A TAXONOMIC REVISION OF THE GENUS

CHAMAESYCE (EUPHORBIACEAE) IN

THE CARIBBEAN

By
DEREK GEORGE BURCH

A DISSERTATION PRESENTED TO THE GRADUATE COUNQL OF

THE UNIVERSITY OF FLORIDA

IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE

DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA
August, 1%5

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. Stearn of the British Museum (Natural History) and Dr.
R. K. Rechinger of the Naturhi stor i skmuseum, 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 1 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

Page

ACKNOWLEDGEMENTS ii

LIST OF MAPS vl

LIST OF PLATES vili

INTRODUCTION 1,

GENERAL DISCUSSION k.

I. HISTORY k.

II, THE SEGREGATION OF THE GENUS CHAMAESYCE 10.

III. MORPHOLOGY \k.

SYSTEMATIC TREATMENT 21.

Chamaesyce 21 .

Key to the species of Chamaesyce in the Caribbean 23.

1 . Chamaesyce vaq i nul ata 29.

2. Chamaesyce a rt i cu lata 33.

3. Chamaesyce buxi fol ia 36.

4. Chamaesyce lecheo ides A-2.

5. Chamaesyce cayensis 51.

6. Chamaesyce porter iana 5^.

7. Chamaesyce myrt i 1 1 i fol ia 63.

8. Chamaesyce nutans 65.

9. Chamaesyce hyper ici fol ia 71.

TABLE OF CONTENTS, Continued

SYSTEMATIC TREATMENT, Continued Page
Chamaesyce

10. Chamaesyce hyssop! fol ia 76.

11. Chamaesyce lasiocarpa 86.

1 2 . Chamaesyce pare! f lora 88 ,

13. Chamaesyce brittonii 90.

14. Chamaesyce cowe 1 1 i i 93.

15. Chamaesyce blodqettii 97.

16. Chamaesyce serpens 107.

17. Chamaesyce centunculoides 111.

18. Chamaesyce torralbasii 117.

19. Chamaesyce camaguayens i s 121.

20. Chamaesyce cord i fol ia ]2k.

21. Chamaesyce pol ygon i fo 1 ia 128.

22. Chamaesyce ammannioides I30.

23. Chamaesyce cumul icola 13^.

24. Chamaesyce prostrata I38.

25. Chamaesyce mendezi i 144,

26. Chamaesyce hepat ica 148.

27. Chamaesyce hel wig! i 150.

28. Chamaesyce h i rta 152.

29. Chamaesyce opthalmica I6O.

30. Chamaesyce berter iana I65.

3 1 . Chamaesyce maculata 1 68 .

TABLE OF CONTENTS, Continued

SYSTEMATIC TREATMENT, Continued Page
Chamaesyce

32. Chamaesyce thymi folia 175.

33. Chamaesyce adenoptera I80.

34. Chamaesyce conferta 190.

35. Chamaesyce garberi 193.

36. Chamaesyce mul t i nodi s 203.

37. Chamaesyce minutula 205.

38. Chamaesyce turp i ni i 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

Page

Map 1. Distribution of _C. vagi nulata 32.

Map 2. Distribution of _C. buxi fol ia 32.

Map 3. Distribution of _C. lecheoides var, wi 1 soni i kS.

Map 4. Distribution of _C. 1 echeoides var. lecheoides 49.

Map 5. Distribution of _C. lecheoides var. exumens i s 50.

Map 6. Distribution of _C. cayens i s 50.

Map 7. Distribution of _C, porter iana var. keyens is 6l.

Map 8. Distribution of _C. porter iana var. porter iana 61.

Map 9. Distribution of C^. porter iana var. scopar ia 62.

Map 10. Distribution of _C. art iculata and _C. myrt i I 1 i fol ia 62.

Map 11. Distribution of _C. nutans 70.

Map 12. Distribution of _C. hyper ici fol ia 70.

Map 13. Distribution of _C. hyssop i fol ia 85.

Map 14, Distribution of _C. las iocarpa 85.

Map 15. Distribution of _C. parci flora 89.

Map 16. Distribution of _C. bri ttoni i 89.

Map 17. Distribution of C_. cowel 1 i i 96,

Map 18. Distribution of _C. blodgetti i 96.

Map 19. Distribution of _C. serpens 112.

Map 20. Distribution of _C. centunculoides 112.

Map 21. Distribution of _C. torral basi i 120.

Map 22. Distribution of _C. camaguayens i s 120,

LIST OF MAPS, Continued

Page

Map 23. Distribution of _C. cordifol ia 127,

Map 24. Distribution of _C. polyqonifol ia 127.

Map 25. Distribution of _C. ammannioides I35.

Map 26. Distribution of _C. cumul icola 135,

Map 27. Distribution of _C. prostrata 143.

Map 28. Distribution of _C. mendezi i I43.

Map 29. Distribution of _C, hi rta 159.

Map 30. Distribution of C. opthalmica I59,

Map 31. Distribution of _C. berteriana I67.

Map 32, Distribution of _C. macuiata I67,

Map 33. Distribution of _C. helwiqi i and _C. paredonensis 179.

Map 34. Distribution of _C. thymifo] ia I79.

Map 35. Distribution of _C. adenoptera ssp. adenoptera 188.

Map 36. Distribution of _C. adenoptera ssp. pergamena 188.

Map 37. Distribution of _C. adenoptera ssp. qundlachi i I89,

Map 38. Distribution of _C. conferta ■ I89.

Map 39. Distribution of _C. garber i 202.

Map 40. Distribution of C. turpini i • 202.

Map 4]. Distribution of _C. pinetorum 215.

Map 42. Distribution of _C. del to idea ssp. serpyl 1 um 215.

Map 43. Distribution of _C. del toidea ssp. del to idea

var. del toidea 223,
Map 44. Distribution of _C. del toidea ssp. del toidea

var. adhaerens 223.

LIST OF PLATES

Plate 1. CHAMAESYCE VAGINULATA (Griseb.) Mi lisp.

Plate 2. CHAMAESYCE ARTiCULATA (Aubl.) Britton

Plate 3. CHAMAESYCE BUXIFOLIA (Lam.) Small

Plated. C. LECHEOIDES (Millsp.) Millsp. var. WILSONli
(Mi 1 1 sp.) i ned,

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

Plate 6. C. LECHEOIDZS (Millsp.) Millsp. var. EXUMENSIS
(Mi 1 1 sp. ) ined.

Plate 7. CHAMAESYCE CAYENSIS (Millsp.) Millsp.

Plate 8. CHAMAESYCE PORTERIANA Small var. KEYENSIS (Small)
i ned.

Plate 9. CHAMAESYCE PORTERIANA Small var. PORTERIANA

Plate

Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate

0. CHAMAESYCE PORTERIANA Small var. SCOPARIA (Small)

I ned.

1. CHAMAESYCE MYRTI LLI FOL I A (L.) Millsp.

2. CHAMAESYCE NUTANS (Lag.) Small

3. CHAMAESYCE HYPERI C I FOLI A (L.) Millsp.
k. CHAMAESYCE HYSSOPIFOLIA (L.) Small

5. CHAMAESYCE HYSSOPIFOLIA (L.) Small

6. CHAMAESYCE HYSSOPIFOLIA (L.) Small

7. CHAMAESYCE BRITTONII (Millsp.) Millsp.

8. CHAMAESYCE COWELLII Millsp. in Britton

9. CHAMAESYCE BLODGETTII (Engelm. ex Hitchc.) Small

Page
31.
35.
38.

^6.

47.

48.
53.

58.
59.

60.
64.
67.
73.
79.
80.
81.
92.
95.
99.

LIST OF PLATES, Continued

Page

Plate 20. CHAMAESYCE BLODGETTI I (Engelm. ex Hitchc.) Small 100.

Plate 21. CHAMAESYCE BLODGETTI! (Engelm, ex Hitchc.) Small 101.

Plate 22. CHAMAESYCE SERPENS (HBK.) Small 109.

Plate 23. CHAMAESYCE CENTUNCULO 1 DES (HBK.) Mi lisp. 115.

Plate 2k. CHAMAESYCE CENTUNCULOI DES (HBK.) Mi lisp. 116.

Plate 25. CHAMAESYCE TORRALBASII (Urb.) Mi lisp. 119.

Plate 26. CHAMAESYCE CAMAGUAYENS I S Mi lisp. 123.

Plate 27. CHAMAESYCE CORDIFOLIA (Ell.) Small 125.

Plate 28. CHAMAESYCE POLYGON 1 FOLIA (L.) Small 129.

Plate 29. CHAMAESYCE AMMANNIOIDES (HBK.) Small 132.

Plate 30. CHAMAESYCE CUMULICOLA Small 137.

Plate 31. CHAMAESYCE PROSTRATA (Ait.) Small l40.

Plate 32. CHAMAESYCE MENDEZII (Boiss.) Mi lisp. 146.

Plate 33. CHAMAESYCE HEPATICA (Urb. & Ekm.) ined. 1^9.

Plate 34. CHAMAESYCE HELWIGII (Urb. & Ekm.) ined. 151.

Plate 35. CHAMAESYCE H 1 RTA (L.) Mi lisp. 154.

Plate 36. CHAMAESYCE OPTHALMICA (Pers.) ined. 162.

Plate 37. CHAMAESYCE MACULATA (L.) Small 170.

Plate 38. CHAMAESYCE MACULATA (L.) Small 171.

Plate 39. CHAMAESYCE THYMIFOLIA (L.) Millsp. 177.

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

(Small) ined. 185.

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

(Small) ined. 186.

LIST OF PLATES, Continued

Page

Plate 42. C. ADENOPTERA (Bertol.) Small var. GUNDLACHI I

(Urban) Ined. 187.

Plate 43. CHAMAESYCE CONFERTA Small 192.

Plate kk. 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 MULTl NODI S (Urb.) Millsp. 204.

Plate 49. CHAMAESYCE TURPINI I (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 (Euphorb iaceae) is represented on all
continents, and, while many species are widespread in temperate areas,
the genus reaches Its greatest diversity in tropical and subtropical
reg ions.

In 1941 Wheeler revised the group in North America exclusive of
Mexico and southern Florida, treating it as a subgenus of the Linnaean
genus Euphorb ia. 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 f lor i st i ca 1 1 y , 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 Euphorb ia 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 (I898-
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, 19^4),

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

UCWl 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 Euphorb ia 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 (175^), and was delimited on the basis of its floral characters.
Reference was made to the earlier names Euphorb ium proposed by D'lsnard
in 1720 for much the same group, and to Ti thymal us and Ti thymaloides ,
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 Euphorb ia was
placed in the third Order of the eleventh Class, DODECANDRIA TRIGYNIA,

k.

5.

since Linnaeus had found that twelve or more stamens were usually present,
and that there were three styles. The description in Genera PI antarum
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- 1 i ke _E. mamm i 1 lar is , 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 (19^3) 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
v/ere named using Euphorb ia 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 Euphorb ia 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
1 dOO ent i t ies.

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 cyathlum is not restricted to this
genus, but occurs in several others recognized by even the most con-
servative taxonomlsts. Among these Pax (who had written the account
of the family for Engler and Prantl's Pf lanzenfami 1 ien In I89O) is
quoted as saying In 1924 that the cyathlum 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
cyathlum 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 (19^3) reports that a survey of

8.

somatic chromosomes in the Euphorb iaceae showed a considerable degree
of uniformity in size, shape, and basic number within all genera except
Euphorb ia , which seemed to include three distinct groups.

Studies on the electrophoresis of latex by Moyer (193^) 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 Pf lanzen-
fami 1 ien (I89O) 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

as a genus without considering the equally valid genera Ti thymal us ,
Aqaloma and Po i n s e 1 1 i a . It is beyond the scope of the present wori<
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 i t wi 1 1 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 Euphorb ieae of the Family Euphorb iaceae has been under-
taken.

I I . The Segregation 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 (19^3), however, supports Millspaugh (1909)
in designating Euphorb ia pep 1 i s L. as type since Gray was clearly describ-
ing this species under his name C. mar i t ima and, in fact, gave _E. pepl is
in synonymy, thus making the later name illegitimate. The type specimen
of _E. pepl i s L. in the Linnaean Herbarium becomes the type of _C. mar i t ima
S. F. Gray and of the genus Chamaesyce S, F, Gray,

Wheeler (19^3) 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 Anisophyl 1 um 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 I763.

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 lov;-growi ng , 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 Euphorb ia by Wiman (1752)

10.

with a similar comment that it was a type of low-growing fig (quas i
humi 1 i s f i cus) ,

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 sect ion Anisophyl 1 um in 1828, whi le
Rafinesque changed his mind after his publication of the subgenus
Chamaesyce in 1817 to treat them as a set of related genera in I838.
They were put bacl< into a single category as Euphorb ia subgenus
Ani sophyl 1 um Caesalp, ex Reichenbach in l84l , treated as the genus
An isophyl 1 um by Klotzsch and Garcke (1859), and finally the definitive
work by Boissier in 1862 followed Roeper by using section Ani sophyl 1 um
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 (19^1), 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

12.

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 i nterpet iol ar 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
Favycett and Rendle maintain Euphorb I a sensu lato. The split has not been

13.

js widely adopted in other regions, however, and an assessment of its
value on a worldwide basis must await a more comprehensive study.

III. 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.
vaq i nulata (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.-

15.

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,

Crolzat (1936b) describes the growth of the secondary axes as
sympodlal 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, 19^1) , 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 "laterals" or "side branches" will

16.

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, 1x1 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 i nterpet iolar 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

17.

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 fol iaceous 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 petal loid 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 cent r I fugal 1 y with respect to the cyathium axis. In our species
five, or rarely fewer, to about thirty is the usual number of staminate

18.

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.
Frui t

This is a trilocular capsule with one seed in each chamber,
usually roundly looed or triangular but sometimes subspher i cal ,
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
Euphorb ia sensu lato, is completely absent in Chamaesyce. The embryo

19.

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

20.

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 vestlture may differ
vyidely 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
Euphorb i a pep 1 i s L. , and the type specimen of this species in the
Linnaean Herbarium (sheet 63O-I8) is thus the type for _C. mar i t ima and
for the genus. Euphorb ia subgenus Chamaesyce (S. F. Gray) House, Bull.
N. Y. State Mus. 25^: 470. 1924,

Anisophyl 1 urn Haworth, Syn. PI. Succ. , I59, I8l2. Type:
Anisophyllum pepHs (L.) Haw., designated by Wheeler, Rhodora 4-3: 110.
1941. Euphorb ia section Ani sophyl lum (Haw.) Pax in Engler £■ Prantl,
Nat. Pflanzenfam., 3(5): 104. I89I.

Euphorb ia subgenus Chamaesyce Raf . , Amer. Monthly Mag. 2: 1 I9.
1817. Type: Euphorbia supina Raf.

Euphorbia section Ani sophyl 1 um Roeper in Duby, A. P. de Candolle,
Bot. Gall, ed 2, 1: 412. I828. Type: Euphorbia pep! is L. , designated
by Wheeler, Rhodora 43: 111. 1941.

Euphorb ia 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. I838. Type: Ditritra hirta
Raf., designated by Wheeler, Am. Mid. Nat. 30: 465. 1943.

Endoisila Raf., Fl. Tell., 4: 114. I838. Type: Endoisila
myrs ini tes Raf.

21.

22,

Aplarina Raf., New Fl. N. Amer., 99. I838. Type: Aplarina
prostrata Raf., designated by V/heeler, Rhodora ^3: 111. 1941.

Annual or perennial herbs, often suf f rutescent , 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 i nter-petiol ar 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 flovyers 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; overy three-celled, each v/ith 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. hyssopi fol ia in the New World
tropics as a v/hole, 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.

23.

The use of the taxonomic categories "species", subspecies", and
"variety" in this work follows conventional lines (see, for example,
Davis and Heywood, I963). 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,

1. Annual or perennial herbs, often woody at the base

and suf f rutescent, but rarely reaching 1 m. 3.

2. Leaves fleshy, subglobose or linear; closely branched

with stems often almost prostrate 1. C^. vaginulata.

2. Leaves membranous, ovate to ligulate; branching

divaricate, stems upright 2. C_, articulata.

3. Capsule glabrous ^«

3. Capsule pubescent 24.

k. Plants erect or ascending 5.

k. Plants prostrate to decumbent 12.

5. Leaf margins manifestly toothed, or, if entire,

leaves ligulate "•

5. Leaf margins entire or obscurely toothed below apex,

leaves never ligulate 10.

Ih.

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

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

7. Cyathia few, borne singly or rarely in groups of two ^_ ..,^^3
or three; plant perennial ^^' ^' £ •

7 Cyathia many, grouped on short leafy laterals or in

glomerules; annual but sometimes perennating

8. Capsule less than l.^f mm. long; cyathia usually in . ^ . -.^U,
peduncled leafless glom,erules 9. C. hvper . ci fol . a.

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 ^ ^ nutans,
with a rippled surface -* *

9 Tips glabrous; leaves glabrous or sparsely pilose

beneath; seed with 1-k lateral ridges ^^^ ^^ hvssopi fol ia.

on each face —

,0. (5) Leaves and young ste.s fleshy; stipules conspicuous

membranous, ca. 1 mm. long, white ->• _•

10 (5) Leaves membranous or somewhat coriaceous, young
stems not fleshy; stipules not conspicuous unless

because of ciliate inner surface, coriaceous, ..11.

ca. 0.5 mm. long, brown

11 Stipules densely white ciliate on surface nearest stem;

leaf and young stem surface minutely white-pap. Hose; .^.^^^
glands dark, appendages obsolete '+• ii* •

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 ^ ^ porteriana.
width of gland -*

12. (4) Stems wiry and stiff, rarely exceeding 0.5 mm.

diameter, usually many from root stock

12 {k) Stems flexible rather than wiry, few to several

from rootstock, either reaching 2-3 mm. diameter or ^ ^^^ ^^^

minute, threadlike

14.

13. Leaf margins enti re

25.

13. Leaf margins serrate, at least In basal leaves 15.

14. Leaves deltoid, margin somewhat revolute; glands fleshy,
appendages obsolete or minute k] , _C. del toidea ssp, del toidea

var. del toidea

14. Leaves orbicular to elliptic, plane; glands cupped,
appendages prominent, to three times

as wide as gland 19. £. camaquayensis.

15. All leaves serrate, most 4-8 mm. long; plants usually

decumbent 18. C_, torralbas I I .

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

lax, prostrate mat 17. C^. centunculoldes.

16. (12) Leaves and stems pubescent 39. £. paredonens i s.

16. (12) Leaves and stems glabrous 17.

17. Stems minute, threadlike, tomentose on upper surface;
cyathia subsessile; styles undivided at apex; capsule

less than 0.7 mm. long 42. _C. tumistyla.

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.

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

parted or laclniate 19.

18. Stipules joined, at least on upper or lower surface of

tips of branches, fringed or entire, never laclniate 22.

19. Seeds strongly angled; appendages conspicuous; leaves

ovate to ovate-oblong, almost as wide as long 20. _C. cordi fol ia.

19. Seeds terete to obscurely angled; appendages incon-
spicuous or absent; leaves narrowly elliptic

to oblong, much longer than wide 20.

20. Capsule more than 2.5 mm. long; seed cuneiform,

more than 2 mm. long 21. C. polygonlfol la.

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

than 2 mm. long 21 .

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

and laterals 22. C. ammannioides.

26.

21. Seed 1-1.4 mm. long; leaves not fleshy, all

similar in size 23. C_. cumul icola.

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 subcor iaceous; leaf and stem surface

somewhat papillose 14. _C. cowe 1 1 i i .

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

smooth 15. £. blodqett i i .

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. las iocarpa.

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.

27.

29. Stem short-pubescent in lines at sides and sometimes on
upper surface; faces of seed with deep,
transverse furrows ..... 2k. C_. prostrata.

29. Stem long-hirsute, at least in lines at sides;

surface of seed rippled 25. C_. mendez i i .

30. Capsule not completely exserted, splitting one

side of cyathium at maturity ..... 32. _C. thymi fol ia.

30. Capsule completely exserted at maturity 31.

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. hepat ica.

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.

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. helwigi i .

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 3^. 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. £. adenoptera.

34. (26) Leaf margins crenate-dentate or roundly

serrate, apex obtuse 30. C. berteriana.

3k. (26) Leaf margins sharply serrate, apex acute 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. hi rta.

35. Cymules terminal and on leafy laterals; stem branching

freely; mostly low, small-leaved plants 29. C,. opthalmica.

28.

36. (24) Plants suf f rutescent; 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_. cayens i s.

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

38. Open shrubs, branched from base; capsules more

than 2 mm. long 7. C^. myrt i 1 1 i fol ia.

38, Suf f rutescent, base mostly unbranched; capsules

1.3~1.5 Tim. long 6. _C, porteriana var. keyens i s.

39. (3o) Stems pubescent on only one surface kO.

39. (36) Stems pubescent on all surfaces k\ ,

kO, Stems tufted, upright, usually less than 0.5 mm.

diameter; leaf surface glabrous 37. £_. mi nutula.

kO. Stems prostrate, usually ca. 1 mm, diameter;
scattered white hairs on
surface of leaf 36, _C. mul t i nod i s,

k] . Cyathia in groups of 2-5 at nodes 43. _C. leonardi i .

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^. garber i .

42, Plants delicate, stems wiry, scarcely reaching 1 mm.
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, vi 1 lous-h i rsute, tips canescent;
capsule reniform, sharply three-lobed,

angles acute 40. _C, p i netorum.

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. turpini i.

29.

1. CHAMAESYCE VAGINULATA (Griseb.) Mi lisp., Field Mus. Bot. 2:
302. 1909. Euphorbia vaqinulata Griseb., Fl. Brit. W. 1. Islands, 52.
1859- Type: Turk Islands, Hjalm. (Lectotype: K 1, 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-
t i sch-Geobotan i sches 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, subart iculate 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;

30.

calyx a triangular pad of tissue; ovary glabrous; styles spreading,
o.k 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
whi te.

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,
(l^Y); 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 768I (GH, NY). Turks Islands: Waterloo, Grand
Turk, 20 Feb. -24 Mar. I9II, Millspaugh & Millspaugh 9055 (GH, NY);
Grand Turk, 27 Aug.-] 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

31

*-»^

new rORK BOTANICAL GAROEK
CLO MUSEUM OF NATURAL HISTORy

I AI'tllKAlK IS I 11 I III n ill iV 1-

2.3":? THE CAICOS GROUP

|itii[iiii|iniiiiii|llii|iiii|iiiii

Plate 1. CHAMAESYCE VAGINULATA (Griseb.) MiHsp.

C vaqinulata (Griseb.) Mi lisp., Long Cay, Ca i cos Group, Bahamas, 7 Mar.
Tgll, Millspa'ugh & Mlllspaugh 9237 (NY).

32.

Map 1. Distribution of _C. vag inulata

Map 2. Distribution of _C, buxi fol ia

33.

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:
51h. 1916, Euphorbia articulata Aubl., PI. Guian., 1: 480, 1775. Type
in Herb. Brit. Mus. if extant.

Euphorb ia 1 i near is Retz. , Observat iones Botanicae, 3: 32. 1783.
Type: Bot. Mus., Lund, Sweden, if extant. _E. linearis heterophylla
Kuntze, Rev. Gen. PI., 605. I89I. _C. 1 i near is (Retz.) Millsp., Field
Mus. Bot. 2: 410. I9I6.

Anisophyl 1 um vahl i i Kl. £• Gke. , Monats. Akad. Berlin 1859: 36.
1859. Type: Specimen by Willdenow, in Bot. Mus. Berlin, if extant.
Euphorbia vahl i i Willd. ex Kl. S- Gke,, loc. cit. 37. 1859. Chamaesyce
vahl ii (Kl. & Gke.) P. Wilson, Bull. N.Y. Bot. Gard. 8: 395. 1917.

Shrub; to 6m., 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.

3^.

Staminate flovvers 8-20 per cyathium, androphores glabrous, to 2.5 nim.
long at maturity. Gynophore glabrous, to 3 n™. 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 nim. 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 S- 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 I96I, Alain 9265 (U); Yauco, 2 Apr. I88O, Garber s.n. (GH) ;
Seashore, Fajardo, 17 Feb. I9OO, Heller 4608 (F, GH, MO, NY); Santurce,
21 Feb, 1900, Heller 4678 (F, MO, NY); Rio Piedras, Fajardo, k July 1913,
Johnston 792 (NY); Guayanilla to Tallahoa, 13 Mar. 1913, Shafer 1993 (F,
NY); Fajardo, 19 May 1885, Sintenis 16)2 (MO, NY). LESSER ANTI LLES.
Culebra, 3-12 Mar. I906, Britton & Wheeler k (F, NY); Culebras Island,
19-20 Jan. 1899, Millspaugh I78I (F); Racoon Bay, St. Thomas, 31 Jan.-
k Feb. 1913, Britton, Britton & Shafer 167 (F, NY); St. Thomas, Jan. I887,
Eggers s.n. (F) ; Tortola, 16 Nov. I9I8, Fishlock 263 (F, GH, NY); Anagada,
19-20 Feb. 1913, Britton S- Fishlock 978 (NY); Barbuda, 6 Apr. 1956, Smith
10466 (GH, IJ). St. Kitts: Canada Estate, 8 Sept. -5 Oct. I9OI, Britton £-

35.

.... .^..- ''^Y

|iiiiiitii|iJM|iiii|iiii|iiimiii|im|iiii|iui|ilii|iili|iiii|iiii|liiiiiiiiiii!n^
3lkyilil!liiilililililJ5iklllJiul5iliii:. ''

Plate 2. CHAMAESYCE ARTICULATA (Aub1.) Britton
E. linearis Retz., St. Thomas, ? Bertero (MO).

36.

Cowell 263 (NY); Sir Timothy's Hill, 24 Dec. 1958, Proctor l84l8 ( 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 (U); Guadeloupe, 1892, Duss
2451 (NY); Martinique, Sieber 392 (F) ; St. Lucia, 22 Apr.-l8 May 1950,
Howard 11417 (iJ). St. Vincent: Mt, Pleasant, 22 Mar. I962, Cooley
8534 (GH); Bequia, 26-31 Mar. 1950, Howard 11269 (GH).

The affinities of this species are with _C. myrt i 1 1 i fol ia (L.)
Mi lisp, and with coastal species of the Bahamas and Florida such as
_C. lecheoides (Mi lisp.) Mi lisp, and _C. porter iana Small. These, however,
differ in size, habit, cyathial and stipular characters.

A number of pubescent collections of _C. art iculata 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, 1 i near i s is a synonym.

3. CHAMAESYCE 3UXIF0LIA (Lam.) Small, Fl. SE. uS., 712. I9O3.
Euphorbia buxi fol ia 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, 1 788 . Type: unknown. A specimen
from Swartz v;ith this name is reported from Herb. Stockholm, Fawcett &
Rendle, Fl. Jam., 4(2): 336. 1920.

_E. 1 itoral is HBK, , Nov, Gen, et Sp., 2: 44. I8I7. Type: Cumana,
prope Bordones (Herb. Mus. Paris, not seen).

_E. f lexuosa HBK., Nov. Gen. et Sp., 2: 44. I8I7. Type: Cumana,

37.

Bonpland (Herb. Mus. Paris, not seen). _E. bux i fol ia Lam. var. f lexuosa
(HBK.) Boiss. 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 nal<ed
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.

38.

Plate 3. CHAMAESYCE BUXIFOLIA (Lam.) Small

C. buxifolia (Lam.) Small, Grassy Key, Monroe County, Florida q Anr
1963, Burch 153 (FLAS). ' ^ ^"^

39.

and of their associated beach-scrub; Bermuda, Florida in all coastal
areas south of Pinellas and Volusia counties, Bahamas, Yucatan, Cuba,
Jamaica, Hispanlola, Puerto Rico, Lesser Antilles and the northern
coast of Venezuela (Map 2 ) .

Representative specimens seen: FLORIDA. Brevard: Cape Canaveral,
2-5 Apr. 190^, Burgess 668 (F, NY); Merritt Island, 18 Apr. 19^6,
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 £■ Krai 53854 (FSU, NSC) ; Lauderdale Beach, 17 Dec. 1929, Moldenl<e
263a (DUKE, MO, NY). Collier: Vanderbilt Beach near Naples, 13 Aug. I963,
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 S- Carter III6 (NY), Flagler: Beach near
Marineland, I8 Nov. I96I, Godfrey 6l675 (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, I9OO, Hitchcock 325 (F,
MO); Captiva Island, 21 May I9OI, 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. I963, Burch 125 (FLAS). Monroe: Cape Sable, Middle Cape,
18 Apr. 1964, Burch & Ward 304 (BM, BUS, E, F, FLAS, GH, I J, 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 I96O, Webster & Samuel 10210 (DUKE, IJ, PUR). Palm Beach:

40.

Beach between Palm Beach and Lake Worth, 7 Aug. I963, Burch 118 (FLAS) ;
Palm Beach, 2k May 1895, Curtiss ^kOk (NY); Juno Beach, 20 Apr, I962,
Lakela 2^955 (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. I963, 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 I887,
Rothrock s.n. (F, NY). Volusia: Turtle Mound, 27 Apr, I96I, Ray, Wood,
Smith & Eaton 10821 (FSU, NCU) . BAHAMAS. Great Bahama, 5-13 Feb. 1905,
Britton S- Millspaugh 2365 (F, NY); Bimini, 27 July 1964, Stimson 776
(DUKE, FLAS); Riding Rocks, Great Bahama Bank, 13 May 1909, Wilson 79l4
(F, MO, NY); New Providence, Nov. I89O, Hitchcock 26 (F, MO) ; Salt Key,
21 May 1909, Wilson 8O89 (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. I96O, Williams 7 (DUKE, PUR); South Caicos,

27 June 1954, Proctor 898I (IJ); Grand Turk, 22-27 Apr. 1954, Lewis s.n.
(IJ). CUBA. Isles of Pines: Caleta Cocodrilos, 8 Mar. I9I6, 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 Brae, South
Shore, 10 Feb. 1899, Millspaugh 1233 (F) ; Grand Cayman, Grape Tree Point,
14 Apr, 1956, Proctor 11973 (U). JAMAICA. Portland: Port Antonio, 21
June 1897, Fredholm 3038 (NY). St, Catherine: Healthshire Hills, 3 May
1913, Harris 11627 (F, MO, NY, UCWl). St. Elizabeth: Long Acre Point,

41.

13 Sept, 1907, Harris 996? (F, NY, UCWl). St. James: Montego Bay,
June 1951, Robbins 423 (UCWl). St. Mary: Galina Point, 7 Dec. 1952,
Proctor 7452 (IJ). St. TInomas: Morant Point, 1-13 Mar. I909, Britton
4098 (F, NY). Westmoreland: Negri 1 area, Homer's Cove, 23 June 1959,
Webster, Ellis & Miller 85OI (PUR). HAITI. Bayeux, 28 Nov. 1924,
Ekman H2676 (IJ); Port de Paix, 28 Apr, 1925, Ekman H3957 (U); 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 HI3I65 (F, PUR), Barahona: San Domingo,
May 1911, Fuertes 1111 (F, NY); Trail between Pedernales and Aceitial,
8-12 Aug. 1946, Howard 8IO6 (NY), PUERTO RICO, Condado, San Juan,
16 Sept. 1941, Blomquist 1 1 789 (DUKE); Fajardo, 4 July 1913, Johnston
791 (NY); Ratones Island, 21 Jan. 1899, Millspaugh 658 (F) ; Salinas de
CaboRojo, 6 Feb. 1885, SIntenis 559 (UCWl); Beach at Santurce, 21 Feb.
1900, Heller 4681 (F, MO, NY), LESSER ANTILLES. Soldier Bay, St.
Thomas, Dec. I88O, Egger s.n. (F); St, Croix, 1 Nov, 1895, Ricksecker
36 (F, MO, NY); Anguilla, I9O8, 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, 1 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. I9IO,
Flynn 101 (MO); Hamilton Bay Islets,. 29 Dec. I898, 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

k2.

as the completely herbaceous species.

Euphorbia mesembr ianthem i fol ia 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. Stearn of the British
Museum (Natural History) was kind enough to search their herbarium, and
Dr. K, H, Rechinger checked that of the Naturh i s tor i sches 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^. mesembr i-
anthemifol ia , it would be irresponsible to accept it and upset the well-
understood use of _C. buxi fol ia , 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

43.

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 - 1 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 \/k 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.

4^.

Key to varieties of _C. 1 echeo ides

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

stem; branches decumbent a. var. wi 1 son i i .

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. exumens i s.

4a, C. LECHEOIDES (Millsp.) Millsp. var. WILSONM (Millsp.) comb,
nov. C^. wi Isoni i Millsp., Field Mus. Bot, 2: 301. 1909. Type: Castle
Island, Bahamas, 22 Dec. 1907, P. Wilson 7795 (Holotype: F !; Isotype:

NY :).

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, I963, Dunbar 239 (GH) ; Inagua, ]k Oct. 1904, Nash
£■ Taylor 1034 (NY); South Caicos, near East Bay, 17 June 1954, Proctor
8828 (iJ); Turks Islands, Salt Cay, 10 Mar, I9II, 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 Mi 1 1 sp. , Field Mus. Bot. 2: I63. I906. Type: Grand Turk,
Turks Islands, Bahamas, 27 Aug.-l Sept. 1905, G. V. Nash & N. Taylor

45.

(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 73^7 (NY); Long Cay, 7-17 Dec. 1905,
Brace 4179 (NY); Crooked Island, 9-23 Jan. I906, Brace 4588 (NY);
Mariguana, 6-8 Dec. 1907, Wilson 753^ (NY); Inagua, 21 Jan. I962, Dunbar
53 (GH), South Caicos: Northern Hills, 19 June 195^, Proctor 8887 (GH,
IJ); Cockburn Harbor, 28 Mar. 1958, Proctor 17^97 (U). Between
Commissioner's House and airfield. Grand Turk, 13 June 195^, Proctor
8799 (GH, I J).

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,
exumens i s .

4c. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS (Millsp.) comb,
nov. C. exumens is Millsp., Field Mus. Bot, 2: 301. 1909. Type: Stocking
Island, Great Exuma, Bahamas, N. L. Britton & C. F. Millspaugh 3071
(Holotype: F '. ; 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 i960, 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

ki.

