. Euphorbia hyssopifolia L. Syst. Nat. ed 10, 1048. 1759. Type: in

Herb. Linn. sheet 630-9 Anisophyllum hyssopifolium (L.) Haw., Syn. PI.

Succ., 161. 1812. E. hypericifolia L."P hyssopifolia L." Griseb. Fl. Br.








W. Ind. Isl., 54. 1859. E. brasiliensis Lam. var. hyssopifolia Boiss.

in DC. Prod. 15(2): 24. 1862.

E. brasiliensis Lam., Encyc. 2: 423. 1786. Type: Mus. Nat.,

Paris, if extant. C. brasiliensis (Lam.) Small, Fl. SE, US., 712. 1903.

C. nirurioides Millsp., Field Mus. Bot. 2: 394. 1914. Type: Malvern,

Santa Cruz Mountains, Jamaica, 5 Sept. 1907, Britton 1186 holotypee: F !,

isotype: NY !). E. nirurioides (Millsp.) Fawcett & Rendle, Fl. Jam., 4(2):

339. 1920.

C. jenningsii Millsp. in Britton, Bull. Torr. Bot. Club 43; 465.

1916. Type: near Vivijagua, Isle of Pines, Cuba, 7 May 1910, Jennings 621

(Holotype: Herb. Carnegie Mus., portion of holotype F !; Isotype: NY 1).

Annual, occasionally perennating; erect or ascending or rarely

decumbent, to 6 dm. Stem to 4 mm. thick at somewhat woody base, internodes

to 3 cm. long; branched at all levels but most in upper part of plant,

glabrous or sparingly long hairy near base, rarely strongly short-pubescent

at lower nodes particularly in plants from more tropical areas, straw-

colored or darker brown. Leaves membranous; blades very variable, linear-

lanceolate, 15-35 x 3-6 mm., with subcordate inequilateral base, margin

serrate in upper half, apex acute, to ovate-elliptic, occasionally some-

what falcate, 10-30 x 6-10 mm., with rounded to subcordate inequilateral

base, margin generally serrate, apex.obtuse, glabrous or sparsely long

pilose, on shoots near base of plant, both surfaces mid-green; petiole 1-2

mm. long; stipules joined or nearly free at upper nodes, almost obsolete to

deltoid, 0.8 mm. long, margin short-fringed. Cyathia borne singly,

terminal and axilary in short-stalked, diffuse or compact, leafy dichasia;

peduncle short, involucre commonly narrowly tubular, 0.7 1 mm. diameter,









sometimes flaring-campanulate, lobes triangular, deep cleft, exceeding

glands, glabrous outside, sparsely pilose within, glands transversely

elliptic or almost circular, 0.1 0.3 mm. long, appendages obsolete to

prominent, elliptic, twice as long and three times as wide as gland,

white or pink, fifth gland obsolete or tiny, linear, sinus shallow.

Staminate flowers 9-20 per cyathium, androphores glabrous, 0.6 1 mm.

long. Gynophore glabrous, ca. 1 mm. long at maturity, exserted, usually

nutant; calyx a triangular pad; ovary glabrous; styles upright or spread-

ing, to 0.4 mm. minutely joined at base, bifid for 1/3 1/2 length.

Capsule glabrous, truncate-cuneiform to broadly ovoid, 1.5 2 mm. long,

1.6 1.8 mm. wide just above base, strongly three-lobed, angles subacute.

Seed oblong-ovate, to 1 mm. long, 0.5 mm. wide, four-angled with ventral

angle less acute than others, faces flat, marked with transverse ridges,

a central longitudinal ridge sometimes on dorsal faces, brown or black,

angles sometimes light.

Widespread throughout Florida and sporadically to South Carolina,

common in all parts of the Caribbean except the Bahamas, where the reports

for New Providence and Eleuthera (Britton and Millspaugh, 1920) are

probably based on collections by Britton and Coker (NY !) which prove to

be misidentifications of plants of C. hypericifolia (L.) Millsp. (Map 13).

Representative specimens seen: FLORIDA. Alachua: Pineland, Gaines-

ville, 15 June 1910, Hood 24465 (FLAS, MO); University of Florida campus,

Gainesville, 10 Sept. 1956, Jackson 707 (FLAS). Baker: Olustee National

Forest, 28 Nov. 1955, Kral 1572 (FSU). Brevard: Melbourne, 22 Aug. 1958,

Kral 7971 (FLAS, GH, NY, USF). Broward: Pinelands, Ft. Lauderdale, 19 &

25 Nov. 1903, Small & Carter 1141 (F, NY); U.S. 27, north of Andytown,









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Plate 14. CHAMAESYCE HYSSOPIFOLIA (L.) Small

C. hyssopifolia (L.) Small, Fairchild Tropical Gardens, Dade County,
Florida, 31 December 1962, Burch 52 (FLAS).


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Plate 15. CHAMAESYCE HYSSOPIFOLIA (L.) Small

Isotype: C. nirurioides Millsp., Malvern, Santa Cruz Mountains, Jamaica;
5 Sept. 1907, Britton 1186 (NY).