*!2K

NCW yORK BOTANICAL CAROCN
CLO MUSeUM OF NATURAL HISTORV

(Ai'i nK\n< t\ < .1 III! n\n \m \s

in 11 i:^

^^'I7H

ilJiiiliuU.

Plated. C, LECHEOIDES (Millsp.) Millsp. van. WILSONII (Millsp.) ined.

Isotype: _C. wi 1 son i i Millsp., Castle Island, Bahamas, 22 Dec. 1907,
Wi Ison 7795 (NY).

hi.

E NEW YONK BOTANICAL CAflDEN

.r^

Grand Turh. Auguftt 27-5cptembe<

Plate 5, CHAMAESYCE LECHEOIDES (Millsp.) Mi lisp, var, LECHEOIDES

Isotype: £. lecheoides Millsp., Grand Turk, Bahamas, 27 Aug.-l Sept,
1905, Nash & Taylor 3888 (NY).

k8.

i. Hitiiiiiin|iiiiMiii|iMi|iiiiliiill \^

IfhiliZiffidlll^

r ' . KC-Uf'1^

Plate 6. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS (MiHsp.) ined.
isotype: E. exumensis Millsp., Stocking Island, Bahamas, 22-28 Feb. 1905,
Millspaugh 3071 (NY).

49.

Map 3. Distribution of _C. 1 echeoi des var. wi 1 son i i

Map k. Distribution of C_. I echeoides var. lecheoides

Map 5. Distribution of ^. lecheoides var.

exumens i s

Map 6. Distribution of C.

ii- cayens i s

Island, 12-13 Mar. 1907, Britton & Millspaugh 61^+3 (NY); Conception
island, 5 Jan. 1932, Fairchild 16 (GH) ; Rum Cay, k 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,

52,

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 1 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. I96O,
Webster IO896 (DUKE, PUR); Abaco, 8 Dec. 1904, Brace 1571 (NY). Berry
Islands: Whale Cay, 29 Jan. I9O5, 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 I909, Wilson 8027 (MO, NY); Water Key, 22 May
1909, Wilson 8139 (MO, NY).

53.

|riti|iiii|MM|iiiHHM|iin|HiijiN:|HH|nH|iiM|iiii|iiii|iiii|iiii|iin|iiiim
ali!:li!^i'laJ.lJllJ)lJuJilJilllililililSlllllLllljllBl!lllill)ljlJa..M

.f-.i^/l.

Plate 7. CHAMAESYCE CAYENSIS (Mi lisp.) Mi lisp.

Isotype: _E. cayens i s Mi lisp., Joulter's Cays, Bahamas, 11 April 190^,
Mi llspaugh 2295 (NY).

5k.

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. cayens i s, 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
k 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
suborb icular , 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,

55<

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 darls red, margin entire or crenulate, fifth gland linear, shorter
than lobes or obsolete, sinus broad and shallow, Staminate flowers 8-lU
or more per cyathium, androphores glabrous, ca. 1 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, porter iana

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- 1 i ke, 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 . va r . scopar ia.

56,

6a, CHAKAESYCE PORTERIANA Small van, KEYENSIS (Small) comb, nov.
Chamaesyce keyensis Small, Torreya 28: 6, 1928. Type; No Name Key,
Monroe County, Florida, 4-5 Feb, I9I6, J, K, Small 7^39 (Holotype NY :).

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,
Burch 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 i960, 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, I9O6, Small & Carter 2504
(NY); Pinelands east of Naranja, 14 Jan. I9O9, Small & Carter 3O66 (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

57.

(BM, BUS, E, F, FLAS, GH, IJ, MO, NSC, NY, PUR, US, USF). Monroe County:
Vacca Keys, 28-29 Jan. 1909, Small & Carter 30^8 (NY); Boot Key, 7-12
April 1909, Britton 5^0 (NY); Big Pine Key, I^ Dec, 1951, Killip ^15^7
(NY); Key West, 19 Sept. 195^, Killip kkkjk (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. keyens is.

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 :).

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 19^2, Eyles & Eyles 8I86 (GH) ; Pine woods. Big Pine Key, 10
Aug. 1963, Burch \kO (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 i ntergradat ion 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

58.

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

Holotype: _C. keyensis Small, No Name Key, Monroe County, Florida, k-S
Feb. 1916, Small 7^39 (NY).

59.

J xi^

|iiiiMiii|iiiiiiiii|iiiimn|iniiiiiijirii|iiii|iiiiniiniifi|TinniiniH
IJlUilililililKililililihlilililililililililSj-

Q. /w4'w. SJ >wi^<' _

?^^

.'^~

Plate 9, CHAMAESYCE PORTERIANA Small \iar . PORTERIANA

Holotype: C, porteriana Small, Miami, Florida, 1877, Garber s.n. (NY)

60.

1 J^ :^..-

a^rft^sg

THE NEW YORK BOTANICAL CAHOC

111 II \2 11 II 1 .',

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

Holotype: _C. scopar ia Small, Big Pine Key, Monroe County, Florida, 17
Nov. 1912, Smal 1 3819 (NY).

Map 7. Distribution of _C. porter iana var. keyens i s

Map 8. Distribution of _C. porter iana var. porter iana

Map 9. Distribution of C. porteriana var

scopar ia

/-,--,- i"

--^ ^M, ,

Map 10. Distribution of C. articulata and C. myrtlllifoM^

63.

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 MYRTI LLI FOL I A (L.) Mi lisp.. Field Mus. Bot. 2:
410. 1916. Euphorbia myrtill ifol ia L. Syst. Nat. ed. 10, 10U8. 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-iO 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- ye I low,
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;

64

I -^ II II I .)

■--riTni. „r Mim, j
ir.r)lt\ or J.v.MAK V

i'",.h SI.. AJiBRrir

Plate 11. CHAMAESYCE MYRTI LL I FOL I A (L.) Millsp.
L' myrtillifoli^ L., Gordon

Proctor 9595 (NY).

Town, St. Andrew, Jamaica, 8 Dec. I954,

65.

calyx a pad of tissue or obsolete; ovary sparsely pilose; styles spread-
ing, to 1 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 nim, 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
189^, Harris 5196 (NY); Valley of Hog Hole River above Gordon Town,
8 Dec. 195^, Proctor 9595 (iJ, NY); lA mile northeast of Gordon Town,
20 Mar. 1956, Proctor 118?^ (GH, IJ) .

The affinities of this species are with the shrubby £. art iculata,
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-
t i 1 1 i f o 1 i a 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, PI. 17. I8l6, Type: "Habit, [at]
in N.[ova] H.[ispania]. Perhaps at Madrid , , ," (Wheeler, 19^+1).

_E. presl i i Guss. , Fl, Sic, Prod, 1: 539- 1827. Type: Palermo,
Italia, Todaro (fragment F seen by Wheeler), C. presl i i (Guss,) Arthur,
Torreya 11: 260, 1912.

_E. hyper ici fol ia L. var. communi s Engelm. in Emory, U.S, & Mex.
Bound, Surv. 2: 188. 1859. Type: New Mexico, 1851-52, C. Wright 18^+2

66.

(fragment F seen by Wheeler).

_E. hyper i c i f o 1 ia L, sensu Michaux, Fl. Bor.-Am., I803; Torrey,
Fl. State New York, 1843.

E^, maculata L. sensu Wheeler. Contr. Gray Herb, 127: 7^. 1939.

Annual; erect or ascending to 8 dm. Stem to k 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
nearly distinct near tip of shoot, triangular to 1 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 - 1 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-l4 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, O.k - 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

67.

91 (1.-1 5

la !'< >•

Plate 12. CHAMAESYCE NUTANS (Lag.) Small

r nut^n^ aaa ) Small, between Marianna and Campbell ton, Jackson County,
?io7fS7^7 Nov! 196U, Godfrey 6U932 (FLAS).

68,

below equator, strongly three-Iobed, 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
1 ight ang les.

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 mis ident i fi cat ions of plants of _E. hyssopi fol ia L. and E^. hyper ici fol ia
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 I898, Hitchcock 1697
(F), Brevard: Okeechobee region, 17 Sept, I9O3, Fredholm 60l4 (GH, NY),
Citrus: June-July I898, Hitchcock 1699 (F, MO), Gadsden: Jim Woodruff
Dam, Chattahoochee, 19 Oct, I963, Burch 207 (FLAS, IJ, US); River Junction,
7 Aug. 1927, Wiegand & Manning I8I8 (GH) ; Chattahoochee, 7 Aug. 1927,
Wiegand & Manning I8I9 (GH) ; Chattahoochee, 8 Sept. 1931, West s.n, (FLAS).
Jackson: Marianna, 19 Oct. I963, Burch 210 (FLAS); near Cottondale, 26 Oct.
1964, Burch 504 (FLAS, MO, NCU, NY); between Marianna and Campbel 1 town,
7 Nov. 1964, Godfrey 64932 (FLAS, FSU) ; Marianna State Park, 15 Sept. I96O,
Mitchell 839 (FSU); Marianna Caverns State Park, 19 Nov, I96O, 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

69.

(F, GH, NY). Levy: June-July I898, Hitchcock 1 698 (F). Wakuila: Newport,
13 Sept. I96U, 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. I963, Manuel 265 (A).

This species bears a strong resemblance to C_. hyper ici fol ia (L.)
Millsp., _C. hyssopi fol ia (L.) Small and C. las iocarpa (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. hyperici fol ia 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 (I8O3) 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.

70.

Map 11. Distribution of C. nutan;

Map 12. Distribution of C. hvper

yperici fol ia

71.

The basis for Smith's actions was sound in that the description of
E. hypericifo] ia 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 (I965)
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 hypericifol ia L., Sp, PI. ed. 1, ^5^. 1753. Type:
Sloane Hist. 1, t. 126. (lectotype, designated by Burch, Rhodora 67. I965,
in press) .

_E. pi lul ifera L. sensu Wheeler, Contr. Gray Herb. 127: 76. 1939.
(But see disclaimer, loc. cit, 78).

_C. qlomer ifera Millsp. Field Mus. Bot. 2: 377. 1913. Type: El
Rancho, Dept. Jalapa, Guatemala, 20 Jan. I908, W. A, Kellerman 8053 (F I).
_E, qlomerifera (Millsp,) L, C, Wheeler, Contr, Gray Herb. 127: 78. 1939.

Annual or perennating in tropics; erect to ascending, or sprawling
to k 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 darl< brown. Leaves membranous, blades ovate-elliptic to

el 1 ipt ic-obovate, at times somewhat falcate, 15-35 x 8-1^ mm., base inequi-
lateral, rounded or cuneate, margin serrate, apex acute, glabrous, green,
sometimes purple-spotted above and lighter green on underside; petiole
to 1 mm. long; stipules joined, sheathing, 1 - 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-k cleft. Cyathia in

72.

short-stalked, lateral and terminal, congested dichasia forming glomerules
with a few basal leaves; involucre short-pedunc led 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,
Stami nate flowers 10-12 or rarely only k 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, subspher i ca I , 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 Gkk (NY), Collier: road
to Marco Island, 15 Apr, 1954, West, Arnold & Cooley 14? (FLAS). Dade:
vicinity of Homestead, 22 Nov, 1964, Godfrey 65583 (FLAS, FSU) ; Buena
Vista, 1 Jan, 1930, Moldenke 330 (DUKE, NY); Miami, 14 May 1904, Tracy
9127 (F, MO, NY). Glades: Ca loosahatchee River bank, 7 Nov. 1964,

73.

iiitittiliiitili

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Plate 13. CHAMAESYCE HYPERI C I FOL I A (L.) Millsp.

C. hypericifol ia (L.) Millsp., University of Florida campus, Gainesville,
Alachua County, Florida, 30 Oct. 196^, Burch 521 (FLAS).

Ih.

Hodgson 230 (FLAS), Highlands: Rd, 25, south of Sebring, 22 June 19^8,
Garrett s.n. (FLAS), Lee: Sanibel Island, 13 Apr, 195^, West & Cooley
2630 (USF). Manatee: U.S. k\ & U.S. 19, 29 Nov. I962, Ward & Burch 3I63
(FLAS). Monroe: Upper Mateconbe Key, June, ?? , Curtiss 2486 (F, MO) ;
No Name Key, 21 Mar. I898, Pollard, Collins & Morris 126 (F, NY); Key
Vaca, 3 Sept. 195^, Thorne 15177 (U); Key Largo, north end, 27 June I96O,
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 I963, 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, I6 Apr. 1904, Millspaugh 2399 (F,
NY). Andros: Mastic Point, 19-28 Mar. I907, Brace 7087 (F, NY); Nicholl's
Town, 4-5 Feb. I9IO, 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. I89O, Hitchcock s.n. (F). Cat
Island, the Bight £■ vicinity, 1-6 Mar. 1907, Britton 5- Millspaugh 58I6
(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).
Oriente: Monte Verdi, Jan. -July 1859, Wright 1422 (F, MO, NY). Santa Clara:
Cienequita, Cienfuegos, 2 May 1895, Combs 4 (F, MO, NY); Limones, Soledad,

75.

Cienfuegos, 27 Sept, 1924, Jack 633^ (MO). GRAND CAYMAN, Georgetown,
2 June 1963, Crosby, Hespenheide, £■ Anderson 50 (DUKE, UCWl); Spot Bay,
13-14 Feb. 1899, Millspaugh 1304 (F), JAMAICA, Portland: Kildare, 21
Aug, 1962, Adams 11555 (UCWl); Port Antonio, 4 Jan. I9O6, Wight 2 (F,
NY). St. Andrew: Mount Pleasant, Stony Hill, 21 July 1912, Harris
11134 (F, NY, UCWl); Long Mountain, June 1953, Robbins 434 (UCWl).
St. Ann: Mosely Hill Cave, 1 mile west of Blacks tonedge, 12 Dec, 1952,
Proctor 7474 (IJ). St, James: Ironshore, May 1953, Asprey 432 (UCWl),
Trelawny: Cowie Park near Troy, 19 Oct. 1917, Harris 12646 (F, MO, NY,
UCWl). Westmoreland: Negri!, 29 Mar, I962, Adams IO98O (UCWl). HAITI.
Jeremie, 13 July 1941, Seibert 1725 (MO) ; Port au Prince, 21-23 Feb. 1920,
Leonard 2778 (NY); St. Michel de I'Abalaye, 1927, Ekman 8394 (F, IJ).
DOMINICAN REPUBLIC, Barahona, June I9IO, 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 I9OI, 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 I886, 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. I9I8, Fishlock 144 (NY). St, Kitts, 8 Sept.-
5 Oct. 1901, Britton & Cowel 1 (NY). Guadeloupe, Poi nte-a-Pi t re, 19 May
1936, Stehle 1190 (NY); Dominica, Soufriere, 1903, Lloyd 490 (NY).

76.

Martinique: Case Pilots, 1879, Duss ^88 (F, MO, NY); Ste. Anne village,
29 July 1939, Egler 39-208 (NY). Barbados: Batinsheba, Apr. -June, 1895,
Waby 101 (F); Waterford, St. Michael, 15 Aug. 1906, Dash 166 (F).
Grenada, St. George's, 29 Sept, I9O6, Broadway s.n. (F). Extra-territorial
specimens: BERMUDA. St. David's Isle, 22 May-6 June 191^, 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. hyper ici fol ia 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
PI antarum, 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 (I768). This matter, and Wheeler's appli-
cation of the name to the plant known here as _C. las iocarpa, are discussed
in more detail by Burch (1965).

The use of the name _E, p i 1 ul i fera L. for this species cannot be
accepted if the sheet (63O-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 character i st ic of seed of _C. hyper i ci fol ia, and
the general aspect of the plant was closer to that of _C. hi rta.

10. CHAMAESYCE HYSSOPIFOLIA (L.) Small,J,N.Y. Bot. Card, 3: ^29.
1905. Euphorbia hyssopifol ia L. Syst. Nat. ed 10, 1048. 1759. Type: in
Herb, Linn, sheet 63O-9 ! Anisophyllum hyssop i fol i urn (L.) Haw,, Syn. PI.
Succ, 161. 1812. _E. hyperi cifol ia L." fi hyssop i fol ia L." Griseb, Fl, Br.

77.

W. Ind. Isl., 5^. 1859. E.. bras i 1 iens i s Lam. var. hyssopi fol ia Boiss.
in DC. Prod, 15(2): 24. 1862.

E. bras I 1 iens is Lam., Encyc. 2: 423. 1786. Type: Mus. Nat.,
Paris, if extant. _C. brasi 1 iensis (Lam.) Small, Fl. SE, US., 712. 1903.

C. ni rurioides Millsp., Field Mus, Bot. 2: 394. 1914, Type: Malvern,
Santa Cruz Mountains, Jamaica, 5 Sept. 1907, Britton II86 (holotype: F 1,
isotype: NY !). _E. ni rurioides (Millsp,) Fawcett & Rendle, Fl, Jam., 4(2):
339. 1920,

_C, jenninqsi i 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 1; Isotype: NY !).

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,

78.

sometimes f lar i ng-campanu late, 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 mi s ident i f icat ions of plants of _C. hyper ici fol ia (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, Krai 1572 (FSU). Brevard: Melbourne, 22 Aug. 1958,
Krai 7971 (FLAS, GH, NY, USF). Broward: Pinelands, Ft. Lauderdale, 19 &
25 Nov. 1903, Small £■ Carter ll4l (F, NY); U.S. 27, north of Andytown,

79.

Plate lU, CHAMAESYCE HYSSOPIFOLIA (L.) Small

C. hyssop i fol ia (L.) Small, Fairchild Tropical Gardens, Dade County,
?iorida, 31 December 1962, Burch 52 (FLAS).

oitAiiAJiSTCS NIRUHIOIDRE sp.noT.

80,

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Plate 15. CHAMAESYCE HYSSOPIFOLIA (L.) Small

Isotype: C, nirurioides Mi lisp., Malvern, Santa Cruz Mountains, Jamaica-
5 Sept. 1907, Britton 1186 (NY). '

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PLANTS OF ISLE OF PINES

Plate 16. CHAMAESYCE HYSSOPIFOLIA (L.) Small

Isotype: _C. jenn i ngs i i Millsp. in Britton, near Vivijagua, Isle of Pines
Cuba, 7 May 1910, Jennings 621 (NY).

82.

26 Jan, I963, 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, I963, Burch I85 (FLAS),
Citrus: Dunnellon, 16 June 1958, Krai 69OI (FLAS, FSU, USF) . Collier:
Immokalee, 26 Sept. 1964, Lakela 27492 (USF). Dade: Buena Vista, 1 Jan,
1930, Moldenke (DUKE, F, MO, NY); Miami-Kenda 1 1 Station, 5 Nov, I9O6,
Small & Carter 2521 (NY), Duval: Jacksonville, 17 Nov, 1929, Moldenke
5232 (NY). Franklin: Apa lach i cola , 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. I96O, Ray, Gleason & Eaton 9728 (USF).
Hillsborough: Tampa, 20 Mar. 1923, Churchill 546 (MO, US); Oldsmar,
12 Aug. i960, Ray, Lakela & Patman IOOO6 (USF), Jackson: between Mari-
anna & Campbell ton, 7 Nov, 1964, Godfrey 64933 (FSU). Lake: Eustis,
16-30 June I894, Nash 1076 (F, MO, NY), Lee: Ft. Myers, July-Aug. I9OO,
Hitchcock 324 (F, MO, NY); 5 miles east of Ft. Myers Beach, 1 Aug. 1958,
Krai 7539 (GH, USF). Leon: Tallahassee, 30 June 1955, Godfrey 53571
(DUKE, FSU, NY). Marion: Silver Springs, 1 Oct. I963, Godfrey 63145 (FSU),
Manatee: Bradenton, 13 June I9I6, 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. I963, Ward £■ Burch 3332 (FLAS). Nassau:
Fort Clinch, 12 Oct. I963, 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 667O (MO, NY), Osceola: St, Cloud,

27 Feb. 1952, Cooley 1032 (USF). Palm Beach: between Palm Beach & Lake

83.

Worth, 7 Aug. I963, 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. 19^0, Laessle s.n. (FLAS). St. Lucie: Ft,
Pierce, 7 Aug. I963, Burch 122 (FLAS). St, Johns: Ponte Vedra, 11
Oct, 1964, Burch 437 (FLAS). Sarasota: Osprey, 10 Mar, 1904, Smith
712 (DUKE). Sumter: Wi thlacoochee River bridge, 8 Nov. 19^6, West &
Arnold s.n. (FLAS), Suwannee: Live Oak, k Oct. 196^, Elam s.n. (FLAS).
Volusia: Daytona Beach Shores, 3 Aug. I962, Ray 11159 (USF). Wakulla:
3 Nov. 1964, Godfrey 64887 (FLAS, FSU). Washington: Chipley, 31 July

1954, Ford & Arnold 386O (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 6I8 (F, MO, NY); Havana, Apr. 1903,
Shafer 91 (F). Isle of Pines: Nueva Gerona, 19 Dec, 1903, Curtiss 244

(F, NY), Oriente: Bayate, I8 July 191^, Ekman 2012 (NY), Pinar del Rio:
Herradura, 26-30 Aug. 1910, Britton, Britton, Earle & Gager 6332 (F, NY);
between Vinales and Pinar del Rio, I6 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. I9IO, 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: Negri 1 to Bloody Bay, 12 Sept. 1962,
Adams II663 (UCWl). Manchester: Lititz, 5 Nov, I96I, Adams 9864 (UCWl).
Portland: Port Antonio, Dec. I89O, Hitchcock s.n, (MO) . St, Andrew: Con-
stant Spring to Bardowie, 2 Aug, 1915, Harris 12112 (F, MO, NY, UCWl);

Sk.

Hope, 18 Aug. 1897, Harris 6803 (F, NY, UCWl); road to Constitution
Hill, 20 Oct. 1905, Harris 9035 (F, NY, UCWl); University of West Indies
campus, 4 June I963, Crosby, Hespenheide S- Anderson 60 (F). St, Catherine:
Bog Walk, 17 Dec. I89O, Hitchcock s.n. (MO). St. Thomas: Morant Bay, 6
Dec. 1959, Adams 5678 (UCWl); Port Morant, 2k Dec. I89O, Hitchcock s.n.
(MO). Westmoreland: Bluefields & vicinity, 6-7 Mar, I9O8, Britton £■
Hollick 19^3 (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); Pol nte-a-p I tre, 12 July I936, Stehle 497 (NY); Port
de Paix, 9 Jan. 1929, Leonard & Leonard 135^3 (NY); St. Michel de I'Atalaye,
1 Jan. 1926, Leonard 8501 (MO); San Michel, 5 Aug. 1905, Nash £■ Taylor
1417 (NY). DOMINICAN REPUBLIC. Barahona, Feb. I9IO, Fuertes l49 (F, NY).
San Juan: Rio Arriba, 9-14 Sept. 1946, Howard & Howard 8821 (GH, NY).
Santo Domingo: Villa Allagracia, 19 Jan. 1929, Ekman Hi 1229 (PUR). PUERTO
RICO. Guayani 1 la-Tal lahoa, 13 Mar. 1913, Shafer I976 (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, I9O6, 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, Manlve, 1893, Duss 2920 (NY); Martinique,
Case-Pi lote, I9OI, 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. -II Dec. 1957, Proctor 16812 (IJ).

_C. hyssopi fol ia (L.) Small may be distinguished from similar species
by its combination of usually glabrous stems, cyathia In a leafy dichasium,

85.

Map 13. Distribution of _C. hyssop i fol ia

\.^-— — I"

tfV'

Map ]k. Distribution of _C. 1as iocarpa

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 dichaslum. 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:
h]k. 1843. Type: Bot. Mus., Berlin, if extant. _E. hyperici fol ia L.
" lasiocarpa Kl." Griseb., Fl. Brit. W. 1. isl. 5^. 1859. (excl. syn.
Ba1b.).

E. hypericifol ia 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-^0 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 nim,
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

87.

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 m.n. 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 I963, Crosby, Hespenheide & Anderson
186 (DUKE, PUR); Liguanea, 7 Feb. I96O, Proctor 20564 (IJ, NY, PUR); Stony
Hill, 21 July 1912, Harris 11136 (NY, UCWl), St. Thomas: Easington
Bridge, 23 Mar. 1954, Proctor 8512 (IJ). Manchester: Warwick, 8 Nov. I962,
Proctor 22886 (IJ). St. Elizabeth: Unity Hill, Aug. I96I, Robertson 9136
(UCWl). HAITI. Port-au-Prince, Sept, 1919, Buch 1713 (U); St. Marc,
25-28 Feb. 1920, Leonard 2954 (NY); Petionville, 15-28 June 1920, Leonard
4967 (NY); St. Michel de I'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 I9OI, Freeman s.n. (UCWl); 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.
hyssop i fol ia do not approach. in most other characters there is a
close resemblance between the two.

Wheeler' s appl i cat ion of the name Euphorbia hyper i ci fol ia L. to
this species was based on a mi s ident i f led specimen in the Linnaean
Herbarium, which was not seen or annotated by Linnaeus until long after
the publication of the name.

12. CHAMAESYCE PARC! FLORA (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 !).

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 g lands, 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.

Map 15. Distribution of _C. parci flora

Map 16. Distribution of _C. br i tton i i

90.

almost equalling lobes, sinus shallow. Staminate flowers 4-12 per
cyathium, androphores glabrous, to 1.3 n\m, 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 63^ (IJ); M, Bonfiere, 29 Sept. 1925, Ekman
H7627 (IJ); Petionville, 24 Aug. 1924, Ekman H1593 (U); 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. hyssopi fol ia 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 BRITTONI! (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 paucipi la (Urb.) Millsp., Field Mus. Bot. 2: 410. I9I6.
Euphorbia paucipila Urban, Symb. Ant., 5: 389. 1908. Type: Prope
Guanabacoa, Habana, Cuba, August, Baker and Hasselbring 7199 (Holotype:

91.

Herb. Estac. Agron,, Havana, Cuba, not seen).

Euphorbia niqueroana Urban in Fedde, Repert. 28: 23^. 1930.
Type: South of Nlquero, peninsula de Cabo Cruz, Oriente, Cuba, I? Jan.
1923, E. L. Ekman I6I76 (Holotype: Mus. Bot. Stockholm, not seen, photo-
graph of holotype and Isotype: NY !). Chamaesyce niqueroana (Urb.) Alain,
Contr. Ocas. Mus. Hist. Nat., Col. 'de la Salle', Habana 11: 12. 1952.

Perennial; erect or ascending to 1.5 dm. Stem to 1 mm. diameter
at base, internodes to 2 mm. long, sparsely branched throughout, sparsely
long-pilose particularly at nodes, brown. Leaves membranous; blades
ovate-elliptic or somewhat orbicular, 2-5 x 1.5 - 3 mm., base inequilateral,
rounded, margin entire or rarely crenate, apex rounded to obtuse, glabrous,
or with scattered long hairs, green; petiole 0.5 mm. long; stipules sepa-
rate, triangular, to 0.5 mm. long, toothed or 2-3 cleft. Cyathia solitary
at upper nodes; peduncles to 0.3 mm. long, involucre campanulate, to 0.7
mm. diameter at mouth, lobes equalling glands, triangular, toothed, glab-
rous on outside, ciliate at mouth within, glands broadly transversely
elliptic, to 0.2 mm. long, cupped and somewhat stipitate, appendages a rim
to twice as wide as gland, fifth gland linear, short, sinus shallow.
Staminate flowers 5-10 per cyathia, androphores glabrous, to 0.7 mm. long
at maturity. Gynophore glabrous to 0.8 mm. long at maturity, exserted,
nutant; calyx a triangular pad of tissue; ovary glabrous; styles spreading,
to 0.3 mm. long, minutely joined at base, bifid for 1/3 length. Capsule
glabrous, broadly ovoid with somewhat flattened base, 1.2 mm, long, 1.5
mm. wide just above base, broadly three-lobed, angles acute. Seed broadly
conical-ovoid, 0.8 mm. long, 0.6 mm. wide below equator, four-angled, dor-
sal faces larger than ventral, dorsal angle more acute than others, brown.

92.

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2| ' 3| '4 ' 6

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Plate 17. CHAMAESYCE BRITTONll (Millsp.) Millsp.

Isotype: E. bri ttoni i Mi 1 1 sp. , Race course, New Providence, Bahamas, 26
August 1904, Britton & Brace 839 (NY).

93,

Sandy areas and barrens of several Cuban provinces, and from
whitelands of New Providence, Bainamas (Map 1 6) ,

Specimens seen: BAHAMAS. New Providence, near race course,

26 Aug. 1904, Britton & Brace 279 (NY). CUBA. Havana, Loma de la Cruz,

27 Oct. 1927, Leon I3O89 (NY). Las Villas: west of Santa Clara, 1-20
July 1950, Howard, Briggs, et al. 4l6-b (A). Matanzas: Corral Nueva,
13 Oct. 1927, Leon I3IOI (NY); San Miguel de los Banos, 10 Oct. 1950,
Alain A 157^ (NY). Oriente: Guantanamo Bay, 17-30 Mar. I9O9, Britton
2159 (NY). Santa Clara: Loma de Belen, 2k June 1932, Leon I5628 (NY);
Placetas del sur, 10 Aug. I9I8, Leon S-Roca 8l45 (NY); Sti. Spiritus,

27 Aug, 1909, Leon 1352 (NY); Serpentine Barrens near Santa Clara, June
1941 , Howard 5106 (GH).

The three species, which are here combined, vary in the degree of
pubescence which they show but have no characters on which to base a
consistent separation. The group differs from the least pubescent forms
of _C. turp in i i only in habit and in the pubescent capsule of that species,
and f rom _C. i nsulaesal is only in its complete lack of pubescence. When
more collections are available, the species limits set here may well need
revi s Ion.

14. CHAMAESYCE COWELLII Mi lisp, in Britton, Bull. Torr. Club 43:
457. 1916. Type: Cayo Muertos, Puerto Rico, 9-12 Mar. 1915, Britton,
Cowell & Brown 5007 (Holotype: F 1; Isotype: NY !).

Perennial from a thickened rootstock; prostrate, forming mats to
15 cm. diameter. Stems many, wiry, ca. 0.5 mm. diameter, Internodes to
1 cm.; branching throughout length, nodes not rooting, glabrous, brown or
ashen. Leaves chartaceous; blades ovate-elliptic, somewhat reniform near

9^.

base of stem, 2-5 x 1-3 mm,, base oblique, cordate or obtuse, margin entire,
apex emarginate, rounded or obtuse, glabrous, obscurely papillose, green
or ashen; petiole 0.4 - 0.6 mm. long; stipules joined at base, 0,3 mm,
long, deeply cleft, sometimes ciliate on surface near stem, Cyathia
solitary at upper nodes; peduncle short, involucre obconical, to 0,6 mm.
diameter at mouth, lobes equalling glands, deltoid, glabrous on outside,
ciliate within, glands transversely elliptic, 0.2 mm. long, somewhat
cupped, appendages a rim to slightly wider than gland, fifth gland short
or obsolete, sinus to one-third depth of involucre. Staminate flowers
6-]k per cyathia, androphores glabrous, to 0.8 mm. long at maturity. Gyno-
phore glabrous, to 1 mm. long at maturity, exserted, nutant; calyx obsolete;
ovary glabrous; styles spreading, to 0,2 mm. long, joined at base, bifid
for half length. Capsule glabrous, ovoid, 1.2 mm. long, 1.4 mm, wide at
equator, sharply three-lobed, angles acute. Seed long-ellipsoid, to 1 mm.
long, strongly four-angled with ventral angle somewhat rounded, other
angles acute, surface slightly convex, obscurely wrinkled, tan or darker
brown.

Found in crevices of coastal limestone cliffs, Antigua and Cayo
Muertos, Puerto Rico (Map 17).

Specimens seen: PUERTO RICO. Sardinera, Mona Island, 20-26 Feb,
1914, Britton, Cowell & Hess 1670 (NY). LESSER ANTl LLES. Antigua: Soldier
Point, Half-Moon Bay, 1 May 1938, Box 1426 (NY); near English Harbor, 18
July 1937, Box 900 (BM, NY) ,

The resemblance between this species and C. turpinii in all charac-
ters except the pubescence of the latter suggests a close relationship.
Future collections may indicate that the two should be separated at only
the varietal level, but until more material is available the existing names

95.

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CATO MuCKTOt

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Plate 18. CHAMAESYCE COWELLII MiHsp. in Britton

Isotype: Z. cowel 1 i i Millsp., Cayo Muertos, Puerto Rico, 9-12 March 1915,
Britton, "Cowe 1 1 & Brown 5007 (NY).

96.

A^^-

Map 17. Distribution of C. cov/e 1 li i

Map 18. Distribution of C. blodqetti

97.

will be malnta ined.

There is a description of the species filed with the type
specimen in F which was probably drawn up by Millspaugh and sent with
the name to Britton for the publication. The citation was made in the
form that reflects this authorship.

15. CHAMAESYCE BLODGETTI I (Engelm. ex Hitchc.) Small, Fl. SE.
US. 712. 1903. Euphorbia blodgett i i Engelm. ex Hitchc, Ann. Rep. Mo.
Bot. Gard. 4: 126, I893. Type: Key West, Florida, Blodgett s.n. (Lecto-
type: MO '. , fragment F !, isotype: NY 1).

_E. bermudiana Millsp., Field Mus. Bot. 2: 65. 1900. Type: Walsing-
ham, Bermuda, 31 Dec. I898, Millspaugh 6OIOI (Holotype: F i, Isotype: F I,
NY i).

E_. cozumelensis Millsp., Field Mus. Bot. 2: 61. I9OO. Type:
East Shore, Cozumel Island, 21 Feb. I899, Millspaugh 6I6O6 (Holotype: F !).

jC. nashi i Small, Fl. SE. US., 709. I9O3. Type: Tampa, Hillsborough
County, Florida, 20 August 1895, Geo. V. Nash 2427 (Distributed as Euphor-
bia maculata) (Holotype: NY 1, Isotypes: F i, GH 1, US i).