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PLANTS OF ISI.I: o01 PINE:

Plate 16. CHAMAESYCE HYSSOPIFOLIA (L.) Small
Isotype: C. ienninqsii Millsp. in Britton, near Vivijagua, Isle of Pines
Cuba, 7 May 1910, Jennings 621 (NY).


I









26 Jan. 1963, Ward & Burch 3338 (FLAS). Calhoun: N.W. of Blountstown,

25 Oct. 1964, Burch 506 (FLAS). Charlotte: Charlotte Harbor, 21 Nov.

1964, Godfrey 65360 (FLAS); Peace River, 13 Aug. 1963, Burch 185 (FLAS).

Citrus: Dunnellon, 16 June 1958, Kral 6901 (FLAS, FSU, USF). Collier:

Immokalee, 26 Sept. 1964, Lakela 27492 (USF). Dade: Buena Vista, 1 Jan.

1930, Moldenke (DUKE, F, MO, NY); Miami-Kendall Station, 5 Nov. 1906,

Small & Carter 2521 (NY). Ouval: Jacksonville, 17 Nov. 1929, Moldenke

5232 (NY). Franklin: Apalachicola, 2 Nov. 1875, Chapman 3844a (MO, NY).

Gadsden: Greensboro, 9 July 1955, Godfrey 53614 (DUKE, FSU, NY). Hendry:

La Belle, 19 Apr. 1930, Moldenke 5911 (NY). Hernando: Brooksville,

3 Sept. 1959, Ray 9461 (USF). Highlands: Childs, 3 Mar. 1945, Brass

14744 (US); Sebring, 25 Apr. 1960, Ray, Gleason & Eaton 9728 (USF).

Hillsborough: Tampa, 20 Mar. 1923, Churchill 546 (MO, US); Oldsmar,

12 Aug. 1960, Ray, Lakela & Patman 10006 (USF). Jackson: between Mari-

anna & Campbellton, 7 Nov. 1964, Godfrey 64933 (FSU). Lake: Eustis,

16-30 June 1894, Nash 1076 (F, MO, NY). Lee: Ft. Myers, July-Aug. 1900,

Hitchcock 324 (F, MO, NY); 5 miles east of Ft. Myers Beach, 1 Aug. 1958,

Kral 7539 (GH, USF). Leon: Tallahassee, 30 June 1955, Godfrey 53571

(DUKE, FSU, NY). Marion: Silver Springs, 1 Oct. 1963, Godfrey 63145 (FSU).

Manatee: Bradenton, 13 June 1916, Cuthbert 1312 (NY); Cortez, 19 July 1958,

Jackson 732 (FLAS). Martin: Stuart, 3 Sept. 1944, West & Arnold s.n.

(FLAS). Monroe: Key Largo, 26 Jan. 1963, Ward & Burch 3332 (FLAS). Nassau:

Fort Clinch, 12 Oct. 1963, Burch 195 (FLAS). Okaloosa: Ft. Walton Beach,

24 Oct. 1964, Burch 470 (FLAS). Okeechobee, 27 Dec. 1964, Burch 589 (FLAS).

Orange: Orlando, 21 June 1900, Curtiss 6670 (MO, NY). Osceola: St. Cloud,

27 Feb. 1952, Cooley 1032 (USF). Palm Beach: between Palm Beach & Lake








Worth, 7 Aug. 1963, Burch 119 (FLAS). Pasco: Land O'Lakes, 10 July

1958, (FLAS, GH, NY). Pinellas: St. Petersburg, 23 Sept. 1907, Deam

2758 (MO). Polk: Lake Hamilton, 30 Sept. 1962, Conard s.n. (FLAS).

Putnam: Welaka, 5 Aug. 1940, Laessle s.n. (FLAS). St. Lucie: Ft.

Pierce, 7 Aug. 1963, Burch 122 (FLAS). St. Johns: Ponte Vedra, 11

Oct. 1964, Burch 437 (FLAS). Sarasota: Osprey, 10 Mar. 1904, Smith

712 (DUKE). Sumter: Withlacoochee River bridge, 8 Nov. 1946, West &

Arnold s.n. (FLAS). Suwannee: Live Oak, 4 Oct. 1964, Elam s.n. (FLAS).

Volusia: Daytona Beach Shores, 3 Aug. 1962, Ray 11159 (USF). Wakulla:

3 Nov. 1964, Godfrey 64887 (FLAS, FSU). Washington: Chipley, 31 July

1954, Ford & Arnold 3860 (FLAS). CUBA. Camaguey: La Gloria, Feb. 1909,

Shafer 285 (F, NY). Cienfuegos: Colonia Limones, Ingenio Soledad, 24 Jan.

1903, Pringle 59 (F, MO, NY); 2 May 1895, Combs 3 (F, MO, NY). Havana:

Guanabacoa, 27 Jan. 1905, Curtiss 618 (F, MO, NY); Havana, Apr. 1903,

Shafer 91 (F). Isle of Pines: Nueva'Gerona, 19 Dec. 1903, Curtiss 244

(F, NY). Oriented: Bayate, 18 July 1914, Ekman 2012 (NY). Pinar del Rio:

Herradura, 26-30 Aug. 1910, Britton, Britton, Earle & Gager 6332 (F, NY);

between Vinales and Pinar del Rio, 16 June 1953, Webster 4670 (PUR). San

Pedro: banks of Ariguanaba lagoon, 5 Apr. 1923, Leon 11417 (NY). Santa

Clara: Hoyo de Manicaragua, 26-28 Feb. 1910, Britton, Britton & Wilson

4660 (F, NY); Rio Hondo Plains, 5 May 1903, Cook & Doyle 528 (F). JAMAICA.

Clarendon: Alley, 11 Feb. 1947, von der Porten s.n. (IJ); Kupuis, 23 June

1955, Proctor 10330 (IJ). Hanover: Negril to Bloody Bay, 12 Sept. 1962,

Adams 11663 (UCWI). Manchester: Lititz, 5 Nov. 1961, Adams 9864 (UCWI).

Portland: Port Antonio, Dec. 1890, Hitchcock s.n. (MO). St. Andrew: Con-

stant Spring to Bardowie, 2 Aug. 1915, Harris 12112 (F, MO, NY, UCWI);









Hope, 18 Aug. 1897, Harris 6803 (F, NY, UCWI); road to Constitution

Hill, 20 Oct. 1905, Harris 9035 (F, NY, UCWI); University of West Indies

campus, 4 June 1963, Crosby, Hespenheide & Anderson 60 (F). St. Catherine:

Bog Walk, 17 Dec. 1890, Hitchcock s.n. (MO). St. Thomas: Morant Bay, 6

Dec. 1959, Adams 5678 (UCWI); Port Morant, 24 Dec. 1890, Hitchcock s.n.

(MO). Westmoreland: Bluefields & vicinity, 6-7 Mar. 1908, Britton &

Hollick 1943 (F, NY). HAITI. Gonaives, Dec. 1899, Buch 278 (IJ); La

Gonave, 9 Aug. 1927, Ekman 8862 (A); Mt. La Porte, near Pilate, 21 Aug.

1903, Nash 628 (NY); Pointe-a-pitre, 12 July 1936, Stehle 497 (NY); Port

de Paix, 9 Jan. 1929, Leonard & Leonard 13543 (NY); St. Michel de l'Atalaye,

1 Jan. 1926, Leonard 8501 (MO); San Michel, 5 Aug. 1905, Nash & Taylor

1417 (NY). DOMINICAN REPUBLIC. Barahona, Feb. 1910, Fuertes 149 (F, NY).

San Juan: Rio Arriba, 9-14 Sept. 1946, Howard & Howard 8821 (GH, NY).

Santo Domingo: Villa Allagracia, 19 Jan. 1929, Ekman H11229 (PUR). PUERTO

RICO. Guayanilla-Tallahoa, 13 Mar. 1913, Shafer 1976 (F, MO); Martin

Pena Station, 20 Feb. 1900, Heller 4665 (F, MO); Punta Arenas, 15 Feb.

1914, Shafer 2896 (F, NY); Rio Piedras, Station Grounds, 2 Oct. 1937,

Otero 123 (MO); Utuado, 15-20 Mar. 1906, Britton & Cowell 409 (F, NY).