Chamaesyce chiogenes Small, Fl. SE. US. 709. 1903. Type: Key
West, Florida, Blodgett s.n. (Holotype: Herb. Columbia College, if extant;
most of this herbarium now in NY, but sheet there not marked with Columbia
stamp, ? isotype 1; portion of isotype F !).

_E. batabanensis Urb., Symb. Ant., 5: 390. I9O8. Type: Prope
Batabano, prov. Havana, Cuba, October, Baker £• Wilson 2212, 2341, 2375
(Syntypes: Herb. Bot. Mus. Berlin, if extant; Baker & Wilson 2301 NY '. -
mistake in number?).

Annual, sometimes persisting; prostrate to decumbent forming mats

to 6 dm. diameter, rarely erect to 3 dm. Stem to 2 mm. diameter, larger
in single-stemmed erect plants, internodes to 2 or rarely 3 cm. long;
branching throughout length, sometimes rooting at nodes, glabrous, green
or brov/n, often with reddish cast. Leaves membranous; blades oblong-
elliptic k-6 X 2-3 mm., base inequilateral, rounded or subcordate, margin
entire or toothed in upper third, apex mucronulate or obtuse, blades of
leaves of laterals smaller, uniform, base usually cuneate, glabrous,
green, often suffused red; petiole to 1 mm.; stipules joined on upper
surface to I mm. long, deeply bifid with each half several-cleft or
toothed, on lower surface longer, triangular, several-cleft, sometimes
deeply bifid, light-colored. Cyathia solitary at upper nodes or appearing
clustered on congested leafy laterals; peduncle ca. 0.5 mm. long, involucre
campanulate, to 0.5 nim. diameter at mouth, lobes equalling glands, tri-
angular, entire or toothed, glabrous on outside, pilose within, glands
transversely elliptic-oblong, 0.2 x 0.05 mm., appendages obsolete or semi-
lunate, to twice as wide as glands, the two nearer the sinus larger than
the other pair, white or pink, fifth gland linear, short, sinus shallow.
Staminate flowers 8-12 per cyathium, androphores glabrous, to 0.5 mm. long
at maturity. Gynophore glabrous, to 0.7 mm. long at maturity, exserted,
nutant; calyx a three-cornered pad of tissue; ovary glabrous; styles
spreading, to 0.3 mm. long, minutely joined at base, bifid for 1/3 to 1/2
length. Capsule glabrous, conical-ovoid, truncate at base and apex,
1.2 - 1.4 mm. long, 1.4 - 1,6 mm, wide above base, roundly three-lobed,
angles acute to obtuse. Seed oblong-ovoid, to 1 mm. long, equally four-
angled with rounded angles, faces flat or slightly convex, obscurely wrin-
kled, brown or gray-brown.

99.

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Plate 19. CHAMAESYCE BLODGETTIi (Engelm. ex Hitchc.) Small

Lectotype: E. blodgett i i Engelm. ex Hitchc., Key West, Florida,
Blodgett s.n. (MO).

100,

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Plate 20. CHAMAESYCE BLODGETTII (Engelm. ex Hitchc.) Small

Holotype: C_. nashi i Small, Tampa, Hillsborough County, Florida, 20 August
1895, Geo. V. Nash 2if27 (NY).

101

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Plate 21. CHAMAESYCE BLODGETTII (Engelm. ex Hitchc.) Small
?|sotype: C. chiogenes Small, Key West, Florida, Blodgett s.n. (NY)

102,

A very common component of beach scrub and dune vegetation, and
also sandy inland areas, in the southern half of Florida, Bermuda, the
Bahamas, Cuba, Jamaica and the Caymans. Only one specimen has been
seen from the Virgin Islands, and the absence of the species from
Hispaniola south through the Antilles is in striking contrast with its
abundance in the other islands (Map 18).

Representative specimens seen: FLORIDA. Brevard: Patrick AFB,
24 Sept. 1957, Crawford s.n. (FLAS) ; St, Johns River valley west of Mims,
24 June 1957, Krai S- Sincock 5008 (FSU). Charlotte: Bermont, 7 Nov. 1964,
Hodgson 213 (FLAS); Charlotte Harbor, 21 Nov. 1964, Godfrey 65361 (FLAS,
FSU), Collier: Naples Beach, 21 Nov. 1964, Godfrey 6544! (FLAS, FSU) ;
Tamiami Trail, 1 Apr. 1930, Moldenke 869 (NY); Vanderbilt Beach, 13 Aug.
1963, Burch 184 (FLAS). Dade: Adams Key, 11 Mar. 1915, Small £• Hosier
5729 (NY); Biscayne Bay, Indian River, 1874, Palmer 500 (NY); Cutler to
Point Creek, 1 July I9I5, Small, Hosier £■ Small 6689 (HO, NY, US);
Homestead Bayfront Park, 20 July 1964, Lakela 27316 (FLAS, USF) ; Hiami
Beach, 11 Jan, 1930, Holdenke 390 (DUKE, HO, NY). Glades: Moore Haven,
28 Dec, 1963, Burch 236 (FLAS). Hendry: Big Cypress Seminole Reser-
vation, 8 June 1964, Brass 33229 (FLAS); near Clewiston, 9 Apr. 1950,
Killip S- Svyallen 40498 (US). Hernando: County line N.E. of Brooksville,
15 Oct. 1964, Burch 449 (FLAS). Highlands: Junction FLA 70 & FLA 17, I8
Nov. 1964, Burch S- Ward 527 (FLAS). Hillsborough: Tampa airport, 7 Sept.
1953. Garrett s.n. (FLAS). Lee: Punta Rassa, 24 June I9I6, Standley 274
(MO, US). Levy: mouth of Wi thl acoochee River, I6 Sept. 1959, Cooley,
Eaton Sr V/ard 6984 (DUKE, FLAS, FSU). Manatee: north end Longboat Key,
1 Aug. 1950, Wilbur £- Webster 2509 (US); Manatee, 11 Sept. 1899, Tracy

103.

6374 (F, MO, NY); Sneeds Island, 12 Sept. 1899, Tracy 6373 (F). Monroe:
Big Pine Key, 27 Feb. 1911, Small, Carter & Small 3563 (NY); Flamingo,
Cape Sable, 26 Nov, I9I6, Small 799^ (NY); Key West, I836, Blodgett s.n,
(MO); Lower Matecumbe Key, 18 Mar. 1930, Moldenke 784 (DUKE, MO, NY);
Pine Crest, 1 Apr. 1930, Moldenke 853 (DUKE, F, MO, NY, US); Upper
Matecumbe Key, Curtiss 2487 (FLAS, GH, MO, NY, US). Okeeclnobee: 3 miles
east of Okeechobee City, 3 Feb. 1945, Brass 14593 (US). Pinellas: Maximo
Point, St. Petersburg, 10 Sept. 1954, Thorne 15411 (FSU, IJ, NCS, US);
north end of Sunshine Skyway Bridge, St. Petersburg, 5 June 1964, Burch
359 (FLAS). St. Johns: Crescent Beach, 14 Oct. 1941, West & Arnold s.n.
(FLAS). Sarasota: Longboat Key, 27 Sept. 1964, Burch 426 (FLAS); 2 miles
south of Venice, 30 Dec. 1956, Krai 3833 (FSU). Volusia: 9 miles south of
New Smyrna Beach at Turtle Mound, 27 Apr, I96I, Ray, Wood et al. IO78O
(FSU), BAHAMAS. Grand Bahama: 3 miles SE of Freeport, 1 Nov. 1964,
McDaniel & Schmiederer 5428 (FSU); Eight Mile Rocks, 5-13 Feb. I905,
Britton £■ Millspaugh 2470 (F, NY). Great Sturrup Cay, 1 Feb. 1905, Britton
& Millspaugh 2257 (F, NY); Bimini, 16 Apr, 1904, Millspaugh 2204 (F, NY);
Andros, I8 Aug. -10 Sept. I9O6, Brace 5115 (F, NY). New Providence: Ocean
Hole, Coral Harbor, 7 July I96O, Webster £• Williams 10328 (PUR); Hog
Island, 30 May I9O9, Wilson 8259 (F, MO, NY). Eleuthera, Glass Window,
26 July i960, Webster & Williams IO636 (PUR); Salt Key Bank, Anguilla
Isles, 15-18 May 1909, Wilson 7946 (MO, NY); Exuma Chain, Ship Channel
Cay, 17 Feb. 1905, Britton S- Millspaugh 2755 (F, NY); Little San Salva-
dor, 25-26 Feb. 1907, Britton & Millspaugh 566O (F, NY); Watling's Island,
27-28 Nov. 1907, Wilson 7286 (F, NY); Crooked Island, 24 Nov. I89O, Roth-
rock 241 (F, NY). Inagua: South of Matthewtown, 15-25 Aug. I96O, Williams

104.

102 (DUKE, iJ, PUR); 8 miles N.E. of Matthewtown, 15-25 Aug. I96O,
Williams 37 (DUKE, PUR). South Caicos: Cockburn Harbour, 26 June 195^,
Proctor 8938 (IJ); East Bay, 24 June 1954, Proctor 8915 (U). Grand
Turk, between Commissioner's House & airfield, 22-27 Apr. 1954, Lewis
s.n. (IJ). CUBA. Camaguay: Cayo Cruz, 27 Oct. 1909, Shafer 2783 (F,
NY); Cayo Paredon Grande, 25 Oct. 1909, Shafer 2756 (F, NY). Havana:
Batabano, 10 Apr. I903, Shafer 514 (NY). Isle of Pines: Vivijagua,
20 Feb, 1916, Britton, Britton & Wilson, 14692 (F, NY). Pinar del Rio:
Coloma, 6 Sept. 1910, Britton, Britton £■ Gager 7007 (F, NY). Santa
Clara: Rio San Juan, 24-25 Mar. I9IO, Britton, Earle & Wilson 5915 (F,
NY). CAYMAN ISLANDS. Grand Cayman: north of Georgetown, 1 June I963,
Crosby, Hespenheide & Anderson 18 (DUKE, UCWl); North Side village, 19
Apr. 1956, Proctor 15123 (iJ); Bodden Bay Road, 13 Feb. 1899, Millspaugh
1333 (F); Jan. I89I, Hitchcock s.n. (MO); Spot Bay, 13-14 Feb. 1899,
Millspaugh 1314 (F). JAMAICA. Clarendon: between Portland & Rocky
Point, 5 Mar. I908, Britton I9II (F, NY). Kingston: Palisadoes, near
Harbour Head, 4 Oct. I96O, Proctor 21392 (IJ, NY, PUR). St. Andrew:
Palisadoes, 5 Nov. 1957, Yuncker 17299 (F, MO) . St. Catherine: Cayamanas,
June 1952, Asprey 437 (UCWl); Ft. Clarence Hill, 9 Dec. I9O6, Harris
9536 (NY, UCWl). St. Elizabeth: Great Pedro Bay, 6-8 Sept. 1907,
Britton 1250 (F, NY); Pedro Bluff, 7 Sept. 1907, Harris 9731 (F, NY,
UCWl). Westmoreland: West Point lighthouse, 26 July 1954, Webster £-
Wilson 5057 (IJ). LESSER ANTILLES. Tortola, Lower Estate, 8 Oct. I9I8,
Fishlock 222 (NY). Extra-territorial specimens: BERMUDA. Hamilton Bay
Islets, 29 Dec. I888, Millspaugh 9 (F) ; Jew's Bay, 23 July 1913, Collins
332 (F, NY); North Shore, 31 Aug. -20 Sept. 1905, Brown & Britton 4

105.

(F, NY); Warwick March, Sept. 1913, Brown £■ Britton 1719 (F, NY).

Hitchcock described this plant after collecting it a number of
times in the West Indies, adopting a name used by Engelmann on herbarium
sheets but not published. A specimen in Herb. Torrey and Herb. Engelmann
is cited as type, and the article is illustrated with a plate in which
the habit drawing was made from the upper right hand plant on a sheet
which he collected in Grand Cayman in I89I (MO i). Detail on the plate
is from sketches probably made by Engelmann.

The sheet from Herb. Engelmann (now Mo. Bot. Gard.# 1792993) was
probably made from a specimen sent by Torrey for identification, and may
well have come from his much fuller sheet which is now in NYBG. it seems
likely that only Engelmann's sheet would have been readily available to
Hitchcock, and it is designated as lectotype for that reason. The sheet
nov; in NYBG is an isotype since it is from the same collection.

The sheet from Millspaugh's private herbarium in F includes two
numbers. The origin of 196079 is not obvious, but the label data clearly
indicates that I96078 is fragments from the Herb. Engelmann specimen and
may be considered part of the lectotype.

The variability of this species is reflected in the number of
attempts which have been made to segregate portions of it as discrete
taxa. None of the criteria by which Millspaugh, Small and Urban distin-
guished their species holds when a long series Is examined, but there
appear to be certain trends which may merit recognition if this seems
desirable at a later date.

The habit of the plant is most commonly prostrate or decumbent,
with several stems from the crown of the root. In Cuba, In Florida on

106.

the coast south of Miami, and particularly in the islands on the Atlantic
edge of the Bahama chain, there are plants of this species which are
strongly ascending or completely erect with one or a few stems from ground
level. In all other respects - stipules, leaf-shape, and cyathlal and
seed characters - they fall within the limits of variability usual for
the species, and often grow interspersed with "normal" plants.

Branching pattern shows a similar break from the usual, rather
sparingly branched form with cyachia on short laterals, to a much-branched,
almost dichotomous form with cyathia solitary and spaced rather widely.
Leaf size is often small in these plants. Others show an extreme reduction
in laterals, and, with their many small leaves, have a superficial resem-
blance to _C. maculata. Branching on some plants shows more than one of
these patterns, but in some areas, such as Hendry and Glades counties in
Florida, the much-branched habit with small leaves is almost the only form
found.

The form distinguished by Small as _C. chiogenes is quite widespread
in dry, saline areas. The plants have a more chartaceous texture, with
the stipules reduced in size, but no way of making a positive identification
of the species was found, and it seemed more in keeping with treatment
elsev/here in the group to reduce it to synonymy.

Chamaesyce blodgettii comes close to C_. serpens in many characters,
but can be distinguished by the obscurely wrinkled seed even when the
difference in habit and size is less clear than usual. The two occupy
a similar set of habitats in their respective ranges, which overlap only
in Cuba and Jamaica.

w

107.

16. CHAMAESYCE SERPENS (HBK.) Small, Fl, SE. US., 709. I9O3.
Euphorbia serpens HBK., Nov. Gen. et Sp., 2: 52. 1817. Type: Cumana,
Venezuela, Bonpland 407 (Herb. Mus. Paris, not seen, photograph G and
fragment F seen by Wheeler, Rhodora 43: I98. 1941.). Anisophy] 1 urn
serpens (HBK.) Kl. S- Gke., Abh. Akad. Berlin, Rhys. 1859: 23. i860.

Euphorbia crassinodis Urb., Symb. Ant., 1: 3^0. I899. Type:
Cuba, Wright 54-7 p.p., 2014 (Herb. Mus. Bot., Berlin, if extant; sheets

i th these collection numbers in GH i, but both are _E. gundlachi i (Urb.)).
C. crassinodis (Urb.) Millsp., Field Mus. Bot. 2: 408. I9I6. Placed
here in synonymy on the basis of other sheets identified as this species
by Urban.

_E. pi leoides Millsp., Field Mus. Bot. 2: 62. 1900. Type: Guanica,
Porto Rico, 22 Jan. I899, C F. Millspaugh 743 (Holotype: F. not found;
Isotype: F '.) . C. pi leoides (Millsp.) Millsp., Field Mus. Bot. 2: 411.
1916.

_E. microclada Urb., Symb. Ant., 9: 215. 1924. Type: prope
Daiquiri, Oriente Province, Cuba, 23 Nov. I9I6, E. L. Ekman 8430 (Holo-
type: Mus. Bot. Stockh. , not seen; Isotype NY i). _C. microclada (Urb.)
Alain, Contr. Ocas. Mus. Hist. Nat., Col.'de la Salle', Habana 11: 12.
1952.

E.. excisa Urb. & Ekman, Arkiv. Bot. Stockh. 22A(8): 66. 1929.
Type: Anse-a-Pi tre, Haiti, 15 Sept. 1926, E. L. Ekman H 6986 (Holotype:
Mus. Bot. Stockh., not seen; Isotypes: iJ i, K 1, NY !).

_E. biramensis Urb. in Feddes, Repert. 28: 232. 1930. Type:
Cienaga de Birama, Oriente Province, Cuba, 10 Jan. 1923, E. L. Ekman
16137 (Holotype: Mus. Bot. Stockh., not seen; Isotypes: NY '. , US i).

108.

_C. bi ramensis (Urb.) Alain, Contr. Ocas. Mus. Hist. Nat., Col.'de la
Sa.le', Habana 11: 12. 1952.

_E. manqjeti Urb. in Feddes, Repert. 28: 233. 1930. Type:
Castillo de Atares, Habana, Cuba, 2 Oct. 1921, E. L. Ekman 13293 (Holo-
type: Mus. Bot. Stockh., not seen; Isotype: NY i). _C. manqlet i (Urb.)
Alain, Contr. Ocas. Mus. Hist. Nat., Col.'de la Salle', Habana 11: 12.
1952.

The unpublished name _E. ant iquens is used by Millspaugh on a plant
collected in Antigua, 16-17 Jan. 1907, Shafer 20 (NY ;, F 1) should be
referred here.

Perennial or winter killed; prostrate, forming mats to k dm,
diameter. Stem several from somewhat thickened root, to 1 mm. wide,
internodes to 3 cm. long; branching throughout length, often rooting at
nodes, glabrous, yellowish-green. Leaves membranous; blades ovate, some-
times orbicular, 2-6 x 1.5 - ^ mn^.. , base inequilateral, rounded to sub-
cordate, margin entire, apex rounded, mucronulate or emarginate, midrib
prominent on underside, glabrous, green; petiole ca. 0,5 mm. long; stipules
united, deltoid, toothed at apex, to 0.5 mm. long, white. Cyathia soli-
tary at upper nodes; peduncle to 1 mm., involucre campanulate, to 0.7 mm.
diameter at mouth, lobes equalling glands, deltoid, toothed, glabrous on
outside and within, glands transversely elliptic to 0.2 mm. long, usually
deep purple, sometimes brown or olive, appendages longer than glands and
narrower to twice as wide, margin entire or crenulate, white, fifth gland
linear equalling lobes, sinus shallow. Staminate flowers 3-10 per
cyathium, androphores glabrous, to 0,5 mm. long at maturity. Gynophore
glabrous, to 0.7 mm. long at maturity, exserted, nutant; calyx three-lobed,

i09.

Plate 22. CHAMAESYCE SERPENS (HBK.) Small

C serpens (HBK.) Small, Vicinity of Port de Paix, Haiti, 1 May 1929,
leonarcl & Leonard 15242 (NY).

no.

each lobe often toothed and recurved, to 0.3 nim. long; ovary glabrous;
styles spreading, 0.2 mm. long, joined at base, bifid for 1/2 length.
Capsule glabrous, ovoid, 1.2 - 1.4 mm. long, 1.4 - 1.6 mm. wide below
equator, three-lobed, angles obtuse. Seed oblong-ovoid, to 1 mm. long,
equally four-angled with rounded angles, faces flat or slightly convex,
smooth, usually brown or gray-brown.

A common plant of the Mississippi valley and of scattered occur-
rence in most states east of the Rocky Mountains. Collected only as a
ballast plant in Florida, and the last collection from the site in
Pensacola was in 1897. Present in Cuba, Jamaica, Hispaniola, Puerto
Rico and islands of the Lesser Antilles to South America (Map 19).

Representative specimens examined: FLORIDA. Ballast ground,
Pensacola, Escambia County, 2 July 1897, Curtiss 5920 (MO, FLAS, NY, US);
Apalachicola, ex herb. Chapman, no further data (MO), CUBA. Oriente
Province. Baitiquiri 27 Dec. 1954, Alain £• Figueras 7 (OH); Daiquiri,
14-16 Mar, 1912, Britton & Cowell 12692 (F, NY); near Cajobabo, 12 Jan.
1956, Alain £• Morton 5015 (GH), Habana Province. Vedado, Feb. 1948,
Clemente 5936 (NY). JAMAICA, St, Thomas, 3 July 1964, Powell & Green
1634 (IJ); St. Catherine, 4 Feb. 1953, Proctor 7607 (U). HAITI. Gonave
island, Aug. 1927, Eyerdam 228 (F, GH, MO); near Hervee, 27 Feb. 1925,
Ekman 3341 (NY); Gonaives, 24 Dec. 1927, Ekman 9453; vicinity Port-de-Pa ix,
9 Jan. 1929, Leonard & Leonard 11776 (GH) ; vicinity Port-de-Pa ix, 1 May
1929, Leonard & Leonard 15242 (NY), DOMINICAN REPUBLIC. Province of
Barahona, I9IO, Fuertes 134 (NY), Fuertes 485 (F, GH, MO, NY), 1911
Fuertes 970b (GH) , Fuertes 992b (GH). PUERTO RICO. Paraquera, 24 Feb.
1924, Britton & Britton 8044 (GH, NY); Aguirre, 14 Feb, 1922, Britton,

Ill

Britton & Brown 6030 (F, NY); San Juan, 14 June-22 July 1901, Underwood
& Griggs 913 (F, NY). LESSER ANTILLES. St. Thomas, 31 Jan. -4 Feb. I913,
Britton, Britton & Shafer 5 (F, MO, NY); St. Croix, 17-25 Mar, I923,
Britton, Britton & Kemp 132 (NY); Mar ie-Ga lante, 26 Mar. I936, Stehle
480 (NY); Martinique, k July I939, Egler 39-65 (NY); Martinique, 11 Jan.
1939, Stehle 3466 (GH) ; Martinique, April I903, Duss 4722 (F, NY); Barbados,
Apr. -June 1895, Waby 94 (F) ; St. Lucia, 27 Nov. I96O, Proctor 2I638 (IJ);
Cannuoan island, 26-31 Mar. 8-20 Apr. 1950, Howard IIII3 (FLAS, GH, IJ,
NY); Grenada, 24 Feb. -6 Mar. 1950, Howard 10547 (GH, IJ).

There seems no reason for the absence of this species from Florida
but an intensive search in most parts of the state has failed to find any.
The plants from the United States usually have brown or pale glands
rather than the deep purple type found in the West Indies, but this is
not considered significant. The slight tendency to enlargement of the
nodes which occurs throughout the range is particulary pronounced in the
plants which Urban named _E. crass i nod i s and _E. exci sa , and _E, exci sa also
has a higher frequency of leaves with an emarginate apex than is common
in the remainder of the range. Both species tend to be of a somewhat
heavier texture than the average run of plants, but in spite of these
differences, all specimens examined appear to fall within the range of
variation expected from the species.

The species is interesting for the marked development of the calyx,
which forms a toothed and recurved three-lobed structure, much more
prominent than in any other species examined.

17. CHAMAESYCE CENTUNCULOI DES (HBK.) Mi lisp.. Field Mus. Bot. 2:
408. 1916, Euphorbia centuncu loides HBK., Nov. Gen. et Sp. 2: 51. 1817.

1 12,

Map 19. Distribution of C. seroen5

AS^-;-

>-sa-^..

Map 20. Distribution of C, centuncu lo i des

113.

Type: Prope Guanavacoa et portum Havanae, Cuba, (Herb. Mus. Paris, not
seen). Fragments of collections by Poeppig "in maritimus Cubae" are In
MO(i) and F(l) - the fragments are from Herb. Martius, Herb. Univ. Prague
and Herb. Bernhardi, so the collection must have been widely distributed,
and may have been seen by Kunth. There is also a fragment in MO(i) of a
collection from Havana by Humboldt. These are regarded as supporting the
use of the name which is maintained here, but their status as possible
type material cannot be decided until Herb. Mus. Paris has been examined.

Euphorbia bal bi s i i Boiss,, Cent. Euph., 11. i860. Placed here in
synonymy on the basis of the description by Boissier in DC. Prod. 15(2):
38. 1862.

Chamaesyce insulaesalis MiHsp., Field Mus. Bot. 2: 395. 191^.
Type: Sandy places, Salt Key, Salt Key Bank, Bahamas, 21 May 1909, P.
Wilson 8082 (Holotype: F, not seen: Isotypes: MO i, NY i).

Euphorbia filicaulis Urban, Symb. Ant., 9: 217. 1924. Type:
Cerro de Fraile prope Holguin, Oriente, Cuba, 28 Oct. 191^, E. L. Ekman
3237 (Holotype: Mus. Bot. Stockholm, not seen; Isotypes: K i, NY i, US 1).
Chamaesyce f i 1 icaul is (Urb.) Alain, Contr. Ocas. Mus. Hist. Nat., Col.'de
la Salle' , Habana II: 12. 1952.

Euphorbia pachypoda Urban, Symb. Ant., 9: 216. 1924. Type: Prope
Vedado, Habana, Cuba, 21 April 1914, E. L. Ekman 146 (Holotype: Mus. Bot.
Stockholm, not seen; photograph of holotype, and Isotype: NY !). Chamaesyce
pachypoda (Urb.) Alain, Contr. Ocas. Mus. Hist. Nat., Col.'de la Salle',
Habana 11: 12. 1952.

Perennial; prostrate forming mats to 2 dm. diameter. Stems many
from crown of root, wiry, 0.3 mm. diameter, internodes to 1.5 cm.;

114.

branching throughout length, sparsely at base, not rooting at nodes,
glabrous, straw-colored or brown. Leaves membranous; blades elliptic,
sometimes broadly so, to subcordate, 2-5 x 1-3 mm., base oblique,
rounded to subcordate, margin entire or crenate in lower leaves of stem,
apex rounded, midrib sometimes prominent on lower surface of leaf,
glabrous, green; petiole ca. 0.3 mm. long; stipules joined, sometimes
only at base, triangular to 0.2 mm., variously toothed or deep cleft.
Cyathia solitary; peduncle to 0.5 mm., involucre broadly obconical, to
0.7 mm. diameter at mouth, lobes slightly exceeding glands, triangular,
entire, glabrous on outside, densely pilose within, glands transversely
elliptic to 0.2 mm. long, cupped, appendages as long as gland and twice
as wide, margin entire or lobed, yellowish, fifth gland linear, short
or obsolete, sinus shallow. Staminate flowers 8-20 per cyathium, andro-
phores glabrous, to 0.5 mm. long at maturity. Gynophore glabrous, to
1.3 mm. long at maturity, exserted, upright or nutant; calyx vestigial;
ovary glabrous; styles 0.2 mm. long, spreading, to 0.3 mm. long joined
at base, bifid for 1/3 - 1/2 length, slightly clavate. Capsule glabrous,
ovoid, 1.2 mm. long, 1.2 mm. wide between base and equator, roundly
three-lobed, angles obtuse to acute. Seed triangular-ovoid, 0.8 mm. long,
0.5 mm. wide below equator, strongly four-angled, ventral angle somewhat
obtuse, others acute, faces flat or slightly convex, obscurely wrinkled,
brown or white.

In sand and coastal rocks. Salt Key Bank and Salt Cay, New
Providence, Bahamas, and Cuba (Map 20).

Specimens seen: BAHAMAS. New Providence: Salt Cay, Feb. 1890,
Northrop & Northrop 277 (F, GH, NY). CUBA. Camaguey: Cayo Paredon Grande,

15.

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WILSON, COLLECTOR

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Plate 23. CHAMAESYCE CENTUNCULO I DES (HBK.) Mi lisp,

Isotype: C. insulaesal is Millsp., Salt Key, Salt Key Bank, Bahamas,
21 May 1909, P. Wilson 8O82 (NY).

lll|ljllll|llll|llll|IMI|lllilHH|IMI|IIH|HIYHt|)tWNI|IIHHtH|mi|Him
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Plate 2k. CHAMAESYCE CENTUNCULOI DES (HBK.) Mi lisp.

Isotype- E. filicaulis Urban, Cerro de Fraile prope Holguin, Oriente,
Cuba, 28 October 191^, E. L. Ekman 3237 (NY).

117.

25 Oct. 1909, Shafer 2753 (F, NY); E. part of Cayo Cruz, 27 Oct. I909,
Shafer 278I (F, NY). Havana: Guanabacoa, 27 Oct. 1927, Leon I3089 (NY).
Matanzas: Boca de Canasi, 7 Dec. 1927, Leon 13173 (NY). Oriente:
Sardlnero, June 19^8, Clemente CI, 6097 (NY); S.E. of Helguin, 7 Apr.
1909, Shafer 1215 (F, NY).

The plant called _C. i nsu 1 aesal i s by Millspaugh is of heavier
texture than is usual in the species, and _E. pachypoda Urban is inter-
mediate in this respect, but no features were found to justify their
separation. Very little material has been seen, and further collections
will allow a more sound assessment of the relationship of these plants.

18. CHAMAESYCE TORRALBASII (Urb.) Millsp., Field Mus. Bot. 2:
412. 1916. Euphorbia torralbasii Urban, Symb. Ant., 1: 339. 1899.
Type: prope Habana ad Vedado, Torralbas 93, Wright 2012 p.p. (Herb. Mus.
Bot., Berlin, if extant; Wright 2012 NY I, but a sheet of Wright 2012 in
MO(i) is not this species but _C. serpens) .

Euphorbia andromedae Millsp., Field Mus. Bot. 2: 63. 1900. Type:
Cape Corientes, Cuba, 17 Feb. 1899, Millspaugh 1466 (Holotype: F sheet
61466, not seen; Isotypes: F !, NY i).

Perennial; prostrate to 2 dm. Stems several to many from crown
of root, wiry or thicker to 0.5 mm. diameter, internodes to 1.5 cm. long;
branching sparsely, mostly at upper nodes, not rooting at nodes, glabrous,
straw-colored. Leaves membranous, sometimes slightly thickened; blades
ovate or somewhat cordate, 4-8 x 3-5 mm., base inequilateral, subcordate,
margin serrate, apex obtuse or rounded, glabrous, green or somewhat yellow-
brown on under surface; petiole to 1 mm. long; stipules joined, sometimes
only at base, subentire or variously parted, at times almost to base.

118.

Cyathia solitary at upper nodes; peduncle to 1 mm. long, involucre broadly
obconical, to 1 mm. diameter at mouth, lobes scarcely equalling glands,
triangular, entire or cleft, glabrous on outside, ciliate within, glands
transversely elliptic to 0.4 x 0.1 mm., cupped, appendages as long as
gland and to twice as wide, margin entire or crenulate, greenish-yellow,
fifth gland linear, short, sinus shallow. Staminate flowers rarely more
than 5 per cyathium, androphores glabrous, to 0.7 mm. long at maturity.
Gynophore glabrous, to 0.8 mm. long at maturity, exserted, nutant; calyx
obsolete or a triangular pad of tissue; ovary glabrous; styles upright,
to 0.3 mm. long, joined at base, bifid for 1/3 length. Capsule glabrous,
ovoid, 1.5 mm. long, 1.5 mm. wide just below equator, three-lobed, angles
acute. Seed oblong-ovate, to 0.9 mm. long, 0,5 mm. wide at equator, almost
equally four-angled, faces slightly convex, smooth or obscurely wrinkled,
brown.

Savannas and sandy areas from all parts of Cuba but not reported
from any other island (Map 21).

Specimens seen: CUBA. Camaguey: Si 11a de Cayo, Cayo Romano, 9-11
Oct. 1909, Shafer 2510 (MO, NY); Si 11a de Cayo, Cayo Romano, 9-11 Oct. 1909,
Shafer 25^3 (MO, NY). Havana: Sea shore, 18 Feb. 1905, Curtiss 652 (NY);
15 July 1928, Roig 4699 (NY); Playa del Vedado, 20 Dept. 1904, Baker I83O
(NY); Havana, Reparto biramar, 10 Feb. 1921, Ekman 13304 (NY); Cojimar, 24
Aug. 1910, Britton, Earle S- Gager 6279 (NY); on walls of Cabana Fortress,
Dec. 1922, Leon 11394 (NY); Morro Castle, Nov. I9O8, Leon 736 (NY); near
Playa de St. Anna, 2 Nov. 1915, Leon & Cazanas 5721 (NY); Playa de Marianao,
22 Feb. 1910, Britton 5- Wilson 4544 (NY). Pinar del Rio: Laguna Jovero and
vicinity, 5-7 Dec. I9II, Shafer 10707 (NY); Bay of Mariel, 21 Sept. 1910,

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plate 25. CHAMAESYCE TORRALBAS I I (Urb.) Millsp.

Duplicate Coll: E^. torra Ibas i i Urban, prope Habana ad Vedado, Cuba,
Wright 2012 p.p.~(NY).

Map 2

Distribution of _C. torralbasii

Map 22. Distribution of C. camaguayens i s

121.

Britton & Earle 7595 (NY). Matanzas: near mouth of Bueyvaca, 28 Aug.
1903, Britton & Wilson ^5 (NY); Boca de Canasi, 7 Dec. 1927, Leon 13173
(NY); 1905, Van Hermann 921 (NY); Playa, 12 Mar. I903, Britton, Britton
Sr Shafer 8O (NY).

Plants of this species are usually larger than those of _C. centun- -
culoides , and have all leaves serrate. The two species are very close,
and the distinction between them is not always absolute, but the strong
growing form represented in the type specimen is of frequent occurrence,
and is so consistent in appearance that it is felt that it merits recog-
nition as a spec ies.

19. CHAMAESYCE CAMAGUAYENS I S Millsp., Field Mus. Bot. 2: 392.
1914. Type: Near Camaguey, Cuba, 2-7 April 1912, N. L. and E. G. Britton
and J. F. Cowel 1 13233 (Holotype: NY !; Isotypes: F 1 , MO '.) . Euphorbia
camaquayensis (Millsp.) Urban, Symb. Ant., 9: 217. 1924.

Euphorbia pinariona Urb. in Fedde, Repert. 28: 23^. 1930. Type:
San Juan y Martinez, Pinar del Rio, Cuba, 18 June 1923, E. L. Ekman I6782
(Holotype: Mus. Bot. Stockholm, not seen; Isotype: NY i). Chamaesyce
pinariona (Urb.) Alain, Contr. Ocas. Mus. Hist. Nat., Col.'de la Salle',
Habana 11: 12. 1952.