LESSER ANTILLES. Charlotte Amalie, 17-18 Jan. 1899, Millspaugh 398 (F);

Antigua, Belvidere, 15 Aug. 1937, Box 964 (MO); Montserrat, 5 Aug. 1939,

Friend 51 (NY); Guadeloupe, Manive, 1893, Duss 2920 (NY); Martinique,

Case-Pilote, 1901, Duss 4641 (F, NY); St. Lucia, 7 Sept. 1935, Potter

5508 (GH); St. Vincent, Kingstown, 2 Dec. 1945, Beard 1397 (MO, NY);

Grenada, St. George's, 30 Oct.-ll Dec. 1957, Proctor 16812 (IJ).

C. hyssopifolia (L.) Small may be distinguished from similar species

by its combination of usually glabrous stems, cyathia in a leafy dichasium,














<\


S-------- --

------0-- -

-0- -- --


Map 13. Distribution of C. hvssopifolia


Map 14. Distribution of C. lasiocarpa








large capsules and seed with prominent transverse ridges on at least

the tangential faces.

The specimen in the Linnaean Herbarium is of a plant with linear

leaves and rather few cyathia in a diffuse dichasium. This form occurs

in most of the islands of the Greater Antilles, and is particularly well-

marked on Hispaniola, but grades into the broader leaf form (often on the

same plant), and the type specimen represents only the extreme leaf shape

of this very variable species.

11. CHAMAESYCE LASIOCARPA (Klotzsch) Arthur, Torreya 11: 260.

1911. Euphorbia lasiocarpa Klotzsch, Nov. Act. Nat. Cur. 19, supp. 1:

414. 1843. Type: Bot. Mus., Berlin, if extant. E. hypericifolia L.

" lasiocarpa K1." Griseb., Fl. Brit. W. I. Isl. 54. 1859. (excl. syn.

Balb.).

E. hypericifolia L. sensu Wheeler, Contr. Gray Herb. 127: 73. 1939.

Annual; erect or ascending to 1 m. Stem to 5 mm. thick at somewhat

woody base, internodes to 3 cm. long; branched throughout, densely tomen-

tose, straw-colored becoming darker with age. Leaves membranous; blades

ovate-lanceolate to obovate, somewhat falcate, 15-40 x 8-20 mm., base

inequilateral, rounded to obtuse, margin sharp-serrate, apex acute to ob-

tuse, densely gray pubescent, rarely glabrate, green; petioles 2 to 3 mm.

long; stipules joined, deltoid to 1 mm., toothed or ciliate, sometimes

deeply bifid near tips of shoot. Cythia solitary in lateral and terminal

short-stalked compound dichasia; peduncle ca. 1 mm. long, involucre

campanulate, 1.2 mm. diameter at mouth, lobes triangular equalling glands,

toothed or cleft, densely pubescent on outside, pilose within, glands trans-

versely elliptic 0.3 mm. x 0.1 mm., appendages lunate, to three times as








long and broad as gland, white, fifth gland deltoid, short or obsolete,

sinus to one fourth depth of involucre. Staminate flowers ten to eighteen

per cyathium, androphores glabrous, to 1 mm. long at maturity. Gynophore

pilose to 1.5 mm. long at maturity, exserted, nutant; calyx a triangular

pad of tissue; ovary strigose; styles upright, to 0.7 mm. long, joined at

base, bifid for 1/3 length. Capsule densely pilose, broadly ovoid or

conical, 1.5 2 mm. long, 2 mm. wide below equator, broadly three-lobed,

angles acute. Seed oblong-ovoid, to 1 mm. long, 0.6 mm. wide, strongly

four-angled, ventral angle more obtuse and rounded than other three, faces

flat, with 2-4 deep transverse ridges, dorsal faces with an additional

central longitudinal ridge, black, often with light angles.

A common roadside weed in Jamaica and Haiti, less common in Puerto

Rico. Collections have been seen from Barbados and Tobago, and the plant

is reported from Central America and northern South America (Wheeler,

1939) (Map 14).

Specimens seen: JAMAICA. St. Andrew: Kingston, 29 Aug. 1908,

Britton 2978 (NY); Irish Town, 17 June 1963, Crosby, Hespenheide & Anderson

186 (DUKE, PUR); Liguanea, 7 Feb. 1960, Proctor 20564 (IJ, NY, PUR); Stony

Hill, 21 July 1912, Harris 11136 (NY, UCWI). St. Thomas: Easington

Bridge, 23 Mar. 1954, Proctor 8512 (IJ). Manchester: Warwick, 8 Nov. 1962,

Proctor 22886 (IJ). St. Elizabeth: Unity Hill, Aug. 1961, Robertson 9136

(UCWI). HAITI. Port-au-Prince, Sept. 1919, Buch 1713 (IJ); St. Marc,

25-28 Feb. 1920, Leonard 2954 (NY); Petionville, 15-28 June 1920, Leonard

4967 (NY); St. Michel de l'Atalaye, 26 Dec. 1925, Leonard 8483 (MO); Jean

Rabel, 27 Jan.-9 Feb. 1929, Leonard & Leonard 12789 (NY). PUERTO RICO. Coamo,

23 Nov. 1899, Goll 756 (NY); Juana Diaz, 3 Nov. 1913, Hess 4002 (NY).









LESSER ANTILLES. Barbados, 30 July 1901, Freeman s.n. (UCWI); Tobago,

6 Feb. 1913, Broadway 4415 (GH, MO).

This species is distinguishable by the dense pubescence of its

stem and leaves, which even the most heavily pubescent specimens of C.