Perennial, erect or ascending to 1 dm. Stems many from root-stock,
wiry, to 0.3 mm. diameter, internodes to 1.5 mm. long; scarcely branched,
glabrous, light brown. Leaves membranous; blades elliptic to suborb i cu lar ,
3-5 X 2-3 mm., base inequilateral, subcordate, margin entire or rarely
somewhat erose, apex rounded rarely obscurely mucronulate, midrib slightly
raised on underside, glabrous, rarely minutely and sparsely puberulous,
green; petiole ca. 0.5 mm. long; stipules joined, deeply 3-5 parted or

122.

bifid, to 0.3 nim. long. Cyathia solitary at upper nodes; peduncle
to 1.5 mm. long, Involucre broadly campanulate, to 1.5 mm. diameter
at mouth, lobes scarcely equalling glands, triangular, entire or toothed,
glabrous on outside, ciliate at mouth within, glands transversely
elliptic 0.5 X 0.2 mm., cupped, appendages obconical, longer than
glands and to three times as wide, fifth gland linear, shorter than
lobes, sinus to 1/3 depth of involucre. Staminate flowers 5-1^ per
cyathium, androphores glabrous, to 1 mm. long at maturity. Gynophore
glabrous, to 1.2 mm. long at maturity, exserted, nutant; calyx a tri-
angular pad of tissue; ovary glabrous; styles spreading, 0.3 mm, long,
joined at base, bifid for 1/3 length. Capsule glabrous, ovate-ellip-
soid, 1.3 mm. long, 1.5 mm. wide at equator, broadly three-lobed, angles
acute. Seed ovate, truncate at base, apex acute, 0.8 mm. long, ventral
angle rounded, dorsal angle acute, dorsal faces wide, slightly convex,
obscurely ridged, chestnut brown, sometimes overlaid white.

Savannas in most parts of Cuba, but not reported from any other
parts of the area (Map 22).

Specimens seen: CUBA. Camaguey: Sabana de la Matanza, 10 July
1948, Leon & Dahlgren 23398 (NY); Camaguey to Santayana, k Apr, 1909,
Britton 2394 (NY); Savannas near Camaguey, 2-7 Apr. 1912, Britton, Britton
& Cowell 13130 (NY). Isle of Pines: May I9IO, Jennings 6O8 (NY); La
Canada, 16 Feb. I9I6, Britton, Britton S- Wilson 14400 (NY). Oriente:
Vicinity of El Morro, Santiago Bay, 10-11 Mar, 1912, Britton 6- Cowell
12537 (MO, NY). Santa Clara: Cayo Frances, 22 Feb. 1924, Ekman 18575 (NY).

The prominent glandular-appendages of this species distinquish
it from all other Cuban species, but there are clear affinities with

123.

J

K

V.-.

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Plate 26. CHAMAESYCE CAMAGUAYENS I S Mi lisp.

Holotype: C. camaguayens i s Mi lisp., near Camaguey, Cuba, 2-7 April 1912,
Britton, sTitton & Cowell 13233 (NY),

124.

some of the other wi ry- stemmed plants of the island and Hispaniola and
the nearby Bahama groups.

20. CHAMAESYCE CORDIFOLIA (Ell.) Small, Fl. SE. US., 709, I903.
Euphorbia cordifol ia El 1 iott, Sketch Bot. South Carol ina £■ Georgia, 2:
656. 1824. Type: Beaufort, South Carolina (Charleston, South Carolina,
Museum) .

Annual; prostrate forming mats to 4 dm. diameter. Stems to 1.5
mm. diameter, internodes to 2 cm. long; branching throughout length, not
rooting at nodes, glabrous, brown. Leaves membranous; blades broadly
elliptic to orbicular or somewhat oblong, 3-8 by 2-4 mm., base inequi-
lateral, rounded or subcordate, margin entire, apex rounded or obtuse,
sometimes obscurely mucronate, midrib distinct on lower surface, glabrous,
green or red; petioles to 1.5 mm. long; stipules parted almost to base
into several filiform segments. Cyathia solitary at nodes of short
dichasia; peduncle to 1 mm. long, involucre campanulate, to 1 mm. diameter
at mouth, lobes exceeding glands, attenuate, toothed or ciliate, glabrous
outside and within or with cilia at mouth, glands transversely elliptic
to 0.3 rnm. long, appendages the same length as gland, up to three times
as wide, rarely obsolete, fifth gland linear, shorter than lobes, sinus
shallow, Staminate flowers 4-18 per cyathium, androphores glabrous, to

1 mm. long at maturity. Gynophore glabrous, to 3 mm, long at maturity,
exserted, upright or reflexed; calyx a triangular pad of tissue; ovary
glabrous; styles upright or spreading, to 0.5 mm. long, minutely joined
at base, bifid for 1/3 length. Capsule glabrous, ovoid, to 1.8 mm, long,

2 mm, wide just below equator, three-lobed, angles acute. Seed oblong-
ovoid, 1.2 mm. long, to 0.8 mm. wide below equator, equally four-angled.

125.

NT

Plate 27. CHAMAESYCE CORDIFOLIA (ED.) Small

_C. cordi fol ia (Ell.) Small, Sebring, Highlands County, Florida,
Ty May 19^8, Garrett 7 (FLAS).

126.

faces flat, smooth or obscurely wrinkled, brown or gray.

Locally frequent in sandy areas from North Carolina to Florida
and west to Texas. Known in Florida only from the northern counties
and from the central ridge, and apparently absent from the Caribbean
(Map 23).

Representative specimens seen: FLORIDA. Alachua: Gainesville,
21 Nov. 1939, Murrill s.n. (DUKE, MO); Gainesville, 9 Nov. 1959, West
s.n. (FLAS). Citrus: Hernando, 15 Oct. 1964, Burch 4^3 (FLAS). Colum-
bia: Lake City, 20 Dec. 1893, Rolfs s.n. (FLAS). Dixie: Along Suwannee
River E. of Oldtown, 14 July 1924, Small, Small & De Winkeler 11470 (NY).
Escambia: Elliott, 2 Sept. 1901, Biltmore 5895b (NY). Highlands: E. of
Sebring, 30 Apr. 1924, Small, De Winkeler & Mosier 11355 (NY). Jackson:
1869, Chapman s.n. (MO). Lake: Eustis, June-July 1894, Hitchcock IO7I
(MO, NY); Eustis, I6-3O June 1894, Nash 1070 (F, MO, NSC, NY); Tavares,
19 Aug. 1901, Biltmore 3848b (NY). Lee: Ft. Myers, July-Aug. I9OO,
Hitchcock 319 (MO). Leon: 6 miles S.E. Tallahassee, 22 Sept. 1957,
Godfrey 56O88 (FSU, GH, NSC, NY). Levy: S.W. of Chiefland, 30 Oct. 1964,
Godfrey, Clewell & Blaisdell 64780 (FLAS). Liberty: Apalachicola National
Forest, 8 Sept. I96O, Godfrey 60234 (FSU). Madison: 9 miles E. Green-
ville, 2 Oct. 1955, Godfrey, Krai & Gillespie 54012 (DUKE, FSU, NY).
Marion: Eureka, 16 Sept. 1941, West & Arnold s.n. (FLAS). Okaloosa:
between Valparaiso £■ Ft. Walton, 4 Sept. 1958, Godfrey 57662a (FSU).
Orange: 1894, Lewton s.n. (NY). Polk: Haines City, 16 Aug. 1897, Curtiss
5959 (F, FLAS, MO, NCU , NY). Putnam: Lake Swan, 5 Dec. 1964, Bratley
s.n. (FLAS). Suwannee: Ellaville, 15 July I96I, Ramsey, Godfrey £■ M i tchell
267 (FSU). Walton: Mossy Head, De Funiak Springs, 23 Sept. 1956, Godfrey

127.

Map 23. Distribution of C. cordi fol ia

Map 2k. Distribution of _C, polygoni fol ia

128.

& Krai 551^1 (FSU, NSC, NY); Summer 1885, Curtiss s.n. (NY).

This temperate species appears to have no strong affinities with
any taxa of the Caribbean.

21. CHAMAESYCE POLYGON I FOL I A (L.) Small, Fl. SE. US., 708.
1903. Euphorbia polyqonifol ia L. , Sp. PI. ed. 1, 455. 1753. Type:
K[alm], no other data, (Herb. Linn, sheet 63O-I9 I). Ani sophyl 1 urn
polyqonifol i urn (L.) Haw., Syn. PI. Succ. , I6O, 1812.

Annual; prostrate, rarely ascending, forming sparse or occasionally
dense mats to 4 dm. diameter. Stem to 2 mm. diameter, internodes to 3
cm. long; sparingly branched throughout length, not rooting at nodes,
glabrous, green or light brown. Leaves membranous or slightly fleshy;
blades ob long- 1 i near to lanceolate, 6-10 x 3-5 mm., base inequilateral,
rounded or obtuse, margins entire, apex acute, often mucronate, midrib
prominent on lower surface, glabrous, green; petiole 1-3 nrn. long;
stipules distinct, rarely joined near tip of stem, entire, toothed or
deeply 2-3 parted. Cyathia solitary, terminal on branches of short
laterals; peduncle 0.5 - 2 mm. long, Involucre broadly obconical, 1 - 1.4
mm. diameter, lobes equalling or exceeding glands, entire or toothed,
glands obsolete and replaced by additional lobes or broadly transversely
elliptic, 0.05 - 3 mm. long, concave, stipitate, usually exappend i cu 1 ate,
fifth gland vestigial or absent, sinus shallow. Staminate flowers 5-10
per cyathium, androphores glabrous, ca. 1 mm. long at maturity. Gynophore
glabrous, 2-3 mm. long at maturity, exserted, nutant; calyx a triangular
pad of tissue; ovary glabrous; styles upright or spreading, 0,3 - 0.7 mni.
long, minutely joined at base, bifid for 1/3 - 1/2 length, somewhat
clavate. Capsule glabrous, truncate-ovoid, 2.8 - 3.2 mm. long, 2.5 - 3 mm.

129,

Plate 28. CHAMAESYCE POLYGON I FOL I A (L.) Small

C. polyqonifol ia (L.) Small, Amelia Island, Nassau County, Florida
6 July 1951, Hood kkO] (FLAS).

130,

wide just above base, three- lobed, angles obtuse or rounded. Seed
ovoid-cuneiform, to 2.k mm. long, ca. 1,5 mm, diameter below equator,
dorsal side rounded, angles apparent only near apex, white.

Beaches and sand dunes of the Atlantic coast of North America,
The collections from northeast Florida represent the southern limit
of its range (Map 24).

Specimens seen: FLORIDA, Curtiss 2^98 (F) ; East Florida, 1874,
Palmer 7^25 (MO). Duval: Little Talbot Island, 12 Oct. I963, Burch 200
(FLAS). Nassau: Amelia Island, 6 July 1951, Hood 4401 (FLAS); W. of
Fort Clinch, 12 Oct. 1963, Burch 194 (FLAS, FSU , GH, iJ) ; Fort Clinch
State Park, 12 Oct. I963, Burch 199 (FLAS, MO, NCU, NY).

Found growing with _C, ammann ioides (HBK.) Small in Florida, but
distinguished from it by the possession of much larger capsules and
seeds, which are also cuneiform rather than ovoid. Other differences,
in stipular and gland characters, are less reliable in separating the
species.

These two species are both able to grow when partially covered
by shifting sand, and are often collected with the root5tocl< and all but
the tips of the branches covered to a depth of an inch or more.

22, CHAMAESYCE AMMANN IOIDES (HBK.) Small, Fl. SE. US., 709.
1903. Euphorbia ammannioides HBK., Nov. Gen. et Sp., 2: 55. 1817.
Type: Cumana, Venezuela, Bonpland 406 (Herb. Mus. Bot. , Berlin seen by
Boissier, and ? Herb. Mus, Paris, not seen; fragment F and photograph G
seen by Wheel er) .

_C. inqal Isi i Small, Fl, SE. US,, 708. 1903. Type: New Orleans,
Louisiana, 1835, Ingalls (NY and photograph G seen by Wheeler).

131

_E. inqal Isi i (Small) Cory, Rhodora 38: 406. I936.

E^. pol iosperma Urban in Feddes, Rep. 28: 234. 1930. Type:
Guanabo, Prov. of Habana, Cuba, 14 Oct. 1921, E. L. Ekman 13336 (Holo-
type: Mus. Bot. Stockh., not seen; Isotype: NY !).

Annual, perhaps perennating in the south; prostrate or decumbent
forming loose mats to 6 dm. diameter. Stems 1-2 mm. thick, internodes
to 4 cm. long; branching throughout length, not rooting at nodes,
glabrous, green, somewhat glaucous and often suffused red. Leaves
membranous or slightly fleshy; blades oblong-elliptic, 6-12 x 2.5 - 6 mm.,
base inequilateral, obtuse, margins entire, apex obtuse, often mucronate,
both surfaces glabrous, green, somewhat glaucous and often suffused with
red; petioles 1-2 mm. long; stipules distinct, deeply laciniate into 3-6
linear segments. Cyathia solitary, terminal on branches of lateral shoots
near tips of main stems; peduncle 1-3 mm. long, involucre campanulate,
1.2 - 1.5 mm. diameter near mouth, lobes triangular, slightly exceeding
glands, glabrous outside, hairy within, glands broadly elliptic or sub-
circular, 0.3 - 0.6 mm. long, concave, somewhat stipitate, appendages
obsolete, rarely to twice as wide as gland, fifth gland linear, often
equalling lobes, sinus shallow. Staminate flowers 8- 1 6 per cyathium,
androphores glabrous, 1 mm, long at maturity. Gynophore glabrous, to
4 mm. long at maturity, exserted, usually strongly nutant; calyx a tri-
angular pad of tissue; ovary glabrous; styles upright, 0.2 - 0.3 mm. long,
joined at base, bifid for 1/2 length, somewhat clavate. Capsule glabrous,
broadly ovoid, wider than long, 1.8 - 2.1 x 2 - 2.4 mm., three-lobed,
angles rounded. Seed ovoid, to 1.8 mm. long, 1.3 mm. diameter at equator,
three-angled from apex, sometimes for only 1/3 length, ventral angle

132.

Plate 29, CHAMAESYCE AMMANNIOIDES (HBK.) Small

C. ammannioides (HBK.) Small, New Point Comfort, Charlotte County,
Florida, 5 June 196^, Burch 3^9 (FLAS).

133.

rounded, obscure, faces convex, p 1 uinp , smooth, white.

Coastal sands, usually at or above the storm line, as far north
as southern Virginia and west to Texas in the United States. Found in
all coastal counties of Florida, in Cuba, and in northern South America,
but not otherwise known from the Caribbean (Kap 25).

Representative specimens seen: FLORIDA. Bay: Mexico Beach, 15
Oct. 1955, Godfrey S-Kral 5^205 (DUKE, FSU , NY). Brevard: Melbourne
Beach, 16 July 19^0, Crevasse s.n. (FLAS). Charlotte: New Point Comfort,
5 June 1964, Burch 3^9 (FLAS). Collier: Vanderbilt Beach, 13 Aug, I963,
Burch 183 (FLAS); Naples Beach, 20 Nov. 1964, Godfrey 65440 (FLAS, FSU).
Dade: Miami, 7 Mar. I9I5, Small & Small 5869 (DUKE, FLAS, FSU, NY).
Duval: Little Talbot Island, 12 Oct. I963, Burch 202 (FLAS); Pablo Beach,
25 July 1896, Lighthipe 427 (NY), Escambia: Pensacola, 24 Oct. 1964,
Burch 479 (FLAS). Flagler: Flagler Beach, 10 Oct. 1940, West & Arnold
s.n, (FLAS). Franklin: Alligator Harbor, 9 July 1951, Jackson 567 (FSU);
Carabelle Beach, 20 Oct. I963, Burch 335 (FLAS). Gulf: Port St, Joe, 20
Oct. 1963, Burch 330 (FLAS), Hillsborough: St. Petersburg, 5 Aug. 1894,
Lewton s.n. (NY), Indian River: Vero Beach, 7 Aug. I963, Burch 113 (FLAS),
Lee: Sanlbel Island, 29 Dec. 1953, Cooley 2489 (FLAS, NY); Ft. Myers
Beach, 8 Aug. 1945, Brass 15481 (US), Levy: Seahorse Key, 1 Dec, I962,
Ward 3255 (FLAS). Manatee: Palmetto, 21-23 Aug. 1895, Nash 2448 (MO, US);
Long Beach, 1 Aug. 1950, Wilbur & Webster 2511 (FSU); Bradenton, 28 Nov.
1901, Tracy 7427 (F). Martin: Jupiter Island, 7 Aug, I963, Burch 127
(FLAS). Monroe: Bahia Honda, 13-14 Feb. 1945, Killip 3l406 (NY), Nassau:
Fernandina, 21 Aug. 1902, Biltmore 3838 (US). Okaloosa: Destin, 24 Oct,
1964, Burch 465 (FLAS), Pinellas: Long Key, 24-28 Dec, 1951, Thorne 10220

13^.

(FSU); Honeymoon Island, 31 May 1S&^, Taylor 5- Taylor s.n. (FLAS).
St. Johns: k miles S. or Pome Vecra, 11 Oct. 1964, 3urch 441 (FLAS);
.'■■.arineland, 2 July 1939, Murrill s.n. (FLAS). St. Locie: Ft. Pierce,
7 Aug. 1963, Surch 121 (FLAS). Santa Rosa: Navarre, 24 Oct. 1964,
Surch 478 (FLAS). Sarasota: Venice 3each, 5 June 1964, Burch 352 (FLAS).
Volusia: Mosquito Inlet, 28 .Nov. 1929, Koldenke 5301 (NY); Daytona Beach,
1 Sept. 1927, West s.n. (FLAS). Walton: 4 miles E. of Oyster La^e, 19
Oct. 1963, Burch 216 (FLAS). CUBA. Habana: Playa de Goanabo, 21 June
1924, Leon II65I (NY); Jibacoa beach, 2 Jan. 1929, Leon 5- Roig 13761 (NY).

Small's action in setting up the new species C^. inqal 1 s i I is hard
to understand, particularly since his early annotations shov; that he
v/as familiar with _C. a: — 5-- " ' " ces as used by most authors, and since
he himself included this species in his Flora of the Southeastern United
States in 1903. Nothing in his later publications indicates v;hy he must
have cec\ced that _C. a~~.ar,r ioides v/as confined to the Caribbean.

Boissier in DC. Prod. 15(2); 28. 1862. includes Euphorbia
'zc~zer.s '. s Jacq., Enum. PI. Carib., 22. I76O. in his synonymy for this
species v/ith the comment "ex descr.". He evidently hesitated to accept
the name without seeing the specimens from which Jacquin v/or^ed, and,
v/hile there is no feature in the description which conflicts with
characters ooserved from specimens assigned by recent workers to _C.
5~~a"~. " z ' ces , some firmer basis is needed oefore this well-understood
name should be displaced. See the commentary ^-cer _C. buxi fol ia ror notes
on another name proposed by Jacquin in the sam.e publication,

23. CrJ\;-'AESYCE CUKULICOLA Sm.all, Kan. SE. Flora, 794. 1933.
Type: Caxambas Island, Florida, 11 Kay 1922, J. K. Sm.all 10490 (Holotype:

135.

Map 25. Distribution of _C. ammannioides

J-s:^-^..

Map 26. Distribution of C. cumul icola

136.

NY :; Isotype: FLAS I, US :).

Annual or perennating; prostrate with several stems from crown
of root forming diffuse mat to 20 cm. diameter. Stem 1-2 mm, thick,
internodes to 3 cm. long; branching throughout length, somewhat flexuous,
not rooting at nodes, glabrous, green or straw-colored. Leaves membranous;
blades long elliptic to lanceolate, 4-8 x 1-2 mm., rather uniform in
size on any plant, base oblique, obtuse or rounded, margin entire, apex
obtuse to acute, somewhat mucronate, midrib prominent on underside,
glabrous, green or red or with a broad green stripe down the center of
an otherwise red leaf; petiole 0.5 - 1 mm. long; stipules joined, 0.5 - 1
mm. long, 3-7 parted into linear segments. Cyathia solitary at upper
nodes; peduncles ca. 1 mm. long, involucre campanulate, 1-1.2 mm.
diameter at mouth, lobes triangular, exceeding glands, glabrous on out-
side, pilose within, glands subcircular, to 0.1 mm. diameter, somewhat
stipitate, appendages obsolete, fifth gland linear, about as long as
lobes, sinus shallow. Staminate flowers 5-8 per cyathium, androphores
glabrous, to 1 mm. long at maturity. Gynophore glabrous, 1.5 - 2 mm.
long at maturity, exserted, nutant; calyx a triangular pad; ovary glab-
rous; styles spreading, 0.2 - 0.3 mm. long, joined at base, bifid for
1/3 length. Capsule glabrous, ovoid or subspher ica 1 , 1.8 - 2 mm. long,
2 - 2.3 mm. wide close to equator, three-lobed, angles acute. Seed
ovoid, to 1.3 mm. long, 1 mm. wide below equator, obscurely angled at
apex, angles rounded below, faces convex, plump, white or gray-brown.

Known only from Florida, usually in sandy areas a little away
from the beaches or far inland (Map 26).

Representative specimens seen: FLORIDA. Collier: Caximbas inlet.

137.

THE NEW YORK BOTANICAL GARDEN

LXPLuRATION (Jl- I'ENINsLl.AH I-I.(J|<I|JA

Plate 30. CHAMAESYCE CUMULICOLA Small

Holotype: C. cumu 1 i cola Small, Caxambas Island, Florida, 11 May 1922,
Small IOU9O (NY).

138.

July, Curtlss IkSS (F, FLAS, MO, NY, UCWI , US). Escambia: Santa Rosa
Island, 10 Dec. 19^1, Blair 6 (FLAS). Highlands: north of Sebring, 16
Sept. 19^+8, Garrett s.n. (FLAS), Indian River: near Sebastian, 26 July
1924, Small, Small & DeWinkeler 115^1 (DUKE, MO, NY). Pinellas: sandy
area near Anclote River, 5 June 1964, Burch 36l (FLAS). St. Johns:
4 miles south of Ponte Vedra, 11 Oct. 1964, Burch 439 (FLAS).

The small seed and diffuse growth habit separate this species
from the closely related _C. ammann ioi des. The two are occasionally found
together, but _C. ammannioides is a shore plant, and _C. cumul icola more
often occurs in areas sheltered from continuous exposure to salt.

24. CHAMAESYCE PROSTRATA (Ait.) Small, Fl. SE. US., 713. 1903.
Euphorbia prostrata Ait.. Hort. Kew. 2: 139. 1789. Type: Nat[lve] of
the West Indies. Cult, 1758, by Mr. Philip Miller (British Museum if
extant). Anisophyllum prostratum Kl. & Gke., Abh. Acad. Berlin Phys. 1859:
26. 1860.

_E. cal 1 itrichoides HBK. , Nov. Gen. et Spec. 2: 52. 1817. Type:
Vera Cruz, Mexico (herb. Mus. Paris, not seen; photograph G seen by
Wheeler). Referred here by Boissier in DC, Prod. 15(2): 47. 1862.

_E. tenel la HBK., Nov. Gen. et Spec, 2: 53. 1817. Type: ad ripas
Orinoci prope Maypures et Carichana, Venezuela (Herb. Mus. Paris, not
seen; photograph G and fragment of isotype from Berlin at MO seen by
Wheeler). Referred here by Boissier, loc. cit., with note that type
specimen and description do not match in pubescence of capsule.

_C. malaca Small, Fl. SE. US., 713. 1903. Type: Kerrville, Kerr
County, Texas, 26-30 June 1894, A. A. Heller I9I8 (NY, photograph G,
isotypes F, G, MO, examined by Wheeler, 1941, who refers the name here).

139.

Annual; prostrate to 2 dm. Stem 0.5 mm. diameter, internodes to
1.5 cm. long or congested; several from base, branching throughout length,
sparsely short tomentose, sometimes only on upper surface, green, brown
or purplish. Leaves membranous; blades obovate or broadly elliptic,
k-8 by 3-5 mm., base rounded, margin serrate, apex rounded, sparsely
short tomentose, upper surface glabrate, green or suffused purple; petiole
to 1 mm. long; stipules separate or united at base, triangular to 0.5 mm.
long, toothed or somewhat lacerate. Cyathia solitary at nodes of somewhat
congested laterals; peduncle ca. 0.5 mm., involucre tubul ar-campanu late,
0,4 - 0.5 mm. at mouth, lobes exceeding glands, triangular, often ciliate,
strigose toward top on outside, glabrous within, glands transversely
elliptic, 0.1 - 0.3 rnm. long, somewhat cupped, rarely somewhat stipitate,
appendages obsolete or a rim to the gland, fifth gland linear, short,
sinus to a third of depth of cyathium. Staminate flowers rarely more than
6 per cyathium, androphores glabrous, 1 mm. long at maturity. Gynophore
sparsely pilose, to 1.5 mm. long at maturity, exserted, nutant; calyx a
three-Iobed pad of tissue; ovary strigose; styles 0.2 - 0.3 mm. long,
spreading, joined at base, bifid for more than 1/2 length; capsule ovoid
1.1 - 1.3 mm. long, 1.5 mm. wide above base, three-lobed, angles acute, with
stiff spreading white hairs. Seed cuneate-ovoid , 0.9 x 0.5 mm., strongly
four-angled, ventral angle more obtuse than others, faces flat or slightly
concave, transversely ridged, the narrow ridges sometimes passing through
the angles, ashen.

Waste places and disturbed ground in the southeast of the United
States and west to Arizona. Poorly collected but probably widespread in
Florida and the Bahamas, and of general occurrence in the remainder of the
Caribbean (Map 27).

\ko.

Plate 31. CHAMAESYCE PROSTRATA (Ait.) Smal

C. prostrate (Ait.) Small, Bahia Honda Key, Monroe County, Florida.
20 Nov. ]9Gk, Ward 4307 (FLAS).

141

Representative specimens seen: FLORIDA. Alachua: University of
Florida campus, Gainesville, 3 June 1S63, Burch 2017 (FLAS). Citrus:
Inverness, 15 Oct. 1964, Burch 444 (FLAS). Collier: Monroe Station,
19 Apr. 1964, Burch 315 (FLAS); Marco, 1900, Hitchcock 318 (F, MO, NY).
Dade: Homestead, 18 Apr. 1964, Burch 308 (FLAS). Dixie: Shamrock, 24
Nov. 1964, Godfrey 65683 (FLAS, FSU). Duval: Jacksonville, I88O,
Curtiss s.n. (MO). Jackson: Marianna, 19 Oct. I963, Burch 212 (FLAS).
Leon: Tallahassee, 5 Dec. 1964, Godfrey 65701 (FLAS, FSU) ; Tallahassee,
13 July 1926, Harper s.n. (NY). Monroe: Bahia Honda, 20 Nov. 1964, Ward,
Burch & Avery 4307 (FLAS); Garden Key, 1904, Lansing 2502 (F, NY).
Nassau: Fernandina, 29 Oct. I89O, Hitchcock s.n. (MO). Okaloosa: Wynne-
haven Beach, 24 Oct. 1964, Burch 474 (FLAS). St. Johns: St. Augustine,
13 Sept. 1898, Curtiss 6426 (GH, MO, NY). Volusia: 5 miles south of
Daytona Beach, 23 Aug. I962, Small, Small £■ deWi nkeler 10568 (NY).
BAHAMAS. New Providence: Nassau and vicinity, Mar. -May 1903, Curtiss
s.n, (F, NY). Grand Turk Island: Ponds and vicinity, 20 Feb. -24 Mar.
1911, Millsp. and Millsp. 8991 (F, NY). CUBA. Camaguey: Atalaya, 20
Mar. 1909, Shafer 999 (F, NY). Habana: Botanical Garden Havana, 22 Mar.
1905, Curtiss 690 (F, GH, MO, NY). Oriente: 1857, Wright 549 (F, GH,
MO, NY). Pinar del Rio: Bay of Mariel, 21 Sept. I9IO, Britton & Earle
7574 (F, NY). Santa Clara: Cienfuegos, 27 Oct. 1928, Jack 6567 (GH, MO).
CAYMAN ISLANDS. Grape Tree Point, 14 Apr. 1956, Proctor 11971 (U).
JAMAICA. Kingston: Race Course, 24 Oct. 1954, Proctor 9378 (IJ, PUR).
St. Ann: Moneague, 22 Jan. 1952, Hunnewell 19765 (GH). Clarendon:
Cockpit, 15 June 1959, Webster, Ellis £• Miller. Portland: Port Antonio,
25 Sept. 1906, Britton 846 (F, NY). St. Andrew: Hope Gardens, 29 Oct.

142.

1914, Harris II803 (GH, MO, NY, UCWl); Un i vers i ty of West Indies Campus,
9 June 1963, Crosby, Hespenheide £■ Anderson 82 (DUKE, F) . St. Catherine:
Bog Walk, 13 Jan. 1958, Yuncker 17950 (F, UCWl). St. Michael: Pine
Estate, 4 Aug. I9OI, Freeman I5I (UCWi). Trelawny: Good Hope, 29 Aug.
1956, Proctor 15771 (GH, IJ). HAITI. Navassa Island, 20-23 June I956,
Proctor 15458 (GH, IJ); La Tortue, La Vallee, 14 June 1925, Ekman H4316
(IJ); Port au Prince, 15 Oct. 1926, Ekman H9125 (A); St. Marc, 25-28 Feb.
1920, Leonard 2858 (GH, NY); St. Louis du Nord, 30 Mar. -7 Apr. 1929,
Leonard & Leonard 14262 (NY). DOMINICAN REPUBLIC. Ciudad Trujillo,
5 Dec. 1945, Allard I3892 (F, MO); Barahona, Sept. 1911, Fuertes 996
(F, GH, NY); Azua, 20 Mar. 1913, Rose, Fitch & Russell 4076 (NY). PUERTO
RICO. Near Soroco, N.E. of Fajardo, 3 July 1959, Webster, Miller & Ellis
8623 (PUR); Rio Piedras, 29 Apr. 1914, Stevenson s.n. (NY); Santurce,
Bayamon, 31 Oct. 1937, Otero 1 78 (MO) ; San Juan, 26 Jan. 1899, Heller
£■ Heller 326 (NY); Isabel Segunda, Vieques Island, 24-27 Jan. 1914,
Shafer 2419 (F, NY); Culubrita Island, 11 Mar. I906, Britton & Wheeler
277 (F, NY). LESSER ANTILLES. St. Thomas: Dec. I88O, Eggers 334 (GH).
St. Croix: New Fort, 11 Nov. 1895, Ricksecker 63 (F, GH, MO, NY); Coakley
Bay, 30 Sept. 1923, Thompson 332 (GH) . St. Barthelmey: Gustaria, 5 Jul.
1938, Q,uestel 434 (NY). St. Kitts: Wingfield Estate, 8 Sept. -5 Oct. I9OI,
Britton & Cowell 578 (NY). Antigua: Ft. James, 7 Oct. 1937, Box 1144
(F, MO, NY). Guadeloupe: Basse Terre, 1894, Duss 3420 (F, GH, MO, NY).
Gosier: Aug. I898, Duss 2739 (NY). Dominica: Roseau, Aug. 1937, Hodge
393 (GH). Martinique: Casepilote, Aug. I869, Hahn IIO8 (GH, NY); St.
Pierre, Aug. 1899, Duss 4056 (NY). St. Vincent: 17 Feb. 1922, Bailey
400 (NY). Barbados: St. John, July 1941, Gooding 276 (NY). Grenada:

143.

Map 27- Distribution of _C, p rost rata

t:

P-^^''

Map 28. Distribution of C_. mendezi i

]kk.

St. Georges, Broadway 1865 (MO). Extra territorial specimens: BERMUDA.
Flatts Inlet, Smiths Parish, 8 July 1905, Moore 2871 (GH) ; Devonshire,
13 Jan. 1912, Robinson 81 (GH).

Wheeler (19^0 proposed the name _E. chamaesyce for this species
on the basis of a sheet (630-17) in the Linnaean Herbarium, which is
annotated with this name and holds a specimen of the plant here called
_C. pros trata. He proposed this as type specimen for the name _E. chamae-
syce, but the sheet was one bought from Patrick Browne, and was not in
Linnaeus' possession until some years after his publication of the name.
The annotation by Linnaeus is a simple mi s ident if icat ion, and the name
_E. chamaesyce should be limited in its application to the Mediterranean
species, quite distinct from our _C. prostrata, for which he first used it.

Plants from Texas and Arizona differ from those of the West Indies
In being more heavily pubescent, with the hairs on the capsule less
rigidly confined to the angles. One specimen from west Florida, Burch
474, is of this form, but differs In no other respect from the usual
characters of the species in the Caribbean. These plants do not merit
recognition at even the varietal level.

25. CHAMAESYCE MENDEZll (Boiss.) Millsp., Field Mus. Bot. 2:
410. 1916. Euphorbia mendezi I Boiss., Cent. Euph. , 15. i860. Type:
ad Leon, prope Guanajuato, Mexico, Mendez s.n. (Herb Candolle, not seen).

E. dorsiventral is Urb. , Symb. Antlll., 5: 391. I908. Type: Madruga,
Cuba; 25 Nov. 1904, Curtiss 535, Batabano, Cuba; 3 Oct. 1904, Baker and
Wilson 2227; Prope Habana ad Vlvora, Torralbas 282 (Syntypes: Herb . Bot.
Mus. Berlin, if extant. Duplicate specimens of Curtiss 535: F '. , GH '. ,
K 1, NY i; Baker £■ Wilson 2227 NY 1, F i, K i). _C. dorsiventral Is (Urb)

145.

Millsp., Field Mus. Bot. 2: 409. 1916.

Perennial; prostrate to decumbent, forming mats to k dm. diameter.
Stems to 2 mm., internodes to 2 cm. long or congested; branching tinrough-
out length, laterals short and compact near tip of stem, not rooting at
nodes, upper surface and sides long-pilose, underside glabrate, green
or reddish. Leaves membranous; blades broadly oblong-elliptic, 4-8 x 2-5
mm., base inequilateral, rounded or cuneate, margin serrate, apex rounded
or truncate, glabrous or rarely with scattered long hairs, green, often
orange-red with age. Cyathia solitary, appearing clumped on congested
laterals; peduncle to 1.5 mm. long, involucre narrov;ly obconical, to 0.6
mm. diameter at mouth, lobes equalling glands, triangular, toothed or
deeply cut, glabrous on outside, long-pilose at throat within, glands
transversely elliptic, 0.2 x 0.1 mm., cupped purple, appendages as long
as glands and 2-3 times as wide, rarely obsolete, white or pink, fifth
gland linear, ciliate, short, sinus shallow. Staminate flowers 6-8 per
cyathium, androphores glabrous, to 1.5 mm. long at maturity. Gynophore
glabrous, to 2 mm. long at maturity, exserted, nutant; calyx vestigial;
ovary strigose; styles spreading, 0.35 mm. long, minutely joined at base,
bifid for 1/2 length, obscurely clavate. Capsule ovoid, truncate, 1.2
mm. long, 1.2 - 1.3 mm. wide below equator, three-lobed, angles acute to
obtuse, sparsely long-pilose at angles and rarely over sides of capsule.
Seed narrowly ovoid, to 1 mm. long, 0.5 mm. wide just below equator,
strongly four-angled, ventral angle more obtuse than others, faces some-
what concave, obscurely wrinkled, dark tan or chestnut-brown.

A plant of waste places and disturbed ground in Mexico and Cuba,
and a recent introduction to Florida (Map 28).

litb.

' ' •'<''»Miiiiii|iiii|i(ii|iiii|liii|iniiiiii|iMi|iiii|ini|iiii|iHi|iin|iiii|iin

1^

iflliil

iJSihiik: ^ai.'

Plate 32. CHAMAESYCE MENDEZII (Boiss.) Millsp.

Duplicate Coll. of Syntype: E. dors i ventral i s Urban, Madruga, Cuba,
25 Nov. 1904, Curtiss 535 (NY).

\kl,

Representative specimens seen: FLORIDA. Collier: Goodland,
26 Sept. 1964, Burch 401 (FLAS, GH, US); Naples, 2? Sept. 1964, Burch 412
(BM, F, FLAS). Dade: Redlands, 5 July 1959, Atwater M- 1 1 3 (FLAS); Old
Cutler Rd., Miami, 29 Nov. I963, Burch 219 (FLAS, NY, US); Homestead,
19 Apr. 1964, Godfrey 6336I (FLAS, FSU). Highlands: Lake Annie, 18 Nov.
1964, Burch & Ward 528 (FLAS, NY), Monroe: Flamingo, Everglades National
Park, 17 Apr. 1964, Burch & Ward 288 (FLAS); Plantation Key, 21 Nov. 1964,
Burch & Ward 563 (FLAS). Polk: Babson Park, 27 Nov. 1964, Burch 58O (FLAS)
St. Lucie: Nigger Jim Scrub, 26 Oct. 1957, Brass 29046 (FLAS). CUBA.
Habana: Chorrera near Havana, 22-23 Feb. I9II, Britton, Cowell S- Leon
9585 (F, NY); Rio Almendares to Playa de Marianao, 22-23 Dec. 1910,
Wilson 9457 (F, GH, NY); Estacion de Cristina, 7 Dec. 1905, Van Hernann
3386 (NY); El Morro to Cojimar, near sea, 8 Dec. I9IO, Wilson 9125 (F,
NY). Pinar del Rio: vicinity of Herradura, 4-5 Sept. 1910, Britton &
Gager 6940 (F, NY); Bay of Mariel, 19-23 Sept. I9IO, Britton £■ Gager 7675
(F, NY).

The first collection of _C, mendezi i in Florida was made in 1957
in St. Lucie County (Brass #29046), and it is now widespread in the
southern part of the state. The plant is a strong grower, reaching full
development in late summer, and, since it occupies the same general
habitats as C_. maculata and _C. prostrata, it will be interesting to
observe its spread in the face of competition from these two.

The species was named by Boissier from Mexican material, but Cuban
collections correspond closely with this, and Urban was not justified in
separating them as a distinct taxon.

1^8.

26. CHAMAESYCE HEPATICA (Urb. & Ekman). comb. nov. _E. hepatica
Urb. & Ekman, Ark. f. Bot. Stockh, 22A(17): 113. 1929. Type: Navassa
Island, Haiti, 17-25 Oct. 1928, E. L. Ekman HIO8O9 (Holotype: Herb.
Naturhistor iska Riksmuseum, Stockholm, not seen; Isotype: IJ i).

Perennial; prostrate, forming mats 10-15 cm. diameter. Stem
wiry, 0.25 - 1 mm. diameter, internodes to 1.5 cm. long; many grov/ing
from crown of taproot, branching near base, but simple in upper part
of stem, not rooting at nodes, sparsely tomentose, straw-color or darker
brown. Leaves membranous; blades oblong to obcuneate, 3.5 - 6 x 1.5 - 3
mm., midrib sometimes prominent on underside, depressed above, base
oblique, margin deeply and coarsely serrate, apex acute, sparsely tomen-
tose, purplish-green above, lighter below; petiole 0.5 - 1 mm.; stipules
joined on lower side of stem, triangular, 3-5 cleft, to 0.3 mm. long,
ciliate, usually distinct on upper surface, subulate, to 0,1 mm. long.
Cyathia solitary at upper nodes; peduncle short, broadening to tubular
involucre, 0.3 mm. diameter at mouth, lobes truncate, scarcely equalling
glands, sparsely tomentose outside, pilose within, glands transversely
elliptic, 0.1 mm. long, deep red, appendages obsolete, fifth gland tri-
angular equalling lobes, sinus shallow. Staminate flowers 3-5 per
cyathium, androphores glabrous, 0.7 mm. long at maturity. Gynophore
strigose, 0.5 mm. long at maturity, exserted, nutant; calyx minute or
obsolete; ovary densely strigose; styles upright or spreading, minutely
joined at base, 0.4 mm. long, bifid for 1/3 length. Capsule strigose,
ovoid, 1.2 mm. long, to I mm. wide below equator, three-lobed, angles
obtuse. Seed ovate, 0.8 mm. long, 0,k mm. wide below equator, strongly
four-angled, the ventral angle somewhat rounded, others very acute, faces

1^9.

Hri|fHiiitii|iiiipi|iimiiii|iiiiiiiiimrmmiiiii|iiii|nii|iiii|^^
JdililiLlililjMllilJiljJ.UklulaJ

ISOTYPE or

N. B 10809

Plate 33. CHAMAESYCE HEPATICA (Urb. & Ekm.) ined.

Isotype: E. hepat i ca Urb. & Ekm,, Navassa Island, Haiti, 17-25 Oct. ]S2i
Ekman H10809 (U).

150.

concave, obscurely wrinkled, dark red-brov/n overlaid with gray.

Known only from Navassa Island, Haiti, where it forms small mats
in gravelly areas between limestone or in cracks in the rocks.

Specimens seen: HAITI, Near lighthouse, Navassa Island, 20-23
June 1956, Proctor 15^57 (BM, GH, IJ).

The affinities of this species are with the group of wiry-stemmed
species of Cuba and Hispaniola, but it is easily distinguished from them
by its tiny holly-like leaves.

27. CHAMAESYCE HELWIGII (Urb. & Ekman) comb. nov. Euphorbia hel-
wigii Urban £■ Ekman, Arkiv. Bot. Stockh. 22A(8): 65. 1929. Type: Road
to Bassin, Les Gonaives, Presqu'ile du Nord'ouest, Haiti, 24 Dec. 1927,
E. L. Ekman H9546 (Holotype: Mus. Bot. Stockh., not seen; isotypes:
NY i, IJ :).

Annual; prostrate or decumbent to 1 dm. Stem to 1 mm. thick,
internodes to 1 cm. long; sparsely branched, mostly from upper nodes,
not rooting at nodes, pilose, straw-colored, becoming darker. Leaves
membranous, or somewhat thickened; blades elliptical or oblong, 2-5 x 2-3
mm., base inequilateral, cuneate or rounded, margin coarsely serrate,
somewhat revolute, sparsely long pilose, midrib prominent on underside,
green suffused red; petiole to 0.5 mm. long; stipules joined only at
base, deeply cleft, to 0.3 mm. long, light colored. Cyathia solitary at
upper nodes; peduncle to 0.3 mm. long, or broadening to campanulate Involucr
to 0.7 mm. diameter at mouth, lobes equalling glands, triangular, entire
or two to three parted, ciliate, glabrous or pilose near apex on outside,
pilose within, glands subcircular, 0.1 mm. diameter, cupped, appendages
slightly longer than and to twice as wide as glands, fifth gland deltoid.

""|irM|iili|llll|ini|ll||||||l||||||||||||||||||||||

■ «^ * '» '« '7

i|iinjiiii|Miijiiii|iMijiiiijiiiijiiiijiiiiHiri|iiii|,iii|ii,|j|in

Jihl^ilihlihlililJZlJiEidl^^

PtMtai Hispiivolje Eip. ill Regnellan..

8tp. Hicli /^^

/'

^^■

Plate 3^. CHAMAESYCE HELWIGII (Urb. & Ekm.) ined,

Isotype: _E. helwigi i Urb. & Ekm., Bassin, Haiti, 21 Dec. 1927,
Ekman H9^56 (IJ) .

152.

short, sinus shallow. Staminate flowers 3-8 per cyathlum, androphore
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
strlgose; styles spreading, to 0,3 mm. long, minutely joined at base,
bifid for 1/3 length. Capsule sparsely pilose, broadly ovoid, 1.5 - 1.8
mm. long, 1.8-2 mm. wide just below equator, roundly three-lobed,
angles obtuse or rounded. Seed oblong-ovoid, base truncate, to 1.3 mm.
long, dorsal face 0.8 mm. wide, strongly four-angled, dorsal angle acute,
ventral angle somewhat rounded, faces flat or slightly convex, smooth,
white or gray-brown.

Known only from type collection (Map 33).

This species is easily distinguished f rom _C. turp i ni i , which it
otherwise resembles closely, by the relatively large seed, with broad,
smooth, light-colored dorsal faces.

28. CHAMAESYCE HIRTA (L.) Mi lisp.. Field Mus. Bot. 2: 303. 1909.
Euphorbia hi rta L. , Sp. PI. ed. 1: 454. 1753. Type: [Euphorbia] hirta 18
(Linn, sheet 630-7 I). E. capitata Lam., Encyc. , 2: 422. 1786. substituted
on the ground that _E. hi rta was a bad name. _E. hi rta L. var. typ ica L.C,
Wheeler, Contr. Gray Herb. 127: 68. 1939.

E.. qlobul ifera HBK. , Nov. Gen. et. Sp., 2: 56. 1817. Type:
Cumana, Venezuela, Bonpland 403 (Herb. Mus. Paris, not seen; fragment
F 1, fragment F and photograph of type GH seen by Wheeler, Contr. Gray
Herb. 127: 68. 1939).

E_. p 1 1 ul ifera L. sensu Jacquin, I cones PI. Rar. 3: t. 478. I986-
93; Boiss. in DC. Prod. 15(2): 21. 1862, and many other authors.

Annual; decumbent to 3 dm. Stems few, to 2 mm. diameter, internodes

153.

to 3 cm. long; sparingly branched, tomentose and with abundant multi-
cellular hairs which are often brown or purple, rarely merely tomentose,
brown or red. Leaves membranous; blades ovate to lanceolate, often
rhombiform, 5-35 x 3-15 mm., base rounded to cuneate, inequilateral,
margin serrate, apex obtuse to acute, sometimes attenuate, strigose,
glabrate on upper surface, green or red often with purple spot; petioles
1-2 mm, long; stipules distinct or joined at base, somewhat lacerate, to
1 mm. Cyathia in dense terminal and axillary glomerules formed of
condensed leafless dichasia; peduncles short, involucre conical or tubu-
lar, to 0.3 mm. at mouth, lobes exceeding glands, attenuate, lacerate,
strigose on outside, subglabrous within, glands minute, purple, stipitate,
cupped, appendages obsolete to three times as wide as gland, fifth gland
linear, shorter than lobes, sinus shallow. Staminate flowers 3-8 per
cyathium, androphores glabrous, to 1 mm. long at maturity. Gynophore
glabrous, to 1 mm. long at maturity, exserted, upright or nutant; calyx
a triangular pad of tissue or obsolete; ovary strigose; styles spreading,
0.2 mm. long, joined at base, bifid 1/2 to 2/3 length. Capsule strigose,
ovate, to 1 mm. long, 1 mm. wide just above base, roundly three-lobed,
angles obtuse. Seed cuneiform, 0.8 mm. long, 0.4 mm. wide above base,
strongly four-angled, angles subequal, faces concave, obscurely trans-
versely ridged or wrinkled, tan, red-brown or gray.

A common weed of cultivated or disturbed ground in Florida and
throughout the West Indies and much of Central and South America (Map 29).

Representative specimens seen: FLORIDA. Alachua: University of
Florida campus, Gainesville, 9 June 1936, Murrill s.n. (MO); University
of Florida campus, Gainesville, 27 Apr. 1965, Wiggins £■ Wiggins 19833

15it.

plate 35. CHAMAESYCE H I RTA (L.) Millsp.

C. hi rta (L.) Millsp., Homestead, Dade County, Florida, 16 April 196^,
"Burch 269 (FLAS).

155.

(FLAS). Brevard: Okeechobee region, 9 Sept. I9O3, Fredholm 599^ (NY) .
Broward: Ft. Lauderdale, 12 Feb. 19^6, Beck H-69 (FLAS). Charlotte:
Charlotte Harbor, 21 Nov. 1964, Godfrey 65359 (FSU); Punta Gorda, 8 July
1948, Garrett s.n. (FLAS). Citrus: Crystal River, I9 Nov, 1964, Godfrey
65081 (FLAS, FSU); Pineola, 3 Dec. 1955, Krai I89I (FSU). Clay: Gold
Head Branch State Park, 1 Nov. 1964, Warren s.n. (FLAS). Collier: 7 mi.
N. of Naples, 29 Dec. 1948, Deam 65587 (FSU); 2.5 miles S.E. of Naples,
22 Nov. 1964, Godfrey 65467 (FLAS, FSU). Dade: Buena Vista, 25 Dec, 1929,
Moldenke 283 (DUKE, MO, NY); between Coconut Grove & Cutler, 31 Oct. -4 Nov.
1903, Small £• Carter 1283 (F, NY); Deering Reservation, Buena Vista,
3 May 1918, Small 8571 (NY); Homestead, 23 June I96O, Webster & Samuel
10093 (DUKE, PUR). De Soto: 7 mi. W. of Arcadia, 29 Nov. I962, Ward &
Burch 3222 (FLAS). Dixie: Shamrock, 24 Nov. 1964, Godfrey 6568I (FSU).
Glades: West side of Lake Okeechobee, 29 Nov. 1958, Ward & Ward 1145
(FLAS). Gulf: Port St. Joe, 20 Oct. I963, Burch 333 (FLAS). Hardee:
Wauchula, 26 Sept. 1964, Burch 390 (FLAS). Hendry: Clewiston, 2 Jan.
1964, Burch 240 (FLAS). Hernando: 4.5 mi. N. of Brooksville, 12 July
1958, Godfrey 57200 (FSU); June-July I898, Hitchcock 1 706 (F). Highlands:
Sebring, 30 July 1934, McFarlin 8l4l (DUKE, FLAS). Hillsborough: Tampa,
13 Aug. 1964, Burch I9I (FLAS). Indian River: Vero Beach, 7 Aug. I963,
Burch 114 (FLAS). Lake: Eustis, June-July 1894, Hitchcock 1705 (F);
Eustis, 16-30 June 1894, Nash 997 (NSC, NY), Lee: Ft. Myers, July-Aug.
1900, Hitchcock 326 (F, NY); Sanibel Island, 30 Apr. 1954, Cooley 2645
(FLAS, NY). Manatee: Bradenton, 13 June I9I6, Cuthbert 1313 (FLAS, NY);
Bradenton, 9 June l840, Simpson s.n. (F). Marion: Silver Springs, 1 Oct.
1963, Godfrey 63144 (FSU). Martin: Jensen, 22 Oct. I898, Rolfs s.n. (FLAS);

156.

Sruart, 5 Aug. I962, Krai 15375 (FSU). Monroe: Everglades National
Park, 15 Aug. I96O, Mitchell £■ Mitchell 66O (FSU); Key West, Stock
Island, 12 Oct. 1950, Whitehouse 24020 (FSU, NSC); Upper Matecumbe Key,
Curtiss 2^96 (F, NY). Okeechobee: Okeechobee, 27 Dec. 1964, Burch 590
(FLAS). Orange: Orlando, 1 Aug. 1947, Schallert £■ Schallert 20768 (FLAS).
Osceola: County line, FLA 60, 7 Aug. I963, Burch 107 (FLAS). Palm Beach:
Palm Beach, 18 May 1895, Curtiss 5395 (FLAS, NY); Palm Beach, 26 Dec.-
11 Jan. 1896, Hitchcock 1 708 (F). Pinellas: St. Petersburg, 23 Sept.
1907, Deam 2762 (F). Polk: Lakeland, 1 Aug. 1948, Garrett S- Garrett
(FLAS). St. Lucie: Ft. Pierce, 7 Aug. I963, Burch 123 (FLAS). Sarasota:
2 miles S. of Venice, 30 Dec. 1956, Krai 3834 (FSU). Volusia: Crows
Bluff, 29 June 1941, Hindery & Hindery s.n. (FLAS). Washington: Chipley,
25 Oct. 1964, Burch 501 (FLAS). BAHAMAS. Bimini: North Bimini, May 1948,
Howard S- Howard 10215 (NY). New Providence: Blue Hills, 14 July I96O,
Webster, Samuel S- Williams 10484 (DUKE, IJ); Farrington Road, 2 June
1909, Wilson 8333 (F, NY); Nassau, Nov. I89O, Hitchcock s.n. (MO). Andros:
Mastic Point, 19-28 Mar. 1907, Brace 7035 (F, NY). Eleuthera: Gov. Harbor
& vicinity, 19-20 Feb. 1907, Britton £■ Millspaugh 5471 (F, NY); Gov.
Harbor, 14 Nov. I89O, Hitchcock s.n. (MO). CUBA. isle of Pines: Caleta
Cocodrilos, 8 Mar. I9I6, Britton, Wilson & Leon 15252 (F, NY). Camaguey:
La Gloria, 1 Feb. 1909, Shafer I83 (NY). Oriente: 1856-1857, Wright 550
(NY). Matanzas: Valley of San Juan, 21 Mar. I9I6, Britton, Britton £•
Shafer 534 (NY). CAYMAN ISLANDS. Grand Cayman, Spot Bay, 13014 Feb.
1899, Millspaugh 1278 (F). JAMAICA. St. Andrew: U.of W.i. campus, 4 June
1963, Crosby, Hespenheide & Anderson 61 (DUKE, PUR); Constant Spring to
Bardowie, 2 Aug. 1915, Harris 12111 (MO, UCWl). St. Michael: Waterford,

157.

10 Aug. 1906, Dash I67 (UCWl). Clarendon: Kupuis, 23 June 1955, Proctor
10329 (IJ). St. Elizabeth: south of Gutters, I6 Sept. 195^, Howard &
Proctor 13886 (IJ). HAITI. Port-de-Pa ix, 17 March 1928, Ekman H9716
(PUR); vicinity of St. Marc, 25-28 Feb. 1920, Leonard 2867 (NY);
vicinity of Gros Morne, 1 ' Art i bon i te, 17 Feb. 1926, Leonard 9797 (NY);
vicinity of La Vallee, Tortue Island, 28 Dec. -9 Jan. 1929, Leonard &
Leonard 11489 (NY). DOMINICAN REPUBLIC. Barahona: Barahona, July
1910, Fuertes 127 (F, NY). San Juan: El Cercado, Hondo Valle, 2 Sept.
19it6, Howard & Howard 8737 (NY). PUERTO RICO. Soroco, Fajardo, 3 July
1959, Webster, Miller £- Ellis 8620 (DUKE, IJ, PUR); Cabra Island, San
Juan, Bayamon, 12 Oct. 1937, Otero 99 (MO); San Turce, 16 Jan. 1903,
Heller 6405 (F, NY); Vieques Island, 24-27 Jan. 1914, Shafer 2513 (F,
NY); Culebra Island, 3-12 Mar. I9O6, Britton £■ Wheeler 173 (NY); Catano,
6-11 Jan. 1899, Millspaugh 1775 (F) ; Toa Baja, 6 July 1912, Johnston
460 (NY). LESSER ANTILLES. Tortola, 13 Oct. I9I8, Fishlock 235 (NY);
St. Thomas, Charlotte Amalie, 17-18 Jan. 1899, Millspaugh 426 (F); St.
Croix, 18 Nov. 1895, Ricksecker 92 (NY); St. Barthelemy: Gustaria, 4 Dec.
1937, Questel 122 (NY); St. Jean, Nov. 1938, Questel 621 (NY). Antigua:
16-17 Jan. 1907, Shafer 21 (NY); Montserrat, 19 Jan. 1907, Shafer 15 (NY);
Guadaloupe, I833, Duss 2453 (NY); Dominica, 1903, Lloyd 229 (NY); Martin-
ique, 29 July 1939, Egler 39-209 (NY); St. Lucia, 1 Dec. I887, Lee s.n.
(F); Carriacou, 7-25 Mar. 1950, Howard 10848 (IJ, NY); Grenada: Tufton
Hall Estate, 30 Oct. -11 Dec. 1957, Proctor 17147 (U). Extra- terri torial
specimen: GEORGIA. Sapelo Island, Mcintosh County, I6 July 1956, Duncan
20301 (NCU, NSC).

Linnaeus described E. hi rta and E_. pi lul ifera in Species Plantarum

158.

edition 1 (1753) under the numbers 18 and 19, referring to an earlier
description by WIman (1752) and to illustrations by Burman (1737).
There are specimens in the Linnaean Herbarium labelled by Linnaeus
(sheet 630-5, 630-6 'hirta', 63O-7 '18 hirta', 63O-8 '19 pi 1 ul i fera' ) ,
and Brown (1911) has pointed out the close correspondence between these
and the descriptions. Burman's illustrations, on the other hand, appear
to be merely two forms of the same species, so that if Linnaeus intended
to describe two distinct species he was In error In citing Burman as
an illustration of both types.

There is no doubt that the specimens In the herbarium represent
two entities. The general appearance of the sheets Is similar but sheet
630-8 ' 19 pilullfera' lacks the multicellular hairs that are so charac-
teristic of _E. hi rta and which occur on all the other specimens. The
plant on this sheet Is glabrous in all parts except the capsules, and
has leaves toothed only in the upper third, and more rounded at the apex
than those of the other sheets.

Some workers have regarded the two species as conspeclfic, arguing
that Burman's illustrations were the source of Linnaeus' Information.
Farwell (1936) has shown that. If one name is to be reduced to synonymy,
this taxon must be called _E. p I 1 ul i fera. If, however, the herbarium sheets
are taken as representative of the concept of these two species which
Linnaeus held, as might well be the case in viev; of their close correspon-
dence with the descriptive phrases, then E_. hi rta Is clearly the name of
the species under consideration. This second argument is accepted here,
and the epithet hi rta will be used for our species.

This species is uniform through the area in most features, but

159.

Map 29, Distribution of _C. hi rta

Map 30. Distribution of _C. opthalmica

160.

occasional plants are found which lack the heavy orange-yellow hairs
found in other specimens. These plants are strikingly different in
the field, but correspond closely on all characters except this one,
and probably do not merit recognition at even the varietal level. Speci-
mens of this form have been seen from Florida, Jamaica, Puerto Rico and
Guatema la.

29. CHAMAESYCE OPTHALMICA (Pers.)comb. nov.E. opthalmica Persoon,
Syn. PI., 2: 13. l807. Type: Rio de Janeiro, Brazil, July I767,
Commerson 238 (Herb. Mus. Paris, not seen; fragment F seen by Wheeler).
Referred to this entity by Boiss. in DC. Prod. 15(2): 21. 1862, and
identity confirmed by Wheeler, Contr. Gray Herb. 127: 69. 1939 with
comment "[fragment Field Mus. 1] A small-leaved plant.".

_E, procumbens DC, Cat. PI. Hort. Monsp., 111. I8I3. Type: probably
a plant from garden at Montpellier, France (Geneva If extant). _E. p i 1 u1 i fera
L. var. procumbens (DC.) Boiss., DC. Prod. 15(2): 21. 1862. _C. pi lul ifera
(L.) Small var. procumbens (DC.) Small, Fl. SE. US., 71^. 1903. E. hirta
L. var. procumbens (DC.) N.E. Brown in Thi sel ton-Dyer , Fl. Trop. Afr.
6(1): 497. 1911. _C. hi rta L. var. procumbens (DC.) Moldenke, Rev. Sudam.
Bot. 6: 178. 19^0. C_. hi rta L. ssp. procumbens (DC.) Croiz. in Moldenke,
Li lloa 6: 299. 19^1.

E_. qemel la Lag., Gen. et Spec. Nov., 17. I8I6. Referred here by
Boiss. in DC. Prod. 15(2): 21. 1862. _C. qemel la (Lag.) Small, Fl. Miami,

110. 1913.

Annual; prostrate or decumbent to 15 cm. Stem to 1 mm. at base,
internodes to 3 cm. long; branched throughout, not rooting at nodes,
densely tomentose, subglabrate, long multicellular hairs throughout,

161.

particularly at tips, hairs often yellow or brown, stems green or red.
Leaves membranous; blades elliptic, often markedly rhombic, 7-15 x 3-7
mm., base inequilateral, rounded to cuneate, margin sharply serrate, apex
attenuate to acute, strigose, glabrate above, green or red; petioles to
1 mm.; stipules distinct or united at base, attenuate to 1 mm., short
cleft or ciliate. Cyathia glomerate, in terminal leafless dichasia;
peduncles short, involucre conical or tubular to 0.3 mm. at mouth, lobes
exceeding glands, attenuate, lacerate, strigose on outside, subglabrous
within, glands minute, purple, stipitate, cupped, exappend i cu late , fifth
gland linear, shorter than lobes, sinus shallow. Staminate flowers 3-8
per cyathium, androphores glabrous, to 1 mm. long at maturity. Gynophore
glabrous, to 1 mm. long at maturity, exserted, upright or nutant; calyx
a triangular pad of tissue or obsolete; ovary strigose; styles spreading,
0.2 mm. long, joined at base, bifid 1/2 to 2/3 length. Capsule ovate,
strigose, to 1 mm. long, 1 mm. wide just above base, roundly three-lobed,
angles obtuse. Seed cuneiform, 0.8 mm. long, 0.4 mm, wide above base,
strongly four-angled, angles subequal, faces concave, obscurely trans-
versely ridged or wrinkled, tan, red-brown or gray.

A common weed of cultivated ground and disturbed areas, Bermuda,
southern and central Florida, Bahamas, and probably through the Antilles
although no specimens have been seen from Haiti or the southern part of
the Antillean chain (Map 30).

Representative specimens seen: FLORIDA. Alachua: Gainesville,
30 June 1964, Burch 382 (FLAS). Brevard: Sebastian inlet, 17 Apr, 1965,
Burch 636 (FLAS). Charlotte: Charlotte Harbor, 21 Nov. 1964, Godfrey
65358 (FLAS, FSU). Collier: Monroe Station, 19 Apr. 1964, Burch 3I6

162.

911 04

^ ii^

Aigrlcultuiftl trpaivttwtt %taJ>-*

PlanU of Fiori<ti

QARaqay-

Plate 36. CHAMAE5YCE OPTHALMICA (Pers.) ined.

C. oDthalmica (Pers.) Burch, Monroe Station, Collier County, Florida,
~3 April 1964, Burch 3l6 (FLAS).

163.

(FLAS); Goodland, 22 Nov. 1964, Godfrey 65529 (FLAS, FSU). Dade: Miami,
1 Apr. 1897, Curtiss 5849 (F, NY); Brickell Hammock, Feb. 1911, Small,
Carter £■ Small 3274 (NY). Dixie: Shamrock, 24 Nov. 1964, Godfrey 65681a
(FSU). Glades: Palmdale, 28 Dec. I963, Burch 234 (FLAS). Highlands:
Sebring, 23 Apr. 1948, Garrett 6 (FLAS). Indian River: Merritt's island,
July, Curtiss 2496 (F, NY). Lake: Lake Louisa, 18 Nov. 1964, Burch &
Ward 525 (FLAS). Lee: Ft. Myers, 27 Sept. 1964, Burch 4l6 (FLAS). Levy:
Cedar Key, Apr. I876, Garber 8 (FLAS). Manatee: Long Boat Key, 27 Sept.
1964, Burch 430 (FLAS). Monroe: Pinecrest, 1 Apr. I93O, Moldenke 865
(DUKE, MO, NY); Big Pine Key, 10 May 1919, Cuthbert s.n. (FLAS); Soldiers
Key, 30 Mar. 1904, Britton 332 (F, NY). Polk: Lake Wales, 18 Nov. 1964,
Burch £■ Ward 526 (FLAS). Sarasota: Long Boat Key, 21 Nov. 1964, Godfrey
65238 (FLAS, FSU). BAHAMAS. Great Bahama: Eight Mile Rocks, 5-13 Feb.
1905, Britton £■ Millspaugh 2471 (F, NY). Berry islands: Little Harbor
Cay, 31 Jan. 1905, Britton £- Millspaugh 2247 (F, NY). New Providence:
Mt. Pleasant, 13 Sept. 1904, Britton £■ Brace 757 (F, NY); Grantstown,
28-29 May 1909, Wilson 8I9I (F, NY). Exuma: Cave Cay, 19 Feb. I905,
Britton £■ Millspaugh 2817 (F, NY). Long Island: Clarencetown S- vicinity,
16-19 Mar. 1907, Britton & Millspaugh 63 1 3 (F, NY). Inagua: Nov. I89O,
Hitchcock 32 (F, MO). South Caicos: Cockburn Harbour, 20 June 1954,
Proctor 8900 (IJ). CUBA. Habana: Rio Almendaras to Playa de Mariano,
22-23 Dec. 1910, Wilson 9470 (F) ; Vedado, 29 Sept. 1914, Ekman 185 (NY);
Batabano, 10 Apr. 1903, Shafer 515 (NY). Camaguey: Cayo Romano, 20 Oct.
1909, Shafer 2623 (NY). Oriente: Forest about Paso Estancia, 27 Apr.
1909, Shafer 1575 (NY). Pinar del Rio: Sierra de Anafe, 17 Dec. I9II,
Wilson 11385 (NY). JAMAICA. St. Andrew: Hope Road Terrace, Kingston,

164.

31 Oct. 1963, Adams II8IO (PUR, UCWl). St. Catherine: Sligoville, I6
June 1963, Adams 12556 (UCWi). St. Elizabeth: Malvern, 5 Sept. 1907,
Britton 1182 (F, NY); Bethlehem, Sept. I9OI, Harris 8291 (UCWl). West-
moreland: Torrington, 22 Feb. I963, Adams 12288 (UCWl). DOMINICAN REPUBLIC.
Santo Domingo City, 13 Jan. 1929, Ekman Hi 1127 (PUR); San Pedro de Macoris,
31 Mar. 1913, Rose, Fitch & Russell 4300 (NY). PUERTO RICO. Station grounds,
Rio Piedras, Bayamon, 2 Oct. 1937, Otero 120 (MO); Culebra island, 3-12
Mar. 1906, Britton & Wheeler 155 (NY). LESSER ANTILLES. St. Barthelemy,
Gustaria, 24 Nov. 1938, Questel 537 (NY); Antigua, Claremont, 2 Jan. 1938,
Box 1332 (BM, F, NY). Extra-territorial specimens: BERMUDA. Harrington
Sound, 4 Aug. 1913, Collins 337 (F, NY); North Shore, 31 Aug. -20 Sept.
1905, Brown & Britton 71 (F, NY); Kllderry Estates, Smith Parish, 26 July
1963, Manuel s.n. (A).

This taxon has often been considered no more than a variety of
_C. h i rta , but is clearly distinguishable, differing in fundamental charac-
ters of branching and cyathial production, and will be accepted as a dis-
tinct species here.

Jacquin's name _E. ob 1 i terata, Enum. Syst. PI. Carib., 22. 1762. and
Sel. Stirp. Am. Hist., 151. 1763., has been suggested as a synonym for
this plant (Boiss. in DC. Prod. 15(2): 21. 1862. and successive authors),
but this cannot be accepted. If this were the plant which Jacquin described
his name would have to be adopted for it, but his description is not
specific enough to be acceptable without authentic specimens to back it,
and, in fact, has features such as "umbel la trifida" which make it unlikely
that this is the plant to which he referred.

165.

30. CHAMAESYCE BERTERIANA (Balb.) MiUsp., Field Mus. Bot. 2:
303, 1909. Euphorbia berteriana Balbis in Spreng., Linnaeus' Syst.
Veg. ed. 16, 3: JSk. 1826. Type: ? "In ins. Guadeloupe et St. Domingue
(Bertero i)" Boiss. in DC. Prod. 15(2): 21. 1862. Specimen probably
in herb, istituto Botanico della Universita, Torino, if extant.

Euphorbia stipi tata Mi 1 Isp. , Field Mus. Bot. 2: 65. 1900.
Type: Guanica, Puerto Rico, 22 Jan. I899, Millspaugh 1782 (Holotype:
F, sheet 6I782, not found; isotype: F i).

Annual, decumbent to 2 dm. Stem 2 mm. thick, internodes to 3 cm.
long; branching throughout length, short- tomentose, brown. Leaves mem-
branous; blades ovate-elliptic or somewhat oblong, 5-15 x 3-8 mm., base
rounded, margin crenulate or blunt-serrate, apex rounded, short pilose,
yellowish green; petiole ca. 1 mm. long; stipules distinct or slightly
joined, linear, toothed, to 1 mm. Cyathia in dense leafless dichasia;
peduncle short, involucre narrowly campanulate, to 0.7 mm. diameter,
lobes triangular exceeding glands, somewhat cleft, strigose on outside
pilose within, glands subcircular, 0.2 mm. diameter, cupped, stipitate,
appendages a rim to twice width of gland, fifth gland linear, shorter
than lobes, sinus shallow. Staminate flowers 5-8 per cyathium, andro-
phores glabrous, to 1 mm. long at maturity. Gynophore pilose, to 1 mm.
long, exserted, nutant or upright; calyx obsolete; ovary pilose; styles
upright, 0,4 - 0.6 mm. long, joined at base, bifid for 1/2 length.
Capsule strigose, elliptic, 1.5 mm. long, 1.5 mm. wide at equator,
shal lowly three-lobed, angles acute. Seed long-cuneiform, to 0.9 mm.
long, 0.4 mm. wide between base and equator, strongly four-angled, angles
subequal, faces somewhat concave, transversely ridged, sometimes obscurely

166.

so, brown, red-brown or gray.