hyssopifolia do not approach. In most other characters there is a

close resemblance between the two.

Wheeler's application of the name Euphorbia hypericifolia L. to

this species was based on a misidentified specimen in the Linnaean

Herbarium, which was not seen or annotated by Linnaeus until long after

the publication of the name.

12. CHAMAESYCE PARCIFLORA (URB.) comb. nov. Euphorbia parciflora

Urban in Fedde, Repert. 15: 411. 1919. Type: prope Marmelade, Haiti,

1-2 Aug. 1905, G. V. Nash & N. Taylor 1269 (in Herb. Bot. Mus. Berlin,

if extant; Isotype: NY 1).

Perennial; erect or ascending to 3 dm. Stem to 2 mm. diameter,

internodes to 3 mm. long; branched sparsely throughout length, glabrous

or sparingly short-tomentose on one surface, dark brown. Leaves membranous;

blades ovate, 6-10 x 3-5 mm., base inequilateral, rounded, margin sharply

serrate, apex acute to obtuse, midrib distinct, glabrous, olive-green;

petiole 1 mm. long; stipules joined, triangular to 1 mm. long, toothed

or deeply cleft, sometimes bifid and almost distinct. Cyathia solitary

at upper nodes; peduncle slender, to 3 mm. long, involucre campanulate,

to 1.8 mm. diameter at mouth, lobes barely exceeding glands, attenuate,

glabrous on outside, densely ciliate within, glands transversely elliptic,

0.6 x 0.1 mm., cupped, appendages as long as gland and up to four times

as wide, creamy white, margin entire or crenulate, fifth gland linear,




89.



--- -








;---r


------------










.M.a.p 52.-... -.: f C -- -
ap ^ ] st i ut on of C p rc fl r


Map 16. Distribution of C. brittonii








almost equalling lobes, sinus shallow. Staminate flowers 4-12 per

cyathium, androphores glabrous, to 1.3 mm. long at maturity. Gynophore

glabrous, to 2.5 mm. long at maturity, exserted, nutant; calyx obsolete

or a subcircular pad of tissue; ovary glabrous; styles spreading, 0.5 mm.

long, joined at base, bifid for 1/2 length. Capsules glabrous, broadly

ovoid, 2 2.2 mm. long, 2.5 2.8 mm. wide above base, narrowly three-

lobed, angles acute. Seed ovate, to 1.3 mm. long, 1 mm. diameter just

above base, four-angled, dorsal faces larger than ventral, dorsal angle

acute, ventral angle flattened, faces flat, wrinkled, white.

Reported only from the mountains of Haiti, at elevations of 500-

2000 m. in pineland or open woods (Map 15).

Specimens seen: HAITI. Badeau, 21 Feb. 1927, Ekman H4968 (IJ);

Gonaives, June 1901, Buch 634 (IJ); M. Bonfiere, 29 Sept. 1925, Ekman

H7627 (IJ); Petionville, 24 Aug. 1924, Ekman H1593 (IJ); San Michel to

Marmelade, 6 Aug. 1905, Nash & Taylor 1466 (NY).

The affinities of this species are not clear. Some of the extreme

forms of C. hyssopifolia in Hispaniola have similar cyathia but are narrow-

leaved, and plants growing strongly usually develop lateral dichasia of

cyathia as they get older.

13. CHAMAESYCE BRITTONII (Millsp.) Millsp., Field Mus. Bot. 2:

303. 1909. Euphorbia brittonii Millsp., Field Mus. Bot. 2: 159. 1906.

Type: Race course, New Providence, 26 Aug. 1904, Britton & Brace 839

(Lectotype: F !; Isotype: NY .).

Chamaesyce paucipila (Urb.) Millsp., Field Mus. Bot. 2: 410. 1916.

Euphorbia paucipila Urban, Symb. Ant., 5: 389. 1908. Type: Prope

Guanabacoa, Habana, Cuba, August, Baker and Hasselbring 7199 (Holotype:




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