Weeds of cultivated ground and pasture, Bahamas, Cuba, Hispaniola
and Puerto Rico. Isolated collections from Antigua and Guadeloupe but
absent from Florida, Jamaica, and the southern part of the Antilles
chain (Map 31 ) .

Representative specimens seen; BAHAMAS. New Providence: Nassau,
Fort Charlotte, 6 July I96O, Webster, Samuel & Williams 10305 (PUR).
Great Exuma: Hayne Road, 22-28 Feb. 1905, Britton £■ Millspaugh 303I
(F, NY). Long Island: Clarence Town & vicinity, I6-I9 Mar. 1907, Britton
£- Millspaugh 6312 (F, MO, NY). Atwood Cay, 3"^ Dec. 1907, Wilson 7^13
(F, MO, NY). Mariguana: Southeast Point, 10-12 Dec. I907, Wilson 7587
(F, NY). Castle Island, 22 Dec. 1907, Wilson 78OI (F, NY). CUBA.
Havana: between Rio Cojimar and Playa de Bacuranao, 26 Dec. I9IO, Wilson
95^0 (F, NY). Oriente: Guantanamo Bay, 17-30 Mar. I909, Britton 2178
(MO, NY); El Morro, Santiago Bay, 10-11 Mar. 1912, Britton £■ Cowell
12588 (F, NY). HAITI. Massif de la Selle, 17 Nov. 1926, Ekman 698I
(PUR); Fond Parisien, Etang Saumatre, 5-13 May 1920, Leonard ^085 (F,
NY); Cabare, Bale des Moustiques, 12-18 Jan. 1929, Leonard & Leonard
119^^ (MO); Port au Prince, 1-2 Apr. 1920, Leonard 3^58 (NY); Petion-
ville, 15-28 June 1920, Leonard 4839 (NY); St. Michel de I'Atalay, 19
Nov. 1925, Leonard 7219 (NY). DOMINICAN REPUBLIC. Barahona: La Salinas,
17 Aug. 1946, Howard & Howard 8407 (NY); June I9II, Fuertes 1055 (F, MO,
NY). PUERTO RICO. Prope Salinas de Cabo Rojo, 8 Feb. 1885, Sintenis
634 (F, NY); Guanica, 5-8 Mar. 1915, Britton, Cowell & Brown 4899 (F,
MO, NY); Cayo Muertos, 9-12 Mar. 1915, Britton, Cowell S- Brown 4967
(F, MO); Guayanilla, 10 Mar. I913, Britton & Shafer 1820 (F, MO, NY);

67.

Map 31. Distribution of C. berteriana

>-:^^''

Map 32. Distribution of _C. rnacu ia ta

168.

Ponce to Penuelas, 22 Mar. I906, Britton & Cowel 1 I3O8 (F, NY); Yauco
to Guayanilla, 14 June-22 July I9OI, Underwood & Griggs 602 (F, NY);
8 miles west of Ponce, 1 Dec. 1902, Heller 6164 (F, MO, NY). LESSER
ANTILLES. Antigua, 1849, Wul 1 schlaegel 518 (MO); Guadeloupe, ex herb.
Grisebach, (MO); Guadeloupe, Desirade, 12 Jan. 1936, Stehle 290 (NY).

The species has clear affinities with _C. hi r ta and _C. opthalmi ca,
but has a somewhat more upright habit, rounder leaves with merely
crenulate margins, and usually smaller glomerules of cyathia. These
are terminal, and for this reason the species is felt to be closer to
_C. opthalmi ca than to _C. hi rta.

31. CHAMAESYCE MACULATA (L.) Small, Fl. SE. US., 713. 1903.
Euphorbia maculata L., Sp. PI. ed. 1: 455. 1753. Type: Herb. Linn,
sheet 630-1 1 i

E_. supina Raf., Amer. Monthly Mag. 2: 119. I8I7. Type: Unknown,
probably not extant. C. supina (Raf.) Moldenke, An annotated and classi-
fied list of H. N. Moldenke's collection numbers from No. 1 to 11, 277
inclusive, 135. 1939.

_C. tracyi Small, Fl. SE, US. 713. 1903. Type: Biloxi, Miss.,
15 July 1894, Tracy 2913 (NY i).

_C. mathews i i Small, Man. SE. Flora, 797. 1933. Type: Dunes
opposite Miami, Florida, 26 Nov. -20 Dec. 1913, Small, J. K. £■ Small,
G. J. 4556 (Holotype: NY !; Isotypes: DUKE i, FLAS !, FSU !, NCU '.) .

Annual; prostrate to decumbent forming loose mats to 5 dm. diameter,
Stem 1-2 mm. thick, internodes to 2 cm. or congested, branching through-
out length, subd i chotomous near base, alternate above with short leafy
laterals, not rooting at nodes, vi 1 lous-pubescent or tomentose or rarely

169.

velutinous, on all parts of stem, or lower surface glabrate or rarely
entirely glabrous, green or red. Leaves membranous; blades ovate-
elliptic or somewhat oblong or linear, 5-15 x 2-5 mm., base inequilateral,
rounded or obtuse, margins serrate, rarely subentire, apex rounded or
obtuse, vi 1 lous-pubescent, often glabrate above, rarely completely
glabrous, green or red, often with a red or purple spot in the middle
of the upper surface; petioles 1- 1.5 mm. long; stipules distinct,
linear, 0.7 - 1 mm. long, toothed or 2-3 parted. Cyathia solitary,
appearing clustered when borne on short congested leafy laterals near
the tip of the stem; peduncle 0.5 - 1 mm. long, cyathia obconical, 0.6 -
0.8 mm. diameter near mouth, lobes triangular, slightly exceeding glands,
entire or minutely toothed, sparsely villous on outside, pilose within,
glands transversely elliptic or lunate, 0.1 - 0,25 mm. long, appendages
narrower than to twice as wide as gland, margin entire or lobed, white
or pink, fifth gland linear, almost equalling glands, sinus deep.
Staminate flowers rarely more than five per cyathium, androphores glab-
rous, 1 mm. long at maturity. Gynophore pilose, occasionally glabrous,
1 - 1.5 mm. long at maturity, barely exserted and bent to lie deep in
sinus; calyx a triangular pad of tissue; ovary strigose, rarely sub-
glabrous; styles spreading, 0.2 - 0.4 mm. long, minutely joined at base,
bifid for 1/3 - 1/2 length. Capsule strigose, often glabrate, ovoid, to
1.5 mm. long, 1.5 mm. wide below equator, strongly three-lobed, angles
acute. Seed oblong-ovoid, 0.8 - 1 mm. long, 0.6 mm. wide below equator,
strongly four-angled, angles subequal, faces flat or somewhat convex,
obscurely transversely ridged or almost smooth, ridges sometimes passing
into angles, tan, red-brown or ashen.

170,

Plaota ui FloixU

Plate 37. CHAMAESYCE MACULATA (L.) Small

C. maculata (L.) Small, Charlotte Harbor, Charlotte County, Florida,
ll Sept. 1964, Burch 4l8 (FLAS).

171

TYPE

Chanaesyce MathewsH onall,
SE ?!., 797, 1505. 1933.

Louis c. (Theeler 19;

Plate 38. CHAMAESYCE MACULATA (L.) Small

Holotype- C. ma thews i i Small, sand dunes opposite Miami, Dade County,
Florida, '26 Nov. -20 Dec. 1913, Small & Small 4556 (NY).

72.

A common weedy plant of the United States. Found In disturbed
habitats throughout Florida except on the Keys where _C. b lodgett i i
replaces It as the characteristic plant of sidewalk cracks and gas
station forecourts. Present in Bermuda but not in the Bahamas or
any part of the Caribbean (Map 32).

Representative specimens seen: FLORIDA. Alachua: University of
-lorlda campus, Gainesville, 9 Oct. 1956, Jackson 702 (FLAS, FSU).
Say: Gulf Resort Beach, 19 Oct. I963, Burch 213 (FLAS). Bradford:
iampton, 29 July 1927, Wiegand and Manning I8I6 (GH). Brevard: Canova
3each, 17 April 1965, Burch 63O (FLAS); Okeechobee region, 6 Sept. 1903,
-redholm 5990 (GH, NY). Broward: New River Sound, 18 Nov. 1903, Small
r Carter 1 1 63 (NY); Deerfleld, 20 March I965, Lakela 28291 (FLAS, USF).
:alhoun: Blountstown, 25 Oct. 1964, Burch 508 (FLAS). Charlotte: New
^olnt Comfort, 5 June 1964, Burch 351 (FLAS); Punta Gorda, 8 July 19^8,
jarrett s.n. (FLAS). Citrus: Hernando, 18 May I965, Ward and Beckner
t676 (FLAS); Lake Tsala Apopka, 8 June I96I, Godfrey S- Reinert 60884
;FSU). Collier: Cape Romano, 28 Aug. 1922, Small, Small & DeWinkeler
10620 (NY); Naples, 6 June 1964, Burch 36? (FLAS); Marco Island, 26 Sept.
1964, Burch 405 (FLAS). Dade: Sand dunes opposite Miami, 26 Nov. -20 Dec.
1913, Small & Small 4553 (DUKE, FLAS, NY); Border of Everglades, 7 March
1892, Simpson 557 (GH); Biscayne Bay, 1874, Palmer 7404 (MO); Key B i s-
;ayne, 6 May 1958, Atwater M-42 (FLAS). DeSoto: Arcadia, 29 Nov. I962,
yard & Burch 3217 (FLAS). Dixie: Shamrock, 24 Nov. 1964, Godfrey 65684
;FLAS, FSU). Duval: Jacksonville, 11 October 1894, Curtiss 5118 (MO,
JY); Jacksonville, 22 Dec. 1924, Small II56I (NY). Escambia: Pensacola,
Jl July 1941, Brinker 356 (MO) . Flagler: Andulusia, 10 Oct. 1940, West &

173.

Arnold s.n. (FLAS). Franklin: Lanark, 6 Oct. 1955, Godfrey 5^128
(DUKE, FSU, GH, NY). Gadsden: Chattahoochee, 13 Sept. 1956, Krai 6-
Godfrey 355^ (FSU, GH) . Gilchrist: Trenton, 5 Oct. 19^0, West £■
Arnold s.n. (FLAS). Gulf: Cape Sand Bias, 8 Nov. 1959, Godfrey 59163
(FSU). Hamilton: White Springs, 26 July 19^6, West & Arnold ih (FLAS).
Hardee: Wauchula, 26 Sept. 196^+, Burch 389 (FLAS). Hernando: Brooks-
ville, 12 July 1958, Godfrey 57208i (FSU). Highlands: Sebring, 23 Apr.
19^8, Garrett 7 (FLAS); Junction US 27 & FLA 64, 2 January I963, Burch
60 (FLAS). Hillsborough: University of South Florida campus, Tampa,
13 Aug. 1964, Burch 192 (FLAS); 5 Nov. 1904, Fredholm 6469 (GH, MO, NY).
Holmes: 9 miles E. of Walton County line, 25 Oct. 1964, Burch 494 (FLAS).
Indian River: Vero Beach, 7 August 1963, Burch 115 (FLAS). Jackson:
Marianna, 19 Oct. I963, Burch 211 (FLAS). Jefferson: Monticello, 4
Oct. 1964, Elam s.n. (FLAS); Capps, 9 Oct. 1954, Godrey & Morrill 52542
(FSU, GH). Lafayette: Dowling Park, 15 July I96I, Ramsey, Godfrey £■
Mitchell 247 (FSU). Lake: Eustis, I6-3O June 1894, Nash 1058 (GH, MO,
NY). Lee: Alva, July-August I9OO, Hitchcock 328 (GH, MO, NY); Sanibel,
outer island, 20 July 1954, Cooley 2330 (USF). Leon: Tallahassee, 30
June 1955, Godfrey 53572 (DUKE, GH, FSU, NY). Levy: Seahorse Key, 1 Dec.
1962, Ward 3256 (FLAS). Liberty: Jackson Bluff, 30 Sept. 1964, Godfrey
64677 (FLAS, FSU). Manatee: Longboat Key, 27 Sept. 1964, Burch 443
(FLAS); Bradenton, 20 Nov. 1964, Godfrey 65192 (FLAS, FSU). Marion:
Lake Kerr, 24 Sept. 1941, West & Arnold s.n. (FLAS). Martin: Jupiter
Island, 7 Aug. I963, Burch 126 (FLAS). Nassau: Fort Clinch State Park,
12 Oct. 1963, Burch 197 (FLAS). Okaloosa: Destin, 24 Oct. 1964, Burch
467 (FLAS). Okeechobee: Okeechobee, 27 Dec. 1964, Burch 588 (FLAS).

174.

Orange: 6 Aug. 1902, Fredholm 5^5^ (GH, NY). Osceola: k miles E. of
Polk County line on FLA 60, 7 Aug. I963, Burch IO6 (FLAS). Palm Beach:
Jupiter Island, 27 July 1956, Cooley, West S- Daggy 4778 (GH, USF) ;
Boca Raton Lake, 19 Nov. 1903, Small & Carter 1146 (NY). Pasco: Land
o'Lakes, 10 July I958, Krai 7257 (FLAS, GH) . Pinellas: Sunshine Sky-
way Bridge, St. Petersburg, 15 Oct. 1964, Burch 452 (FLAS); Fort DeSoto
County Park, Mullet Key, 10 Sept. I963, Thorne, Long & Lakela 26325
(USF). Polk: Timberlane, Lake Wales, 1 Oct. 1964, Conard s.n. (FLAS).
Putnam: Welaka, 12 July 1940, Laessle s.n. (FLAS). St. Johns: North
River, St. Augustine, 12 Oct. I963, Burch 203 (FLAS). St. Lucie: South
side Ft. Pierce, 7 Aug. 1963, Burch 124 (FLAS). Santa Rosa: Navarre,
24 Oct. 1964, Burch 477 (FLAS). Sarasota: Longboat Key, 21 Nov. 1964,
Godfrey 65239 (FLAS, FSU). Sumter: Wildwood Exit from 1-75, 18 Nov.
1964, Burch & Ward 523 (FLAS). Suwannee: East of Ellaville, 15 July
1961, Ramsey, Godfrey &■ Mitchell 286 (FSU). Taylor: Adams Beach, 21
Oct. i960, Godfrey & Houk 60458 (FSU, GH) . Union: Worthington, 24
Aug. 1964, Warren s.n. (FLAS). Volusia: Green Mound, south of Daytona
Beach, 23 Aug. 1922, Small, Small £■ DeWinkeler 10571 (GH) ; Daytona
Beach Shores, 3 Aug. I962, Ray 11144 (USF). Wakulla: St. Marks Wild-
life Refuge, 6 Oct. 1955, Godfrey & Krai 54104 (FSU, GH, USF). Wash-
ington: Careyville, 25 Oct. 1964, Burch 498 (FLAS). Extra-territorial
specimens: BERMUDA. Old Ferry Road, 9 Aug. 1913, Collins 217 (GH, NY);
North Shore, 31 Aug. -20 Sept. 1905, Brown & Britton 12 (GH, NY).

A very variable species in growth habit, color, pubescence and
appearance of seed. The very similar Euphorbia humistrata, reported by
Wheeler (1941) from Alabama, has not been found in Florida although

175,

some collections are suggestive of this species in their pubescence
and seed.

Small's species, _C. t racy i and C_. ma thews i i are based on
extreme forms of this species, and examination of a long series has
shown a range of intermediates that span the differences,

Wheeler's rejection (1939, I960) of the long-established name
for this species has been mentioned in the commentary on _C, nutans.
His case was never proved, and the traditional application of the
epithet to this prostrate species should be maintained. The matter
is discussed at some length in Burch (1965), which also gives a list
of contributions to the controversy stirred up by Wheeler's proposal.

32. CHAMAESYCE THYMIFOLIA (L.) Millsp., Field Mus. Bot. 2:
412. 1916. Euphorbia thymifol ia L., Sp. PI. ed. 1: kSk, 1753. Type:
not sheet 63O-IO in Herb. Linn. No specimen known to have been seen
by Linnaeus has been found. Anisophyllum thymifol ium (L.) Haw., Syn.
PI. Succ, 160. 1812. "_E. maculata L. (var. thymifol ia L.)" Griseb.,
Fl. Brit. W. I. Islands, 53. 1859., cited in this form.

Annual; perhaps perennating by thick rootstock, prostrate,
forming mats to k dm. diameter. Stem 1-2 mm. thick, internodes to 1.5
cm. long or congested; sparingly branched near base, and with short,
congested laterals or unbranched at tips of stem, not rooting at nodes,
vi 1 lous-pubescent or tomentose on upper surface, often glabrate, naked
on under surface, green or light brown. Leaves membranous; blades
oblong-elliptic, often narrow and very uniform on side shoots, 4-10
X 2-5 mm,, base oblique or subequ i latera 1 on side shoots, margin coarsely
serrate, apex rounded to acute, upper surface usually glabrous, lower

176.

sparsely tomentose, glabrate, green or yellowish; petioles 0.5 - 1 mm,
long; stipules distinct, sometimes partly joined on underside of stem
near tip, linear and toothed or 2-3 parted, 0.7 - 1 mm. long. Cyathia
solitary or in pairs or even few-flowered glomerules at nodes by
reduction of branches, sometimes appearing clustered on congested
laterals; peduncle short, involucre obconical, O.k - 0.6 mm. diameter,
lobes triangular, shorter than or equalling the glands, strigose out-
side, glabrous inside, glands broadly elliptic or subcircular, 0.05 -
0.2 mm. long, pink or red, appendages small, rimming gland or to twice
width, the pair nearest sinus sometimes extended in length toward sinus,
pink, fifth gland absent, sinus extending almost depth of involucre.
Staminate flowers rarely more than five per cyathium, androphores glab-
rous, 1 mm. long. Gynophore strigose, ca. 0.5 mm. long, not exserted
but forcing cyathium apart as it bends to push capsule through sinus;
calyx obsolete; ovary strigose; styles upright, O.k - 0.6 mm. long,
minutely joined at base, bifid for 1/3 length. Capsule strigose, long-
ovoid, 0.8 - 1 mm. long, 1 mm. wide below equator, three-lobed, angles
acute. Seed conical-ovoid, to 0.9 mm. long, 0.5 mm. wide below equator,
strongly four-angled, faces concave, obscurely transversely ridged,
ridges extending somewhat into angles, tan-colored.

Present in three counties on the west coast of Florida. Not
known from the Bahamas, and no specimens seen from Cuba, but a weed
of waste places, often near salt water, through the remainder of the
Antilles and in Central and South America (Map 3^).

Representative specimens seen: FLORIDA. Charlotte: Charlotte Har-
bour, 27 Sept. 1964, Burch 417 (BM, FLAS, GH, NY, US). Lee: Fort Myers,

177.

Plate 39. CHAMAESYCE THYMIFOLIA (L.) Mi lisp.

C. thymi fol ia (L.) Millsp,, Charlotte Harbor, Charlotte County, Florida,
27 Sept. \3Gk, Burch 417 (FLAS).

178.

13 Aug. 1963, Burch l86a (FLAS) ; Fort Myers, 27 Sept. 196^, Burch 4l3
(BM, F, FLAS, FSU, GH, NY, US). Pinellas: Dunedin, 15 Oct. 1964, Burch
451 (FLAS). JAMAICA. St. Mary: Green Castle, 5 July I963, Crosby,
Hespenheide £• Anderson 476 (DUKE, PUR, UCWl); Sea Wall, 3 Sept. 1957,
Robertson 3333 (UCWl). St. Andrew: Mona, 25 Mar. I96O, Adams 6598
(UCWl). DOMINICAN REPUBLIC. Peninsula de Samana, 28 Apr. 1930, Ekman
HI4792 (A, F); Higuey, Seybo, 14 Apr. 1929, Ekman H12179 (PUR). PUERTO
RICO. Catano, 1 Apr. 1930, Britton & Britton 9550 (NY); Mayaguez, 17
Oct. 1884, Sintenis 34b (GH, NY). LESSER ANTILLES. Guadeloupe,
Desirade, 1893, Duss 2738 (NY); Dominica: Resale Valley, I903, Lloyd
739 (F); mouth of Pagoua River, 18 Apr. 1940, Hodge 5- Hodge 306I.
Martinique: Ancien Riviera Blanche, 18 Feb. 1939, Stehle 3468 (A);
Military Cemetery, St. Pierre, 1899, Duss 4056 (F, NY); Fort de France,
Marigot, Aug. I89O, Duss 4455 (NY). St. Vincent: Byera, Charlotte
Parish, 9 Mar. I962, Cooley 8479 (GH). Grenada: vicinity of St.
Georges, 30 Oct. -11 Dec. 1957, Proctor 16824 (A, IJ); The Bower,
St. Georges, 26 Dec. 1904, Broadway s.n. (F).

It is not certain whether Linnaeus ever saw the plant to which
his name _E. thymi fol ia is usually applied, since the sheet labelled
"20 E. thymifolia" in the Linnaean Herbarium is not that species but
C_. maculata. The descriptions to which he refers in synonymy are
scarcely explicit enough to give a positive identification of the plant
involved, but the application of the name is so uniform, at least in
the New World, that it should be maintained unless it can be demonstrated
that Linnaeus had some other intention.

The species is very close to the group of species with greatly

179.

Map 33. Distribution of _C. helwiqi i and _C. paredonens i s

Map 34. Distribution of _C. thytnifol la

180.

enlarged glandular-appendages in two pairs of unequal size. it differs
from them in having very small glands, and in its much smaller appendages,
which are usually insignificant even when one pair can be seen to be
larger than the other. The capsule in _C, thymi fol ia is not exserted,
but splits the cyathium as it enlarges. This is sometimes seen in other
species, but not to the same degree as in _C. thymi fol ia where the cyathium
finishes life as a fringe at the lower end of one side of the mature
capsule.

33. CHAMAESYCE ADENOPTERA (Bertol.) Small, Fl. SE. US., 71^.
1903.

Perennial; prostrate or decumbent, forming tufted mats to 2 dm.
diameter, or more open, to 3 dm. Stem ca. 1 mm. diameter, internodes to
2 cm. long or much shorter in Florida plants; sparsely branched, mostly
from upper part of stem, not rooting at nodes, strigose or rarely pilose
on upper surface, glabrous below, straw-colored or darker. Leaves mem-
branous to coriaceous; blades oblong-elliptic or reniform, 4-10 x 2-6
mm., base oblique, rounded or cordate, margin serrate, sometimes recurved,
apex obtuse, glabrous or strigose on lower or both surfaces, green, often
suffused purple; petiole ca. 1 mm. long; stipules separate or joined at
base, attenuate, to 1 mm. long. Cyathia solitary at nodes, often
collected on short laterals; peduncle short, involucre campanulate, to
0.6 mm. diameter at mouth, lobes scarcely equalling glands, deltoid,
strigose on outside, ciliate within, glands transversely elliptic, cupped,
to 0.5 mm. long, the pair flanking the sinus longer than the other,
appendages prominent, the pair by sinus larger, to 2 mm. long and three
times width of gland, margin entire or crenulate, white, fifth gland short

181

or obsolete, sinus deep. Staminate flowers 8-18 per cyathium, andro-
phore glabrous, to 1 mm. long at maturity. Gynophore strigose, to
1,2 mm. long at maturity, exserted, strongly nutant, capsule almost
hidden by appendages; calyx obsolete; ovary strigose; styles upright,
0,6 mm. long, joined at base, bifid for 1/3 length. Capsule strigose,
globose-ovoid, base truncate, to 1,2 mm. long, 1.4 mm. wide just above
base, angles rounded. Seed ovoid, 1 mm. long, 0.4 - 0.6 mm. wide below
equator, angular or with angles obscure, faces flat with transverse
ridges or broadly convex with short transverse grooves, dark red or
wh i te.

Key to subspecies of Z_, adenoptera
1, Stems few, sparingly branched, usually ca. 1 mm. diameter, often
strongly decumbent; leaves usually 5-10 mm. long; seed angular,
faces flat with transverse ridges a. ssp, adenoptera.

1. Stems several to many, often branching freely in upper part, rarely
exceeding 0.5 mm. diameter, usually prostrate; leaves rarely reach-
ing 5 mm. long; seed plump, angles rounded, obscure, faces marked
with short transverse grooves 2.

2. Stems several, internodes short forming a compact plant; appendages
of glands prominent, larger pair usually more than 1 mm. long

b. ssp. perqamena.

2. Stems many, internodes often long, giving plant an open appearance;
appendages of glands prominent but larger pair usually less than 1
mm. long c. ssp. qundlachi i.

182.

33a. C. ADENOPTERA (Bertol.) Small ssp. ADENOPTERA Euphorbia
adenoptera Bertol., Misc. Bot., 3: 20, t. 23. 1843.

Pinelands and dry rocky areas in the mountains of Hispaniola
(Map 35).

Representative specimens seen: HAITI. Hinche, Artibonite,
23 Sept. 1955, Proctor 10931 (j); Bassin Bleu, 14-27 April 1929,
Leonard & Leonard 15010 (GH, NY); Fond Parisien, Etang Saumatre, 5-13
May 1920, Leonard 4092 (NY); Marmelade, 25 Aug. 1903, Nash 799 (F, NY);
Morne des Commissa i res , 14 Sept. 1955, Proctor 10712 (IJ); Morne
Rochelois, Miragoane, 17 Nov. 1926, Ekman H7242 (A, F) ; Port au Prince,
1-2 Apr. 1920, Leonard 3459 (NY); San Michel, 5 Aug. 1905, Nash £■
Taylor 1436 (F, NY). DOMINICAN REPUBLIC. Barahona: Trail between
Pedernales and Aceitial, 8-12 Aug. 1946, Howard & Howard 8I36 (GH,
NY); 1912, Fuertes 693 (F, GH, NY). Constanza, February I9IO, von
Turckheim 2930 (F, GH, NY); Moncion, Monte Cristy, 9 June 1930, Valeur
92 (MO, NY); S. Jose, Santiago, 8 Apr. 1933, Valeur 970 (NY); Sabana
de Santa Rosa, Santo Domingo, 11 Oct. 1947, Allard 15936 (MO, NY).

This Is a stronger and much more diffuse plant than the other
two subspecies. It is maintained at this level because some collections
come close to ssp. pergamena in leaf shape and pubescence, but the seed
differs so much that a good case could be made for separating it at
the level of species.

33b. C. ADENOPTERA (Bertol.) Small ssp. PERGAMENA (Small) comb,
nov. Euphorbia pergamena Small, Bull. Torr. Bot. Club 25: 615. I898.
Type: Miami, November I878, A. P. Garber s.n.; Lemon City, 2 March I892,
J. H. Simpson 523 (Syntypes: NY 1); duplicate collections Simpson 523:

183.

F i, GH 1, NY 1, US I). Chamaesyce perqamena (Small) Small, Fl. SE. US.,
713. 1903.

Euphorbia lindeniana A. Rich, in Sagra, Hist, Cub., II: 197.
1850. Type: Cuba, 18^3-44, Linden 1706 (Herb. Mus. Paris, not seen;
isotypes: F 1, NY i). Anisophyllum lindenianum Kl. & Gke., Tricocc.,
33. 1859.

Chamaesyce monensis Mi lisp.. Field Mus. Bot. 2: 389. 191^.
Type: Limestone Plateau, Ubero, Mona Island, 20-26 Feb. 191^, Britton,
Cowell £- Hess l84l (Holotype: F !; isotype: NY i).

Crevices in limestone rock or in sand, usually at low altitudes,
Dade and Monroe counties of southern Florida, Cuba, Hispaniola and Mona
Island (Map 36).

Representative specimens seen: FLORIDA. Dade: Campbell Drive,
Homestead, 23 June I96O, Webster and Samuel 1009^ (DUKE, PUR); Arch
Creek Prairie, 3 July I9I5, Small, Mosier & Small 6809 (DUKE, FLAS,
NY); Lemon City, 30 July I9IO, Harper 83 (MO, NY); Miami, 17 March
1930, Moldenke 5827 (NY); Between Cocoanut Grove and Cutler, 31 Oct.-
4 Nov. 1903, Small & Carter 757 (NY); Long Pine island between Camp
Jackson and Long Key, Everglades, 9 Jan. 1909, Small 5- Carter 3I89 (NY);
Pinelands south of the Miami River, 20 Nov. 1912, Small 3851 (NY).
Monroe: Big Pine Key, k Feb. 1952, Killip 41882 (F); Big Pine Key, 4
Sept. 1957, Brass 28985 (FLAS); Big Pine Key, 2 Dec. 1912, Small 3980
(NY). HAITI. Jean Rabel , 2 7 Jan.-9 Feb. 1929, Leonard & Leonard 12867 (NY).

This subspecies is very close to the next, and is separated
mostly because the Florida plants form such a homogeneous group. They
differ on the size of appendages and in having a more compact growth

\8k.

habit than ssp. gundlachl i .

_C. monens i s is known only from the type collection, and is
included here until it can be shown that the lack of appendages on
the glands is a constant character.

33c. C. ADENOPTERA (Bertol.) Small ssp. GUNDLACHII (Urban) comb,
nov. Euphorbia qundlachii Urban, Symb. Ant., 5: 392. I9O8. Type: Cuba,
1865, Wright s.n. (Herb. Mus. Bot., Berlin, if extant).

In sand or rocky places, often at the coast, Cuba (Map 37).
Representative specimens seen: CUBA. Havana: Cojimar, 2k Aug.

1910, Britton, Earle & Gager 6281 (F, NY); Guanabo, August I9I6, Cazanas
340 (F, NY); Between Rio de Cojimar and Playa de Bacuranao, 26 Dec. I9IO,
Wilson 9532 (F, NY). Isle of Pines: Caleta cocodrilos, 8 March I9I6,
Britton, Wilson £■ Leon 15250 (F, NY); Pedernales Point, I6 Feb. 1899,
Millspaugh 14-36 (F). PInar del Rio: Laguna jovero and vicinity, 5-7 Dec.

1911, Shafer 10840 (F, NY); Las Martinas to the coast, 19 Dec. I9II,
Shafer 11093 (F, NY). Oriente: Baracoa, 26 Jan. 1915, Ekman 4436 (NY).
Santa Clara: Castillo de Jagua, Sept. I9II, Cuesta 248 (F, NY); Cien-
fuegos, Aug. 1913, Cuesta 746 (F, NY); Playa de Buenaventura, Bahia de
Cochinos, 14 Aug. 1920, Leon & Loustalot 9547 (F, NY).

The application of this name seems to be well-understood, and
has been followed here although no type material has been seen. The
usual appearance of the plant is very diffuse with many fine wiry stems,
and long internodes holding the small leaves well apart. Other specimens
approach ssp. pergamena in general appearance, but have much smaller
appendages to the glands than is typically found In that taxon.

These subspecies were combined as a single species, C. adenoptera ,

185.

)PTERA (Bertol.) Small var. PERGAMENA (Small) ined.

Plate '.O. C. ADENOf

p oerqamena Small, Miami, Florida, Nov. I878, Garber s.n. (NY)
Syntype: E. £3^^

186.

i^-4.-

r^

ilililililililililililililili

Jll|llll|liil|ilil|iiii|illljnnpilt|iiii|iiiniiii|iMinnijiiH|iiii|Mn|mmi«m^

fflilllllllilillUhllllllllllJllllllllllllllRllllllllllllIll!!

i

i.iirvn or

FiciA Nji.K'dt- 14,-j Ku to? ^ J»r. '

IME NEW YORK BOIANICAL CAKOt >. ' '
Wt-LuSATlOfi f.r »^p»Ti> tilt,

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

Isotype: C. monensis Mi lisp,, Mona Island, Porto Rico 20-?fi f^k low
Britton, Cowell & Hess 18^+1 (NY). ' ^° ' '9"^,

187.

J*

<|ill|MMI|IMI|llll|ltll|l|ll|IMI|lltl|ltll|l'i'

' YORK BOTANICAL QAROCN

ilniliiJiiiliiimi

1,1,1, i,l,iJ^,liiJ,i,uf^~'"

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

_C. adenoptera (Bertol.) Small, between Rio Coj imar and Playa de Bacuranao,
Prov. of Havana, Cuba, 26 Dec. 1910, Wilson 9532 (NY).

^ \

Map 35. Distribution of C. ad

enoptera ssp. adenoptera

Map 36. Distribution of C. adenopt

noptera ssp. pergamena

189.

Map 37. Distribution of _C. adenoptera ssp. gundlachi

Map 38. Distribution of C. conferta

190,

by Millspaugh, Field Mus. Bot. 2: 388. 1914. In this he followed
Boissier in DC. Prod. 15(2): 49. 1862., but differed in accepting
_E. d ioica Kunth as a distinct species, while Boissier rejected even
the name on the grounds that the plants were monoecious. Millspaugh's
treatment was, perhaps, too sweeping in the combination of taxa, although
the three separated as subspecies here are certainly too close to be
considered distinct at the specific level.

34. CHAMAESYCE COiMFERTA Small, Fl. SE. US., 713. 1903. Type:
Cocoanut Grove, Florida, 2 S- 5 Nov. 1901, J. K. Small & G. V. Nash s.n.
(Lectotype: lower RH plant on sheet, NY 1; Isotypes: NY '. , F !). Euphorb ia
conferta (Small) B. E. Smith, J. Elisha Mitchell Sci. Soc. 62: 82. 1946.

Annual or perennating; prostrate or decumbent, one to several stems
from crown of root, to 2 dm. long. Stem to 1.5 mm. diameter, nodes to
1.5 cm. or congested; branched throughout length, upper laterals short
and congested, villous on upper surface, naked below, red or rarely green.
Leaves membranous, somewhat coriaceous; blades oblong-oblanceolate, 5-10
X 3-5 mm., base markedly oblique, margin serrate, somewhat thickened, apex
acute to obtuse, sparsely pilose, quickly glabrate, red or rarely green;
petiole 0.5 - 1 mm. long; stipules distinct or minutely joined at base,
subulate, to 1 mm. long, sometimes toothed or slightly parted. Cyathia
solitary at nodes of congested laterals in upper part of stem; peduncles
short, involucre narrowly campanulate, 0.7 mm. diameter at mouth, lobes
scarcely equalling glands, deltoid, strigose on outside, pilose within,
glands transversely elliptic, to 0.3 mm. long, curved to follow rim of
cyathium, cupped, dark red, appendages unequal, the pair near the sinus
twice as wide as gland and elongated in length toward sinus concealing

191

the cyathium, the other pair to twice as wide as gland but not elongated,
margin crenate, pink or red, rarely white, fifth gland obsolete or small,
sinus deep. Staminate flowers 5-8 per cyathium, androphores glabrous,
to 1 mm. long at maturity. Gynophore pilose, to 0.8 mm. long, bent
and short exserted through sinus but not distorting cyathium; calyx a
triangular pad of tissue; ovary strigose; styles upright to 0.6 mm. long,
joined at base, bifid for 1/3 length; capsule strigose, conical or
truncate-ovoid, 1.2 mm. long, 1.2 mm. wide above base, three-sided,
angles obtuse. Seed ovoid, 1 mm. long, 0.^ - 0.6 mm. wide between base
and equator, angular, faces flat or convex with transverse ridges, some-
times very plump and faces grooved rather than ridged, red or white.

Pinelands and oolitic limestone of Everglades Keys and lower
Florida Keys (Map 38).

Representative specimens seen: FLORIDA. Dade: Pinelands about
Sykes Hammock, 4 Mar. 1915, Small, Mosier, £■ Small 5655 (DUKE, FLAS, GH,
NY); Biscayne Bay, 19 June 1895, Curtiss 5^86 (FLAS, GH, MO, NY); Jenkins
to Everglades, 10 Nov. 1903, Eaton 206 (F) ; Pinelands near Long Prairie,
2k Mar. 1904, Britton 211 (F, NY); Florida City, 23 Nov. 1964, Godfrey
65653 (FLAS, FSU); Long Pine Key, 9 Mar. 1956, Craighead s.n. (FLAS);
Junction of Old Cutler Road and Red Road, 11 Aug. I963, Burch I56 (FLAS);
Brickell Hammock, Feb. I9II, Small, Carter & Small 3277 (NY). Monroe:
Big Pine Key, 25 Mar. -7 Apr. 1950, Killip £- Swallen kOkGj (BUS); Big
Pine Key, 1 Sept. 195^, Thorne 15085 (U); Big Pine Key, 20 Nov. 1964,
Burch & Ward 539 (FLAS).

This taxon may prove to be referable to Chamaesyce dioica (HBK.)
Mi lisp.. Field Mus. Bot. 2: 384. 1914. In making the combination

192.

iJ0M

4^

A

, < |niiMiii|iiii|nii|iiii|iiii|iniiirii|iiii|nii|iiiiimi|iiii|nii|iin|iiiiinn|iiim^

EVERGUDE KEYS

*ORK aOTANICAt. OAnOih

r^j

,a.^..-^^

Pla

te i+3. CHAMAESYCE CONFERTA Smal

Lectotype: C. conferta Small, Cocoanut Grove, Florida, 2 & 5 Nov. 1901
Small & Nash s.n, (NY).

193.

Millspaugh gives the range of the species as Brazil to Colombia and
through Central America to Lower California, and also cites a single
collection from Santo Domingo. No authentic material of the species
has been seen, but the description suggests that it has a strong
morphological resemblance to the plants here carried under Small's
name _C. conferta. and the possibility that the two are conspecific
cannot be ignored.

Some collections of _C. thymi fo1 ia come very close to this species,
but can usually be distinguished by the marked splitting and distortion
of the cyathium as their capsules enlarge, the smaller appendages to
the glands, and the small glands which rarely show any of the cupping
or enlargement to outline the rim of the cyathium that is found in
_C. conferta. The phylogenetic relationship of the two species is a
matter for conjecture, and an examination of _C. dioica and the other
species of Central and South America discussed by Millspaugh (loc. cit.)
might be illuminating in this respect.

35. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small, Fl. SE. US.,
710. 1903. Euphorbia garberi Engelm. ex Chapm., Fl. Southern US. ed. 2,
646. 1883. Type: Coast of South Florida, undated, Garber (Lectotype:
MO 1; Isotype: MO 1, F i).

Chamaesyce adicioides Small, Fl. SE. US., 710. 1903. Type: E. of
Bahia Honda, Monroe County, Florida, 20 Jan. 1920, J. H. Simpson 464
(Holotype: NY 1; Isotypes: F i US 1).

Chamaesyce brachypoda Small, Fl. SE. US., 709. 1903. Type: Miami,
Dade County, Florida, 4-7 Apr. I898, C. L. Pollard and G. N. Collins 244
(Holotype: NY 1; Isotype: US 1).

19^.

Chamaesyce mosieri Small, Torreya 27: 93. 1927. Type: Pine-
lands about Brogdon Hammock, Dade County, Florida, 19 June 1915, J. K.
Small and C. A. Hosier 63^7 (Holotype: NY I).

Annual or perennating; prostrate to strongly decumbent, forming
mats to 4 dm. diameter. Stems few to several from root stock, to 2 mm.
diameter, internodes to 3 cm. long or congested on laterals; branching
throughout length with few, subequal branches near base and small con-
gested laterals near tip, not rooting at nodes but occasionally acting
as runners to form new plants, strigose to long-pilose, sometimes woolly
and canescent, tan or purple-brown. Leaves membranous; blades ovate to
oblong-elliptic, 4-9 x 3-6 mm., base inequilateral, obtuse or rounded,
margin entire or rarely obscurely toothed near apex, apex rounded or
obtuse, sometimes minutely apiculate, midrib sometimes prominent on under
surface, strigose or long-pilose, sometimes glabrate, yellow-green or
suffused with red or purple; petiole 0.5 to 1 mm. long; stipules joined,
deeply bifid, toothed, to 0,6 mm. long. Cyathia solitary, collected on
congested laterals; peduncle short, involucre campanulate, to 1 mm.
diameter at mouth, lobes exceeding glands, deltoid, somewhat attenuate,
entire or few-toothed, strigose, sometimes glabrate on outside, ciliate
within, glands broadly transversely elliptic, 0.3 x 0.2 mm., appendages
minute to approximately equalling glands, margin entire or crenulate,
fifth gland obsolete, sinus narrow, to half depth of cyathium. Staminate
flowers 8-20 per cyathium, androphores glabrous, to 0.8 mm. long at
maturity. Gynophores glabrous, to 1.5 mm. long at maturity, exserted,
upright or nutant; calyx a triangular pad of tissue; ovary strigose; styles
spreading, to O.k mm. long, joined at base, bifid for half length. Capsule

195.

strigose or pilose, broadly ovate, to 1,6 mm. long, 2.1 mm. wide below
equator, broadly three-lobed, angles acute to obtuse. Seed oblong-ovoid,
to 1.2 mm. long, 0.6 to 0.8 mm, wide between base and equator, four-
angled, angles subequal, ventral faces slightly concave, smooth, dorsal
faces flat or convex, obscurely wrinkled, red-brown or ashen.

Restricted to pinelands and coastal sands of Dade and Monroe
counties, Florida (Map 39).

Representative specimens seen: FLORIDA. Dade: Pineland south
of Miami River, 26 Nov. -20 Dec. 1913, Small and Small 4752 (NY); Between
Miami and Kendall Station, 5 Nov. 1906, Small & Carter 2508 (NY); Between
Homestead and Camp Jackson, 1 Nov. 1906, Small & Carter 2517 (NY).
Monroe: Middle Cape Sable, 18 Apr. 1964, Burch 306 (BM, DUKE, E, F,
FLAS, GH, IJ, MO, NCU, NSC, NY, PUR, UCWI , US, USF) ; Middle Cape, Cape
Sable, 2 Apr. I960, Small 7687 (DUKE, FLAS, MO, NY); Upper Matecumbe Key,
26 Nov. 1912, Small 3900 (NY); No Name Key, 30 July 1895, Curtiss 5511
(F, GH, MO, NY); Long Key, 28 Feb. 1921, Hunnewell 7316 (GH); Grassy
Key, 28-29 Jan. 1909, Small & Carter 3195 (F, NY); Vacca Keys, 28-29
Jan. 1909, Small & Carter 31^5 (F, NY); Big Pine Key, 27 Feb. I9II, Small,
Carter and Small 3568 (NY); Summerland Key, 28 Feb. 1911, Small, Carter
and Small 3591 (NY); Key West, 28-30 Mar. I906, Hitchcock s.n. (F) ;
Boca Grande Key, Marquesas Group, 13 Mar. 1904, Lansing 2271 (F, NY).

Chapman gives little help in typifying this species since his
Flora gives no collector and only "Sandy coast. South Florida" as the
locality. There are a number of collections extant which come from such
an area. Curtiss distributed (#2479) a plant labelled "E. garberi, Chapm.
n.sp. Sandy Field at Cape Sable, Florida". The date of distribution of

196.

•;■' ■I.i.. <.:.rl,i,i,

';;:^:'l'''"'i"j'»wii"fNi|infjflnp(pii^

^ ^ ^^-^^IF

8| » I c

I!

Uiiilihljhll

Plate kk. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small

197.

JHI|HII|IIM|llll|llll|llll|llll|IMI|IIM|llimilllllH|WljHHIHH|ini|IIM|ini{IMIIIIIIii;;lL

■^i| l«| I 31 Nl l»| l>| I7I ' H ^ 4| I i|d ' ill
iS^UJilillllSlllliltllllillllilillUlilJSn.,

12 1 n II

^ .'^ ■ ^A^

Plate i+5. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small

Holotype- C. adicioides Small, E. of Bahia Honda, Monroe County, Florida,
20 Jan. 1920, Simpson U6i+ (NY),

. i: cl.^etc t,i*... very 3!-.ort Ir.terr.odes: Inavesnucerous; Hades
- ;•: or elliptic, nalnly 3-6 n.-i. lone, acute or acutish, entire, hir-
"", r.ll.-htlv lne<]ullaceral, short-pstloled: Involucres approxlr^te
■ : rar.cMets, about 1 rea. hl^h, hirsute, the lobes deltoid; n-''*'"^-
" t 0.5 nm. wide, the appendacen very nario-.i nndi rrecular : capsuleo
"- . ~lde, hircute: seeds ovoid, 1 mt, lone, rediiish, t!^^art!>ier. obt-ise,
tto races scarcely wrinkled.

In plnelands, southern peninsular Florida.

>•

6 i^^^y^^ -r/iy/T-^ t^<

PLANTS OF SUBTROPICAL FLORID

MMt^ bu. Cv

Plate it6. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small

Holotype: _C. brachypoda Small, Miami, Dade County, Florida, 4-7 Apr
1898, Pollard & Collins 2kk (NY).

199.

'.L\?

if|iin|iiiniiiipli|(i"|iiii|Hii|iiijiH^"iPfiTi"|"¥"|^

rrT^r.iii.i.i^.u.i.i.i.i.iVi.iii.ui.i,

Plate 't?. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small

intvoe- C mosieri Small, pinelands about Brogdon Hammock, Dade
county^ FloridTTirJune 1915. Small & Mosier 6347 (NY).

200.

this number has not yet been established, but he is known to have
collected Cape Sable before 1880 (Rodgers, IS^). There is a sheet
from Herb. Engelmann, now in MO, which may well be from this same
collection although labelled only "Florida, A. H. Curtiss". This
sheet is named _E. qarber i Engelm. ap Chapm. and dated I88O.

There are sheets in F and MO (ex herb. Engelmann) collected on
Cape Sable by Garber in October I878 (misread by Millspaugh as I888),
and also an undated sheet by Garber from "Coast of South Florida" which
was bought by MO as part of Chapman's herbarium. The name on this is
_E. qarber i Engelm.. A comment on Garber's Cape Sable sheet "nowhere
else seen" might indicate that these three were part of one collection.

The presence of these plants in their private herbaria seems to
indicate that both Chapman and Engelmann were familiar with this species,
and that Chapman was given the name in correspondence probably on the
basis of the Garber material. In any case, since Garber died in I88I
his specimen was probably in Chapman's hands before this time and during
the preparation of the supplementary list that constitutes the changes
in second edition of the Flora. The description in the Flora was probably
based at least in part on this specimen and it is possible that only the
name should be credited to Engelmann. It is for this reason that author-
ship will be cited in the form used. The same argument governs the
choice of type specimen. Chapman is assumed to have used the specimen
in his herbarium in writing the description, making this the obvious choice
as lectotype. The specimen of Garber's in the possession of Engelmann is
an isotype on the assumption that it is part of the same collection. The
specimens distributed by Curtiss may never have been available to Chapman,
and cannot be allowed any status as types.

201.

Chamaesyce qarberi, as here delimited, shows a wide range of
pubescence and branching characters. Examination of herbarium material
and observations in the field do not support Small's separation of the
group into three or four species, although his type specimens represent
rather distinctive portions of the spectrum of variation. The name
_E. qarberi was applied by Engelmann and Chapman to a plant from the coast
of south Florida, and specimens that show the same characters have been
collected at intervals from various dry areas behind the beach on Cape
Sable at the southwestern extremity of the Everglades. The branching of
these plants rarely goes beyond the production of laterals from the main
branches, with the cyathia developed at the upper nodes of these, in
contrast with specimens from other parts of south Florida where the
laterals themselves often carry congested sideshoots on which most of
the cyathia form.

Pubescence on the plants from Cape Sable is of long, somewhat
velutinous hairs, while in the plants recognized as _C, adicioides by
Small the pubescence is short and more appressed. Small's C. brachypoda
and _C. mosieri have straight pubescence even longer, stiffer, and more
erect than that of the Cape Sable plants. This type of pubescence is
almost confined to collections from the pinelands immediately south of
Miami, but is occasionally found in plants from the Lower Keys. The
close-appressed pubescence occurs on plants from all parts of the Keys
and from pinelands south of Miami, together with plants whose pubescence
falls between the three types described. Small's C. brachypoda and C.
mosieri sometimes have a tendency towards a wiriness of stem, particularly
when sprouting back after a fire, but plants occur which show grades of

202.

Cy \«n

Map 39. Distribution of _C. garberi

Map ^0. Distribution of _C. turpini

203.

heaviness of stem between this and the stouter, less brittle type
which is more common on the Keys.

The plants do not root at the node, but instances were noted
on Cape Sable of stems acting as runners with the buried tips rooting
to produce new plants. This has not been observed in any other species
treated in this study.

The type specimen of _C. adicioides appears to be a s i ngl e- stemmed,
erect plant, but closer examination suggests that it may simply be a
very old specimen of a decumbent plant, from which any other branches
which were present have been lost.

36. CHAMAESYCE MULTINODIS (Urb.) Mi lisp.. Field Mus. Bot. 2:
410. 1916. Euphorbia mul tinodis Urban, Symb. Ant., 1: 34l. 1899. Type:
prope Vieux-Fort, Guadeloupe, 1895, Duss 27^0 (Herb. Mus. Bot., Berlin,
If extant; Isotypes: GH 1, NY i).

Perennial; prostrate, forming mats to 15 cm. diameter. Stem to
1 mm. diameter, internodes to 1 cm. long; branched throughout length or
mostly from upper part, not rooting at nodes, glabrous on lower surface,
tomentose or short-pilose above, green or brown. Leaves membranous or of
somewhat heavier texture; blades broadly ovate to orbicular 3-5 x 3-5 mm.,
base oblique, subcordate , margin obscurely serrate in upper half, apex
blunt or emarginate, scattered long white hairs on both surfaces, green;
petiole 0.5 - 1 mm. long; stipules joined, short 3 to 5-cleft, often
longer on lower surface of stem and merely toothed, ciliate on surface
toward stem. Cyathia solitary at upper nodes; peduncle to 0.5 mm. long,
involucre campanulate, to 1 mm. diameter at mouth, lobes barely exceeding
glands, deltoid, strigose on outside, ciliate within, glands transversely

20^+,

jiijnM|iiii|nn|im[MjM[im|iiti|iiiiliin|im|iiii|iiiip

•Vi^ulJSil.hliilaJWa^^

Symb. hntUI. / J4i. t9^~j

Plate ^8. CHAMAESYCE MULTINODIS (Urb.) Millsp.

Isotype: E. multinodis Urb., prope Vieux-Fort, Guadeloupe, 1895,
Duss 27^0 (NY).

205.

elliptic, 0.3 mm. long, dark and fleshy, appendages a crenulate rim
or as wide as gland, light-colored, fifth gland linear, short, sinus
shallow. Staminate flowers 5-16 per cyathium, androphore glabrous, 0.7
mm, long at maturity. Gynophore pilose, to 1 mm. long at maturity,
exserted, nutant; calyx a triangular pad of tissue, to 0,6 mm, broad;
ovary pilose; styles upright or spreading, 0.3 mm, long, joined at base,
bifid for half length. Capsule sparsely pilose, ovoid, 1,6 mm, long,
1.8-2 mm. wide just below equator, broadly three-lobed, angles rounded.
Seed oblong-ovoid, 1.1 mm, long, 0.6 mm. wide below equator, strongly
four-angled, angles subequal, dorsal more acute than ventral, faces al-
most flat or slightly sunken, obscurely wrinkled, chestnut brown.

Known only from type locality on Guadeloupe, by type specimens,
and by a collection from IJ (Vieux Fort, Basse-Terre, Guadeloupe, 30
Nov. 1959, Proctor 20204).

The species is distinct from others encountered in the Caribbean,
and has no obvious close affinities with them; but bears a general
resemblance to _C. br i ttoni i , C_. cowe 1 1 i i and _C. centunculoi des. It will
be carried under Urban's name unless it can be shown later that it is
an introduction from some other area.

37. CHAMAESYCE MINUTULA (Boiss,) comb. nov. Euphorbia minutula
Boissier in DC. Prod. 15(2): 1263. 1862, Type: Cuba, 1864-65, Wright
2013 (Herb, Conserv, et Jard, Bot,, Geneva, not seen; Isotype: GH i).
The specimen examined has, in addition to Wright's standard label, a
handwritten annotation "Euphorbia liliputiana. On rocks in river San
Sebastian, Lagunillas, Jurisdiccion Pinar del Rio, Dec. 14" (punctuation
added) .

206,

Known only from collections by Wright (type and #3707) from the
same locality in Pinar del Rio, Cuba.

The glabrous, entire leaves separate these plants from depauperate
specimens of _C. thymi fol ia or _C. adenoptera ssp, gund lachi i , and the stems
pubescent on the upper surface, and the pubescent capsules distinguish
between this and the glabrous _C. centunculoides. The species is accepted
here, with some hesitation, on the basis of Wright's material.

The name _E. 1 i 1 iput iana Wr. has not been validly published, it
appears on the herbarium sheets, and is mentioned in a note by Urban
(Symb. Ant., 1: 3^2. 1899), as a presumed synonym for _E. mi nutula Boiss.,
but has no standing beyond this.

Perennial, ascending to 5 cm. Stems tufted, wiry, 0,3 mm, diameter,
internodes to 5 mm. long; almost unbranched, tomentose on upper surface,
glabrous below, brown. Leaves membranous; blades ovate, 2-3 x 1-2 mm.,
base oblique, rounded, margin entire or obscurely toothed, apex rounded,
midrib sometimes prominent on lower surface, glabrous, green; petiole 0.3
mm. long; stipules joined, deltoid, toothed, 0.3 mm. long on upper nodes;
peduncle 0.2 mm. long, involucre obconical, 0.6 mm, diameter at mouth,
lobes equalling glands, deltoid, strigose on outside, ciliate within,
glands transversely elliptic, 0.2 mm. long, appendages to twice as wide
as gland and longer, pairs perhaps of unequal size, fifth gland linear,
short, sinus deep. Staminate flowers ca. 5 per cyathium, androphores
glabrous, to 0.5 mm. long at maturity. Gynophore glabrous, to 0.5 mm.
long at maturity, scarcely exserted, bent through sinus; calyx obsolete,
ovary strigose; styles spreading, 0.2 mm, long, joined at base, bifid at
apex. Capsule not seen, described by Boissier as "adpresse hirtula". Seed
not seen, "obsolete ruguloso".

207.

38. CHAMAESYCE TURPINll (Boiss.) Mi lisp.. Field Mus. Bot., 2:
412. 1916. Euphorbia turpinii Boissier, Cent. Euph. 12. i860. Type:
in Ins. Santo Domingo, Turpin (Herb. Lenormand, not seen).

_E. qarkeana Boiss. in DC. Prod. 15(2): 38. 1862. Type: In
Santo Domingo, Schomb. 167 (Herb. Bot. Mus., Berlin, if extant). Chamae-
syce qarkeana (Boiss.) Millsp., Field Mus. Bot., 2: 409. 1916.

E_. portoricensis Urban, Symb. Ant., 1: 338. 1899. E^. portoricens is
Urb. var. genuina Urb., loc. cit. 339. Type: in Portorico prope Ponce,
prope Guanica, prope Salinas de Cabo-Rojo, et ad Los Morillos, Sintenis
635, 637, 3^59, 3492b, 4807 (Syntypes: Herb. Mus. Bot., Berlin, if ex-
tant; duplicate coll., Sintenis 637, F !, NY 1; Sintenis 3492b, F !).
_E. portoricensis Urb. var. albescens Urb., loc. cit. 399. Type: in Porto-
rico prope Guanica, Sintenis 3439 (Herb. Mus. Bot., Berlin, if extant;
isotype: F !, GH !). C_. portoricensis (Urb.) Millsp., Field Mus. Bot.
2: 394. 1914, C. albescens (Urb.) Millsp., Field Mus. Bot. 2: 408. 1916.

Chamaesyce aneqadensis Mi 1 Isp. , Field Mus. Bot. 2: 394. 1914.
Type: near settlement, Anegada, Dutch West indies, 19-20 Feb. 1913, N. L.
Britton & W. C. Fishlock 998 (Holotype: F !; isotypes: GH i, MO i, NY !).

Dry sites, often among rocks, at low altitudes, Cuba, Haiti, Puerto
Rico and Anagada (Map 40) .

Representative specimens seen: CUBA. Guantanamo, Oriente province,
24 Sept. 1914, Ekman 2901 (NY). HAITI. Les Gonaives, 24 Dec, 1927, Ekman
H9455 (A); Passe David, May 1900, Buch 392 (iJ); Ennery, Artibonite, 13
Jan. 1926, Leonard 88O6 (NY). PUERTO RICO. Salinas de Cabo Rojo, 27 Feb.
1915, Britton, Cowell and Brown 4703 (F, GH, MO, NY); Los Morrillos,
Cabo Rojo, 14 Feb. 1885, Sintenis 636 (A, F, GH, K, MO, NY); Guanica,

208,

5-18 Mar. I915, Britton, Cowell and Brown kS]k (F, NY); Guanica, 19
Sept. 1913, Stevens and Hess 303^ (F, NY); Guanica State Forest, 23
June 1962, Alain 9138 (IJ); Mona Island, 20-26 Feb. 1914, Britton,
Cowell and Hess 1844 (F, NY); Punta Aguila, 27 Feb. I9I5, Britton,
Coweil and Brown 4692 (F, NY).

Perennial; erect or sprawling to 1 dm. Stems several to many
from heavy rootstock, to 1 mm. diameter, internodes to 1 cm. long;
branching from base and heavily in upper part of stem, sparsely or
densely short-pilose on all surfaces, sometimes canescent at tips, brov;n.
Leaves somewhat chartaceous or thickened; blades long-ovate or lanceolate,
2-6 X I-3 mm., base inequilateral, cordate to obtuse, margin entire,
somewhat recurved, apex obtuse, midrib sometimes prominent on lower
surface, both surfaces pilose or tomentose, sometimes sparsely so, some-
times almost woolly-pubescent, green, often suffused red or gray; petiole
0.5 - 1 mm. long; stipules separate or joined at base, to 0.3 mm. long,
toothed. Cyathia solitary at upper nodes; peduncle to 0.5 mm. long,
involucre obconical, to 0.6 mm. diameter at mouth, lobes scarcely exceed-
ing glands, deltoid, toothed, glabrous or sparsely pilose on outside,
ciliate within, glands transversely elliptic, 0.1 mm. long, appendages
obsolete to as wide as gland, fifth gland linear or obsolete, sinus
shallow. Staminate flowers 5-12 per cyathium, androphores glabrous, to
0.8 mm. at maturity. Gynophore glabrous, to 1 mm. at maturity, exserted,
nutant; calyx a triangular pad of tissue; ovary strigose; styles spread-
ing, 0.3 mm. long, minutely joined at base, bifid for 1/2 - 2/3 length.
Capsule broadly ovoid, base truncate, 1.2 mm. long, 1.8 mm. wide just
above base, broadly three-lobed, angles obtuse. Seed ovoid, 0.8 - 1 mm.

209.

1
h

iill|llil|lllllllll{llll|llli|ini|illl|ilii|lMl|illl|llll|lill|liii{iHi|nii|iiii|iiiiMi>iiiiii|iii|-<.f.i .,,. ,|. {;,ii<;|iH
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Plate U9. CHAMAESYCE TURPINI I (Boiss.) Mi lisp.

Duplicate Coll. of Syntype: E. portor i cens i s Urban. Salinas de Cabo-Rojo,
Porto Rico, 17 Feb. 1885, Sintenis 637 (NY).

210.

long, 0.5 mm. wide below equator, strongly four-angled, dorsal angles
more prominent than others, faces flat or convex, sometimes slightly
sunken, obscurely ridged, brown.

The variation in degree of pubescence of this species was
recognized by Urban in setting up the two varieties which he published.
The range of material examined in the present study showed no point
at which to make such a separation consistently, but confirmed that a
wide range of pubescence states exists. No authentic material of
_C. turp i n i i was seen, but specimens given this name by Urban appear to
fall within the variation of his _E. portor i cens i s. There is an overall
similarity between the plants given these names, and it seemed more
reasonable to combine them as one species than to formulate diagnostic
characters which would separate only the extremes of the range.

39. CHAMAESYCE PAREDONENSIS Mi lisp.. Field Mus. Bot. 2: 393. 191^.
Type: Cayo Paredon Grande, Cuba, 25 Oct. 1909, J. A. Shafer 2763 (Holo-
type: NY i, Isotype: F i).

Perennial; prostrate forming dense mat to 2 dm. Stems wiry, many
from crown of root, to 1 mm. diameter, internodes to 1 cm, long; branching
throughout length, not rooting at nodes, tomentose, brown. Leaves mem-
branous or slightly coriaceous; blades elliptic to obovate, 2-4 x 1.5 - 2.5
mm., base strongly inequilateral, rounded to cuneate, margin coarsely
serrate, somewhat revolute, apex blunt, tomentose, green or reddish;
petiole to 0.5 mm,; stipules joined at least at base, broadly triangular,
margin toothed or lacerate, inner surface densely ciliate. Cyathia soli-
tary at uppermost nodes; peduncle to 0.3 mm. long, involucre obconical,
to 0.8 mm. diameter at mouth, lobes triangular, equalling glands, sparsely

211

pilose outside, densely ciliate within, glands transversely elliptic,
0.45 X 0.1 mm., dark brown, cupped, appendages longer than glands and
to twice as wide, white, margin entire or somewhat crenate, fifth gland
tiny or obsolete, sinus shallow. Staminate flowers rarely more than 8
per cyathium, sometimes absent, androphores glabrous, to 0.5 mm. long
at maturity. Gynophores glabrous, to 1.5 mm. long at maturity, exserted,
upright; calyx a triangular pad of tissue; ovary densely pilose; styles
spreading, to 0.3 mni. long, joined at base, bifid for more than half
length, somewhat clavate. Capsule sparsely pubescent, glabrate, ovoid,
1.3 mm. long, 1.6 mm, wide just above base, three-lobed, angles acute.
Seed not seen, described by Millspaugh as "seed pink, farinose, ovoid-
triangular, 1.2 X 0.8 mm., acute, the dorsal angle very sharp, the facets
smooth, the two ventral about equal to one dorsal".

Chamaesyce helwigii from Haiti, and this species are very similar,
and, to judge by Millspaugh's description, share a very distinctive
type of seed. The type specimens differ in several respects - stems
of _C. helwi qi i fractionally stouter than those of _C. paredonens i s , leaves
of the former ap i cu late- toothed near the apex and sparsely long-pilose
while the latter has leaves with serrate edges which are sparsely tomen-
tose, ciliate stipules are not as evident in _C. helwiq i i as in _C. paredon-
ens is, and the appendages of the glands are less prominent. Only the
type collections have been seen, and a wider range of material might
indicate that these two taxa would be better considered one species, but,
until further evidence Is available, they will be maintained under the
separate names used here (Map 33).

212.

^7^.3

Plate 50. CHAMAESYCE PAREDONENSIS Mi lisp.

Holotype: C. paredonensis Mi lisp., Cayo Paredon Grande, Camaquev fuh;,
25 Oct. 1909, Shafer 2763 (NY). ^ ^' ^'

213.

40. CHAMAESYCE PINETORUM Small, Bull. N.Y. Bot. Card., 3:
429. 1905. Type: Homestead trail between Cutler and Camp Longvlew,
Dade County, Florida, 9-12 Nov. 1903, J. K. Small 6- J. J. Carter 836
(Holotype: NY i).

Perennial from root stock to 5 mm. diameter; erect or decumbent
to 10 cm. Stems wiry, to 0.5 mm. diameter, internodes to 2 cm. long;
branching sparse throughout, vi 1 lous-hi rsute, tips canescent, dark
brown. Leaves membranous or slightly thickened; blades reniform to
ovate 2-5 X 2-k mm., base inequilateral, cordate to rounded, margin
entire, somewhat revolute, apex obtuse or rounded, vi 1 lous-hi rsute,
gray-green above, suffused red-brown below; petioles 1 mm. long; stipules
distinct, to 0.3 mm., deltoid, sometimes deep-cleft, ciliate, light-
colored. Cyathia solitary at upper nodes; peduncle to 1,5 mm., involucre
campanulate to 1.3 mm. diameter at mouth, lobes triangular equalling
glands, sparsely pilose on outside, densely hairy within, glands sub-
circular, O.k by 0.3 mm., appendages a rim to slightly wider than glands,
fifth gland triangular, short, sinus broad and shallow. Staminate flowers
8-14 per cyathium, androphores glabrous, to 0.5 mm. long at maturity.
Gynophore sparsely pilose, to 1.5 mm. at maturity, exserted, upright;
calyx a triangular pad of tissue; ovary strigose; styles spreading, to
0.4 mm. long, bifid for 1/2 length, minutely joined at base, obscurely
clavate at tips. Capsule tomentose, reniform, to 1,5 mm. long, 2 mm,
wide just above base, sharply three-lobed, angles acute. Seed ovoid,
0,8 mm. long, 0.5 - 0.6 mm. wide above base, strongly angled, ventral
angle usually obscure, faces convex, obscurely wrinkled, red-brown.

Found only in the pinelands on oolitic limestone of Dade County,
Florida (Map 4l).

21^+.

UAP
EVERGLADE KEYS

I I 12 1.1 M 1ft

.III M W ViillK l»iT»Ml\l. I.»li|.l \ I '■'^Z,

Plate 51. CHAMAESYCE PINETORUM Small

Holotype: _C. pi netorum Small, between Cutler and Longview Camp, Florida
9-12 Nov. 1903, Small & Carter 836 (NY).

215.

Map 41. Distribution of _C. pi netorum

Map k2. Distribution of _C. del toidea ssp. serpyl 1 urn

216,

Representative specimens seen: FLORIDA. Dade: Coral Gables,
25 June i960, Webster S- Williams 1015^ (DUKE, IJ); Everglades National
Park on FLA 27, 11 Aug. I963, Burch I76 (FLAS) ; Florida City, 29 Nov.
1963, Burch 223 (FLAS); Hammer Key, Everglades, 12 May I9I8, Small
8605 (NY, US); Homestead, I8 Kay 195^, Killip 44241 (US); Long Key,
Everglades, 18-26 Jan. 1909, Small & Carter 2930 (NY); near Long Prairie,
24 Mar. 1904, Britton 193 (NY); near Murden Hammock, 22 June 1915, Small,
Hosier S- Small 6451 (FLAS, NY); pinelands about Sykes Hammock, 2 July
1915, Small, Hosier S- Small 676 1 (FLAS, NY).

The strongly decumbent or ascending habit of this species
together with its overall vi 1 lous-pubescence, distinguish it from the
prostrate and glabrous or tomentose _C. del toidea. In most other respects
the species are very close, but maintain their integrity although sym-
patric over the whole range of _C. p i netorum.

Some collections of _C. mos ier i (here placed under _C. garber i ) ,
perhaps even the type sheet, should be referred to this _C. p inetorum.

41. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small, Fl. SE. US.,
710. 1903.

Perennial with swollen rootstock to 1.5 cm. diameter, 4 cm. long;
prostrate forming dense mats to 3 dm. diameter. Stems many from crown
of root, wiry, 0.3 - 0.6 mm. thick, internodes to 1 cm. long; sparingly
or much branched throughout length, not rooting at nodes, glabrous to
sparsely or densely tomentose, light or dark brown. Leaves membranous,
somewhat thickened; blades reniform, orbicular or ovate, 2-4 x 1-4 mm.,
base inequilateral, cordate or subcordate, margins entire or rarely
isolated leaves deeply and coarsely serrate, somewhat revolute or plane.

217.

apex rounded to obtuse, glabrous to sparsely or densely tomentose,
young leaves and stem tip sometimes canescent, yellowish-green or
gray; petiole 0.5 - 1 mm. long; stipules joined 0,2 - 0.3 mm, long,
subentire or several times parted almost to base, tine linear segments
sometimes dark-colored. Cyathia solitary at tips of branches; peduncles
1 - 1.5 mm. long, involucre broadly campanulate, 1.5 mm. diameter, lobes
triangular, equalling or slightly exceeding glands, pilose on outside,
sometimes sparsely so or glabrous, densely hairy within, glands fleshy,
yellow-green or occasionally drying black, transversely elliptic, 0.4 mm.
long, 0,3 mm. wide, appendages obsolete or a rim to gland, fifth gland
linear, shorter than lobes, sinus up to 1/3 depth of cyathium. Staminate
flowers 8-14 per cyathium, androphores glabrous, to 1 mm. long at maturity.
Gynophore glabrous, 1 mm. long at maturity, exserted, nutant; calyx a
subcircular pad of tissue; ovary sparsely to densely strigose; styles up-
right or spreading, 0.4 mm. long, joined at base, bifid for 1/3 length,
somewhat flattened at tip; capsule pilose, glabrate, broadly deltoid or
reniform, 1.2 mm. long, 1.5 mm, wide just above base, sharply and narrowly
three-angled. Seed oblong-ovoid, 0.8 - 1 mm. long, 0.4 - 0.6 mm. wide
below base, three angles prominent, obtuse, ventral angle rounded, faces
convex, plump, obscurely wrinkled, brown or deep red-brown.

Key to subspecies of _C. del toidea
1. Plants forming mats, but becoming diffuse with age; leaves much

longer than wide a, ssp, serpyl lum.

1, Plants maintaining tight mat form with age; leaves about as long

as wide ^- ^^P* del toidea.

218.

4la. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small ssp. SERPYLLUM
(Small) comb, no v. Chamaesyce serpyllum Small, Fl. Florida Keys, 8l. 1913.
Type: Big Pine Key, Monroe County, Florida, 17 Nov. 1912, J. K. Small 3811
(Holotype: NY '.) .

Found only in pinelands on oolitic limestone on Big Pine Key,
Monroe County Florida (Map 42).

Representative specimens seen: FLORIDA. Monroe: Pinelands, Big
Pine Key, 17 Dec. 1913, Small & Small 5027 (DUKE, FLAS, NY); PInewoods,
Big Pine Key, 30 Jan. -4 Feb. 1940, Martin I368 (DUKE, NY); near Watson
Hammock, Big Pine Key, 1 Sept. 1954, Thorne 15011 (IJ); Big Pine Key,
10 Dec. 1921, Small, Small & Matthews IOI96 (NY); Big Pine Key, 4 Sept.
1957, Brass 28978 (FLAS); bridge over Pine Channel, Big Pine Key, 10 Aug.
1963, Burch 139 (FLAS).

The diffuse habit distinguishes this from the other subspecies,
which also has much broader leaves on young parts of the stems. The
seed of this subspecies is usually somewhat plumper, and sometimes longer,
than that of ssp. del to idea.

4lb. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small ssp. DELTOIDEA

Key to varieties of ssp. del to idea
1. Stems and leaves glabrous or with a few tiny hairs on underside of

'eaves 1^ var. deltoidea.

1. All parts of plant tomentose, often densely so, and young shoots

canescent II. var. adhaerens.

4lb. 1. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small ssp. DELTOIDEA
var. DELTOIDEA Euphorbia deltoidea Engelm. ex Chapm., Fl. Southern U.S.
ed. 2: 647. I883. Type: Biscayne Bay, Florida, June, Curtiss 2474

219.

.'? 4 •^|X|"t;i'"'i"^""iT"'"r«; ^ '

i lL!lhli!J3iJ:il!JuJW.i.i::S:... _ ''■

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

Holotype: £. serpyl 1 um Small, Big Pine Key, Monroe County, Florida,
17 Nov. 1912, Smal 1 381 1 (NY) .

220,

(Lectotype: MO i; Isotypes: F '. , NY !). _C. del to idea (Engelm. ex Chapm.)
Smal 1 , Fl. SE. US., 710. 1903.

Locally frequent on oolitic limestone of pinelands and Everglades
Keys of Dade County, Florida (Map 43).

Representative specimens seen: FLORIDA. Dade: Pinelands south
of the Miami River, 20 Nov. 1912, Small 3850 (NY); rocky pine woods near
Miami, 11 July 1895, Curtiss 5468 (F, FLAS, GH, MO, NY); Miami, 13 May
1904, Tracy 9114 (F, MO, NY); Buena Vista, 26 Dec. 1929, Moldenke 291
(MO, NY); Miami, 28 Oct. -28 Nov. 1903, Small £■ Carter 520 (F, NY);
Perrine, 23 Mar. 1904, Britton 145 (F, NY); between Cocoanut Grove and
Cutler, 31 Oct. -4 Nov. 1903, Small £■ Carter 755 (F, NY); N.E. of Perrine,
7 June 1964, Burch 376 (FLAS).

Often found growing with the next variety and resembling it
closely in all features except its pubescence and rather more compact
growth habit,

41b. 11. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small ssp.
DELTOIDEA var. ADHAERENS (Small) comb. nov. Chamaesyce adhaerens Small,
Torreya 27: 104. 1927. Type: Pinelands between Peter's Prairie and
Homestead, Florida, 10 Nov. 190&, J. K. Small & J, J. Carter 2531 (Holo-
type: NY i).

Locally frequent on oolitic limestone of pinelands and Everglades
Keys of Dade County, Florida (Map 44).

Representative specimens seen: FLORIDA. Dade: 2 miles south of
Florida City, 30 Aug. 1954, Thorne 14868 (IJ); north of Goulds, 10 Aug.
1963, Burch l44a (FLAS); Homestead, 22 Nov. 1964, Godfrey 65587 (FLAS,
FSU); Naranja, 23 June I96O, Webster £■ Williams 10107 (PUR); between

221

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plate 53. CHAMAESYCE DELTOIDEA (Engelm, ex Chapm.) Small
ssp. DELTOIDEA var. DELTOIDEA

Isotype- E deltoidea Engelm. ex Chapm., Bay Biscayne, Florida,
undated'. CurTiTTl^--'-- (NY).

222,

EVERCUCE Y' ^L

■ -~- >c^ a^

Plate 5h. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm ) Small
ssp. DELTOIDEA van. ADHAERENS (Small) ined.

Holotype: C. adhaerens Small, Pinelands between Peter's Pralr;« ^
Homestead, Florida, 10 Nov. I9O6, Small s Carter 253I (NY) '"'

Map 43. Distribution of _C. del to idea ssp. del toidea var. deltoide

Map kk. Distribution of C. del toidea ssp. deltoidea var. adhaerens

224,

Peter's Prairie and Homestead, 10 Nov. 1906, Small & Carter 2530 (NY);
near Shields Hammock, 10 July 1915, Small, Hosier 5- Small 6948 (DUKE, NY).
Between Homestead and Naranja, 29 Nov, I963, Burch 226 (FLAS).

Tine type specimens of tlnese two varieties represent the two ex-
tremes of pubescence, and could stand convincingly as the types of two
distinct species. Examination of a long series of collections, however,
has shown a range of intermediates, and it is more consistent with obser-
vations of pubescence difference within other species of the genus to
consider the presence or absence of hairs as no more than a varietal
character! sti c.

Chapman refers only to Curtiss as the collector of this species,
but his collection (#2474) has been generally accepted as the type. The
sheet in Missouri Botanical Garden is designated lectotype because it
is from Engelmann's herbarium, and because no other sheet seen is in
better condition or seems to have any stronger claim to be selected.

Merrill (1948) discusses the pagination and dates of publication
of Chapman's Flora. Author citation is made in this form since there
is no reason to suppose that Engelmann supplied more than the name to
Chapman.

This species has a strong resemblance to _C. turp Ini I , C_. paredonen-
sis, and _C. helwigl i of the Greater Antilles, but differs in having fleshy
glands with appendages much reduced, and In the del toi d-reni form capsule
with three narrow lobes.

42. CHAMAESYCE TUMISTYLA sp. nov. Type: aviation field north
of city. Mole St. Nicholas, Haiti, 13-19 Feb. 1929, E. C. S- G. M. Leonard
13146 (Holotype: NY i; isotype: GH I),

225.

Annual or perennating; prostrate, forming mats to 2 dm. diameter.
Stem to 0.5 mm. diameter, internodes to 5 mm. long; branched freely in
upper part, rooting at nodes, nodes thickened, sparsely strlgose on
upper surface, glabrous beneath, light-colored. Leaves membranous,
somewhat thickened; blades ovate-orbicular, 2-3 x 2-3 mm., base oblique,
rounded or obtuse, margin entire or somewhat erose in drying, apex
rounded, glabrous, green; petiole 0.3 mm. long; stipules joined at base,
sheathing parted into linear segments 0.1 mm. long. Cyathia solitary
at upper nodes; sessile or minutely peduncled, broadly campanulate, to
0.6 mm. diameter at mouth, lobes scarcely equalling glands, deltoid,
glabrous on outside, sparsely hairy within, glands transversely elliptic,
fleshy, 0.3 mm. long, appendages obsolete, fifth gland short, deltoid,
sinus shallow. Staminate flowers 5-12 per cyathium, androphores glab-
rous, to 0.4 mm. long at maturity. Gynophore glabrous, to 1.2 mm. long
at maturity, exserted, upright; calyx obsolete or a rim of tissue; ovary
glabrous; styles upright or spreading, 0.3 mm. long, joined at base,
swollen at apex, undivided. Capsule glabrous, broadly ovoid, 0.6 mm.
long, 0.7 mm, wide below equator, broadly three-lobed, angles rounded.
Seed ovoid, 0.4 mm. long, 0.2 mm. wide below equator, four-angled, ventral
angle sometimes obscure, others rounded, faces convex, plump, smooth,
dark tan.

Known only from the type collection, Haiti.

There are marked differences between this tiny plant and any other
species examined. The prostrate habit with stems rooting at the enlarged
nodes is reminiscent of C. serpens , but the stems are tomentose on one
surface, and are threadlike and much more slender than those of that

226.

glabrous species. The stipules differ, too, in being somewhat sheathing
and yellow, rather than simply fused in pairs and white.

The subsessile cyathium has not been seen elsev^here, but the
most distinctive feature is the undivided style, sv^ollen for the upper
thi rd of i ts length.

This is considered to be a distinct species on the basis of these
characters, and will be published as such under an appropriate name.
The epithet "tumistyla" (which is used here for purposes of identification
within the work, and not in any sense as an attempt at publication) refers
to the enlarged apical portion of the undivided style and stigma.

43. CHAMAESYCE LEONARDll sp. nov. Type: bluff east of Bord de
Mer, vicinity of Jean Rabel , Haiti, 27 Jan. -9 Feb. 1929, E. C. S- G. M.
Leonard 12888 (Holotype: NY i; isotypes: GH !, K i).

Annual; prostrate, fruiting when stems 2-3 cm. long. Stem to
0.5 mm. diameter at base, internodes to 5 mm.; branches reduced to
groups of cyathia, not rooting at nodes, tomentose on all surfaces,
greenish-yellow suffused purple. Leaves membranous or of a heavier tex-
ture; blades ovate-elliptic, 5-8 x 2.5-^, base oblique, subcordate or
rounded, margin obscurely serrated, thickened, apex obtuse, midrib
prominent on lower surface, both surfaces strigose or sparsely tomen-
tose, upper surface glabrate, lower surface minutely papillose, green
suffused purple; petiole ca. 1 mm. long; stipules joined at base, 0.3 mm.
long, deeply bifid, strigose. Cyathia in leafless clusters of 2-5 at
nodes; peduncle 0.3 mm. long, involucre obconical, to 0,6 mm. diameter
at mouth, lobes equalling glands, deltoid, strigose on outside, densely
ciliate within, glands transversely elliptic, 0.05 mm. long, somev/hat

227.

stipitate, appendages obsolete, fifth gland short, deltoid, sinus broad,
shallow. Stamlnate flowers 3-10 per cyathia, androphores glabrous, to
0.6 mm, long at maturity. Gynophore strigose, to 1 mm, long at maturity,
exserted, upright or nutant; calyx a triangular pad of tissue; ovary
densely whi te-str igose; styles spreading, 0.4 mm. long, joined at base,
bifid for half to two-thirds length. Capsule densely whi te-strl gose,
broadly ovoid, 1 mm. long, 1.2 mm. wide at equator, broadly three-lobed,
angles rounded. Seed ovoid, 0.7 mm. long, 0.4 mm, wide, strongly angled,
ventral angle rounded, faces convex, strongly wrinkled, red-brown or
ashen, angles often lighter.

Known only from type collection, Haiti,

The cyathia of this species are borne in a manner not encountered
in other species examined. The laterals near the tip of the stem are
given over entirely to cyathia production, and consist of a group of one
to a few cyathia congested on a stem' a few m i 1 1 imeters long. This
probably represents an extreme reduction of the type shown by _C. maculata
or _C. thymi fol ia, in which cyathia are formed on congested leafy laterals
in the upper part of the stem, rather than a further reduction from the
glomerules of _C. berter iana or _C. hi rta, since close observation shows
that these cyathia alternate on the short stems, while in glomerules
they fall at the nodes of a dichasium.

The white, stri gose-pubescence of the capsules and the prominent
midrib below the purple-flushed leaves are further distinctive features
of the plant, and the wrinkled seed is also unusual. The epithet "leonardii'
in honor of the very active collector of the Haitian flora, will be used
here for identification, and not in an attempt to publish the name.

228.

No close affinities a.-nong species of tiie area are apparent for
this species, but more mature plants may show features which suggest
possible relationships.

DOUBTFUL SPECIES

CHAMAESYCE GYMNADENIA (Urb.) Mi lisp.. Field Mus. Bot. 2:
409. 1916. Euphorbia gymnadenia Urban, Symb. Ant., 5: 390. I9O8.
Type: Cuba, Wright 3706 (Herb. Mus. Bot., Berlin, if extant; Isotypes:

GH :).

The sheet from GH which was seen holds three plants; one is
_C, qund 1 ach i i , and the other two are referrable to _C. mi nutul a as under-
stood here. Urban makes comparisons with these two species in his notes,
but the points of difference are not apparent on the specimens. The
name will probably fall into the synonymy of one of these two species,
but, until the original material has been seen, cannot be assigned to
either one with confidence.

CHAMAESYCE INFLEXA (Urb. & Ekm.) comb. nov. Euphorbia inflexa
Urban & Ekman, Arkiv. Bot., Stockh. 22A (8): 64. 1929. Type: Pineland
toward Peu-de-Temps , Morne Vi ncent ,Gaul thier , Massif de la Selle, Haiti,
3 Mar. 1927, E. L. Ekman H7764 (Holotype: Herb. Naturhistor i ska Riksmuseum,
Stockholm, not seen; Isotype: K i).

No fruiting material of this species has been seen. The vegetative
plants seen at K were suggestive of the Cuban species _C. camaqueyens is,
but cyathial characters must be checked before any decision on the dis-
tinctness of the species can be made.

CHAMAESYCE URBANl! Millsp., Field Mus. Bot. 2: 412. I9I6. Euphorbia
vi 1 losula Urban, Symb. Ant., 1: 340. 1899. Type: in Ste. Domingo, prope
Tamboril, May, Eggers 1944c (Herb. Mus. Bot., Berlin, if extant), non

229.

230.

E. vj] losula Pax, Bot. Jahrb. 19: 118. 189^.

Urban's comment on the species compares it with E. turp i n I i ,
but the name seems to have been used for plants which resemble _C. adenoptera,
perhaps because the description mentions that the glands are not all the
same size. Since no authentic material has been seen the position of this
species cannot be decided with confidence.

PHYLOGENY

The relationships within Euphorbi a sensu lato are far from clear.
No study of the genus at the specific level has been completed since
that of Boissier in 1862, and the treatments of the tribe Euphorb ieae
by Pax (18S0) and Pax and Hoffman (1931) in Die N'aturlichen Pflanzen-
fami 1 ien, and of the family by Scholz (1964) in the Syl labus , are limited
by the format laid down by Engler and Prantl when they originated these
works.

The study by Haber (1925) of the anatomy of the cyathium did not
provide much basis for speculation on the relationship of the divisions
proposed by various authors, although she suggests that there are con-
siderable differences between species in the degree of specialization
shown, A similar pattern of demonstrating the possible utility of
characters without drawing phylogenetic conclusions exists in the chromo-
some studies by Perry (19^3) and the work on electrophoresis of latex by
Moyer (193^). It was disappointing to find that Punt (I962) could give
so little indication of possible phylogenetic paths within the genus
Euphorb la in view of some of the remarkable results obtained from pollen
studies in the rest of the family and many other parts of the plant
ki ngdom.

Wheeler (19^0 has correlated the type of branching described by
Crolzat (1936b, 1937) for the genus Chamaesyce with the suggestions for
the origin of the group first made by Roeper in 1824. He feels that the
development of branches from the crown of the stem after the abortion of

231.

232.

the main axis in Chamaesyce is parallel with the formation of a
pleiochasium in Euphorbi a subgenus Esula Pers. after the main axis has
produced a terminal cyathium. The stems of Chamaesyce would, then, be
homologous to the rays of subgenus Esula which are usually d ichotomous ly
branched. This explanation is in accord with Croizat's description of
the growth of the stems in Chamaesyce by successive production of nodes
terminating in a bud or cyathium, and with extension beyond this by the
development of laterals.

A pleiochasial development similar to that in subgenus Esula is
shown by some members of Euphorbia subgenus Aqaloma. Both groups
have alternate leaves on the main axis but opposite leaves on the rays,
and it is suggested that Chamaesyce has developed from one or the other
by the reduction of this main axis to one or two nodes. This is illustra-
ted in Figures 1 and 2 of the series of diagrams (adapted in part from
Wheeler) in Plate 55.

This origin for Chamaesyce seems entirely reasonable, and implies
that dichotomous branching is a primitive characteristic within the genus.
Dichotomous branching for the full length of the stems (secondary axes)
is rare in the species under consideration, but many show a tendency toward
dichotomy at the tips of the branches and apparently lateral branching
in the lower parts of the stem. This transition is illustrated in Figure
3. The lateral branching comes about by the unequal development of the
lateral buds at each node, and in most cases the partial suppression of
one lateral alternates up the stem. This type of branching Is found in
most of the upright, large-leaved species such as _C. hyssop i fol ia and
_C. hyper i ci fol ia.

233.

The production of cyathia is usually confined to the upper parts
of the stem, and the solitary cyathium is terminal at each node, although
a lateral may extend the axis so that it appears to be lateral. In
some species such as C_. hyssopifol ia. production is concentrated on the
partially suppressed member of the pair of laterals, which develops into
a compound dichasium bearing many cyathia. Condensation of the internodes
of the dichasium, and reduction or loss of leaves gives rise to the
glomerulate grouping of cyathia in _C. hyper i ci fol i a and to the tight
glomerules of C_. hi rta and similar species (Figures k and 5).

in many of the prostrate species, such as _C. maculata, there is
very little tendency to dichotomy in any part of the stem, and there is
apparently a normal alternate lateral branching system. This represents
an extreme suppression of one or other lateral, which then shows the
same suppression in its own branching. Reduction of internodes in these
laterals gives rise to the congested production of cyathia which is often
seen at the upper nodes of these species (Figure 6).

Two paths can thus be traced in the evolution of branching pattern
in the group of species under consideration. Dichotomous branching
appears to be the primitive condition, and, in almost all cases, has given
way to unequal development of the two branches of the dichotomy. In one
series the partially suppressed branch has retained its dichotomous tendency,
and the cyathia are carried in compound dichasia, which, by uniform reduc-
tion of internodes, have given rise to tight glomerules. The second series
Involves a continuation of the suppression of one of the branches of the
dichotomy into the branching of these sideshoots themselves. Reduction of
the internodes of these shoots has given the congested branchlets on which

23^.

Fig. 1

Plate 55. EVOLUTION OF BRANCHING PATTERN IN CHAMAESYCE

235,

most of the cyathia are developed in many species. These are usually
leafy, but further reduction in _C. 1 eonard i i has resulted in the produc-
tion of small groups of cyathia on a leafless axis.

There is no evidence to suggest that reduction in either of these
series has proceeded to the point at which there has been much loss in
the number of cyathia, and in particular it seems safe to assume that
the production of a single terminal cyathium at a node is a primitive
rather than a derived condition.

No other characters in the genus show transitions which are as
easily recognizable as are those in the branching and the position of
cyathia production.

Croizat pointed out, while describing the abortion of the primary
axis which characterizes the genus, that the ability to produce buds at
this aborted apex, or for them to be held back by factors of the environ-
ment, makes the plant very efficient under adverse conditions. Many
species grow very slowly under drought or an unfavorable temperature
regime, but produce heavy growth and abundant seed when conditions im-
prove for even a short period. This must have represented a selective
advantage over plants which had to produce a large basic branch system
before forming cyathia, such as those of Euphorbia subgenus Esula.

An aborted primary axis from which grow many fruiting branches,
homologous to the rays of subgenus Esula, is taken to be the primitive
state in Chamaesyce. Once this character was established, evolution was
probably from a small to a much enlarged root in the unfavorable habitats
in which the appropriate selection pressure was present, and this type
of root, and the associated long life-span, are probably derived characters,

236.

The situation is not clear-cut, however, since in less stringent
environments it is unlikely that the same pattern of development would
have occurred. The maximum seed production under uniformly favorable
conditions would come from steady growth and fruiting, and in this case
a single- or few-stemmed plant might be derived from the basic type
in which many stems grow from the crown of the root. The shrub habit
is probably also derived, and represents another response to environment,
perhaps in species in which the ability to produce multiple stems from
ground level had already been lost.

Pubescence is widespread in the family Euphorb iaceae, often with
complex stellate or branched hairs. it is not common in the groups close
to Chamaesyce, but presumably represents a primitive condition. All
pubescence in the genus is of simple hairs, although most are multicelled,
and in some cases are large and heavily pigmented.

Most characters of the inflorescence are rather stable in the
genus. The perianth of the female flower is reduced to a vestigial
calyx represented by a pad of tissue and in only one species, _C, serpens,
do the three lobes develop to about 0.3 fTini. long, and recurve. it would
be necessary to examine the vascular traces to decide if this is more
than a casual elaboration of tissue.

The number of male flowers varies greatly from species to species,
and Haber (1925) traced a series through Euphorbia sensu lato in which
the numbers in each fascicle in the cyathium were reduced, from the
primitive state with many present, by loss of terminal and lateral flowers
in various combinations. The examples that she gives from Chamaesyce
fall throughout the series.

237.

The same is true of the elaboration or reduction of the bracts
which subtend the male flowers in their inflorescences inside the
cyathium. These bracteoles are fused, large and much divided in most
species of Chamaesyce and this is probably derived from the simple, un-
joined type that Haber suggests as the basic form for the genus Euphorbia.

The fifth gland and the sinus in which it lies seem variable with-
in species but are broadly similar through the genus Chamaesyce. The
fifth gland occasionally develops in the full glandular form in C_, vagin-
ul ata, indicating perhaps, that the primitive condition with five equal
glands is not completely lost from the group.

The glandular appendages are petal loid through most of the group,
and their loss is a derived condition which is found consistently in only
a few species. It is more common for pairs of appendages to develop to
a different extent. This is of very frequent occurrence, and in one set
of species the condition has developed to such a marked degree that the
cyathium is covered at maturity by the greatly elongated pair which flank
the sinus.

These characters, and others in which changes are even less marked,
give little insight into possible phylogenetic paths in the genus. With
the large number of species to consider it is surprising that no grouping
can be made, at the taxonomic level of ser i es for example, within the genus,
The only groups which suggest themselves are those such as the species
with expanded appendages, or those in which all cyathia are carried on
congested laterals, which have no features to give a constant separation
from the remainder of the genus. It is interesting to note that Wheeler
(19^-1), in his study of approximately the same number of species from the

238.

area north of the Caribbean, was equally unwilling to lay down lines,
and Boissier (1862) divided his section An isophyl 1 um on patently arti-
ficial grounds, and as a matter of convenience rather than phylogeny.

The impression gained from this work with the genus in the
Caribbean is that speciation is proceeding at a relatively rapid pace.
The treatment here has been conservative in that many taxa recognized
as species by earlier authors have been reduced to a lower taxonomic
rank. This is due in some cases to the availability of a wider range
of material than other workers used, which has shown their species to
be merely the extremes of a continuous series, but more often to a
different assessment of the degree of interchange of genetic material
still taking place. The potential for complete separation in many of
these cases appears to be high.

Neither the pollen nor the seed is adapted for wide distribution,
and this may account for the high degree of endemism shown by the group.
All parts of the area studied had endemic species, in some cases
restricted to a single island. In many cases it was possible to recog-
nize similar species from the area, but others must relate more closely
to taxa from other parts of the world, and emphasize the difficulties of
phylogenetic speculation based on even as apparently contained an area
as the Caribbean.

BIBLIOGRAPHY

ALAIN, Hermano. 1953. Flora de Cuba, 3. Contr. Ocas. 13, Mus. Hist.
Nat. "Col. de la Salle", Habana,

BOISSIER, E. 1862. Euphorbieae, in de Candolle, Prodromus systematis
natural is regni vegetabilis, 15(2): 3-188. Paris.

BRITTON, N. L. and MILLSPAUGH, C. F. 1920. The Bahama Flora. Pub-
lished by the authors, New York.

BRITTON, N. L. and WILSON, P. 1924. Scientific Survey of Porto Rico
and the Virgin Islands, 5: Botany of Porto Rico and the Virgin
Islands. New York Academy of Sciences, New York.

BROWN, H. E. 1911. Euphorbia, in Thi s t 1 edon-Dyer , Flora of Tropical
Africa, 6(1): 470-603. Reeve & Co., London.

BROWN, R. I8l8. Observations on the natural family of plants called
Compositae. Trans. Linn. Soc. 12: 76-142.

BURCH, D. G. 1965. The application of the Linnaean names to some New
World species of Euphorbia subgenus Chamaesyce. Rhodora,
in press.

BURMAN, J. 1737. Thesaurus Zeylanicus. Amsterdam.

CROIZAT, L. 1936a. On the classification of Euphorbia. I. How important
is the cyathium? Bull. Torr. Bot. Club 63: 525-531.

. 1936b. Chamaesyce. in Degener, Fl. Hawaii, family 190,

Chamaesyce 1 £• 4. Published by the author, Honolulu.
. 1937. Chamaesyce, in Degener, Fl. Hawaii, family 190,

Chamaesyce 2 & 3. Published by the author, Honolulu.

DAVIS, P. H. and HEYWOOD, V. H. I963. Principles of Angiosperm Taxonomy.
Van Nostrand Co., New York.

D'ISNARD, A. T. D. 1720. Mem. Acad. Roy. Sci., Paris.

ERDTMAN, G. 1952. Pollen Morphology and Plant Taxonomy. Chronica
Botanica Co., Waltham, Mass.

FARWELL, 0. A. 1936. Euphorbia pi 1u1 ifera in Michigan. Rhodora 38:
331-332.

239.

2^0.

FAWCETT, W. and RENDLE, A. B. 1920. Flora of Jamaica, 4. British
Museum, London.

GRAY, S. F. 1821. A Natural Arrangement of British Plants, 2.
Baldwin, London.

GUNTHER, R. T. 1959. The Greek Herbal of Dioscorides. Oxford Uni-
versity Press, Oxford.

HABER, J, M. 1925. The anatomy and morphology of the flower of
Euphorbia. Ann. Bot. 39: 657-707.

HAWORTH, A. H. 1812. Synopsis Plantarum Succu lentarum. Taylor,
London.

. 1827. Descriptions of new succulent plants. Philos. Mag.

S- Ann. Philos. n.s. 1: 271-277.

JACQUIN, N. J. 1763. Select. Stirp. Amer. Hist. Vienna.

KLOTZSCH, J. and GARCKE, H. 1859. Linne's Naturliche Pf lanzenklasse
Tricoccae. Abh. Akad. Berlin 1859 (Phys.): I-IO8.

LANJOUW, J. and STAFLEU, F. A. 1964. Index Herbariorum: 1. The
Herbaria of the World, edition 5. International Bureau for
Plant Taxonomy and Nomenclature, Utrecht.

LINNAEUS, C. 1753. Species Plantarum, ed. 1. Stockholm.

. 1754. Genera Plantarum, ed. 5. Stockholm.

MERRILL, E. D. 19^8. Unlisted binomials in Chapman's Flora of the
Southern United States. Castanea 13: 6I-83.

MICHAUX, F. A. 1803. Flora Boreal i-Americana. Paris.

MILLER, P. 1754. The Gardeners Dictionary, abridged ed. k, London.

. 1768. The Gardeners Dictionary, ed. 8. London.

MILLSPAUGH, C. F. 1909. Praenunclae Bahamensis II. Field Mus. Bot. 2:
289-322.

MOVER, L. S. 1934. Species relationships in Euphorbia as shown by the
electrophoresis of latex. Am. J. Bot. 21: 293-313.

PAX, F. 1890. Euphorbiaceae, in Engler £- Prantl, Die Naturlichen
Pf lanzenfami 1 ien, 3(5): 1-119. Leipzig.

. 1924. Die phylogenie der Euphorbiaceae. Bot. Jahrb. 59:

129-176.

2k]

PAX, F. and HOFFMANN, K. 1931. Euphorb iaceae, in Engler & Prantl, Die
Naturlichen Pf lanzenfam i 1 ien, 2 Aufl,, 19c: 11-233, Leipzig.

PERRY, B. A. 19^3. Chromosome number and phylogenetic relationships
in the Euphorbiaceae. Am. J. Bot. 30: 527-5^1.

PERSOON, C. H. 1806. Synopsis Plantarum. Paris.

PUNT, J. 1962. Pollen morphology in the Euphorbiaceae with special
reference to taxonomy. Wentia 7: 1-1 16.

RAFINESdUE, C. S. 1817. Second decade of undescribed American plants.
Amer. Monthly Mag. 2: 119-120.

. 1838. Flora Telluriana, k. Philadelphia.

REICHENBACH, H. G. L. 1841. Repertorium Herbarii s i ve Nomenclator
Genera Plantarum. Dresden,

RODGERS, A. D. 1944, American Botany 1873-1892. Princeton University
Press, Princeton.

ROEPER, J. 1824, Enumeratio Euphorb iarum. Gottingen,

1828, Euphorb ia, in de Candolle, Botanicon Gallicum, 1:

412-417.

SCHOLZ, H, 1964, Euphorbiaceae, in Engler, Syllabus der Pf 1 anzenfami 1 ien ,
II. Dicoty ledoneae, 255-261. Gebruder Borntraeger, Berlin.

SMALL, J, K. 1933. Manual of the Southeastern Flora. Published by the
author. New York.

TORREY, J. 1843. A Flora of the State of New York, Albany, New York,

TOURNEFORT, J. P, de. 1700. 1 ns t i tut iones Rei Herbariae. Paris.

URBAN, I. 1898-1928. Symbolae Antillanae seu Fundamenta Florae Indiae
Occidental i s. Nine volumes. Berlin.

URBAN, 1. and EKMAN, E. L, 1929. Ark. f, Bot,, Stockh. 22A: parts 8-I6,

WHEELER, L, C. 1939. A miscellany of the New World Euphorbiaceae.
Contr. Gray Herb. 127: 48-78.

. 1941. Euphorb ia subgenus Chamaesyce in Canada and the United

States exclusive of southern Florida. Rhodora 43: 97-154, 168-205,
223-286. Reprinted as Contr. Gray Herb. I36.

242.

WHEELER, L. C. 1943. The genera of living Euphorbieae. Am. Mid. Nat.
30: 456-503.

_. i960. Typification of Euphorb ia macul ata. Rhodora 62:

134-141.

WIMAN, J. 1752. Euphorbia ejusque Historia Natural is. Uppsala. Re-
printed in Amoen. Acad. 3: IOO-I3I.

APPENDIX

The following new names and combinations are used in this
work, and will be validated by publication at a later date.

New species

Chamaesyce 1 eonard i i
Chamaesyce tumi styl a

New combi nat ions

Chamaesyce lecheoides (Millsp.) Millsp. var. wi 1 soni i (Millsp.)

Chamaesyce lecheoides (Millsp.) Millsp. var. exumensis (Millsp.)

Chamaesyce porteriana Small var, keyens is (Small)

Chamaesyce porteriana Small var. scopar i a (Small)

Chamaesyce parci flora (Urb.)

Chamaesyce hepatica (Urb. S- Ekm.)

Chamaesyce helwigii (Urb. & Ekm,)

Chamaesyce opthalmica (Pers,)

Chamaesyce adenoptera (Bertol.) Small ssp, pergamena (Small)

Chamaesyce adenoptera (Bertol.) Small ssp. qundlachi i (Urb.)

Chamaesyce minutula (Boiss.)

Chamaesyce deltoidea Small ssp. serpyl 1 um (Small)

Chamaesyce deltoidea Small ssp. deltoidea var. adhaerens (Small)

243,

BIOGRAPHICAL SKETCH

Derek George Burch was born on June 26th, 1933, in Caerphilly,
Glamorgan, Great Britain. He attended school in London, and college
at University College of Wales, Aberystwyth, where he received a
B.Sc. (Hons.) in Agricultural Botany in 195^. He carried out graduate
work at that institution and at the University of Nottingham leading
to an M.Sc. in 1957.

After working for a sugar company in the Dominican Republic,
and in various areas of horticulture in Miami and Montreal, Mr. Burch
was admitted to the Graduate School of the University of Florida in
September, I96I. He has been employed as a teaching and research
assistant and as an Instructor In the Department of Botany, and as
herbarium correspondent for the Agricultural Extension Service while
pursuing his work toward the degree of Doctor of Philosophy.

Mr. Burch is a member of the Association of Applied Biologists,
the Society for Experimental Biology, the Classification Society, and
the Royal Geographical Society.

Mr. Burch is married to the former Nancy Hudgins of Lake Wales,
Florida, and they have no children.

2^.

This dissertation was prepared under the direction of the
chairman of the candidate's supervisory committee and has been approved
by all members of that committee. it was submitted to the Dean of the
Coli.^cje of Agriculture and to the Graduate Council, and was approved as
partial fulfillment of the requirements for the degree of Doctor of
Ph i losophy.

August, 1965

yC)P>>d€a^n, College of Agriculture

Dean, Graduate School

Supervisory Committee:

Cha i rman

^242