A TAXONOMIC REVISION OF THE GENUS
CHAMAESYCE (EUPHORBIACEAE) IN
DEREK GEORGE BURCH
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARIIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
I should like to express my thanks to Dr. Daniel B. Ward
for his advice and helpful criticism in his capacity as Chairman
of the supervisory committee for this study. I should also like to
thank the other members of the committee, Dr. W. 0. Ash and Dr.
D. A. Roberts, for reading and evaluating the manuscript.
I am grateful to the Curators and staff of the herbaria listed
in the introduction for lending their material, including type-
specimens, for use in this study, and my thanks are also due to
Mr. W. T. Steam of the British Museum (Natural History) and Dr.
R. K. Rechinger of the Naturhistoriskmuseum, Vienna, for searching
their collections for additional specimens of interest.
The Department of Botany and the Graduate School of the Uni-
versity of Florida contributed to the expenses of a visit to several
important European herbaria, which added greatly to the coverage of
Finally, my thanks are due to the anonymous assistants for the
final stages of the preparation of the dissertation, and particularly
to my wife Nancy, who rounded out two years of "herbarium-widowhood"
by finding out just how I had been spending my nights during that time.
All photographic work in the dissertation was done by Mr. Jerry Uelsmann,
and I am grateful to him both for the long hours of work that went into
the plates, and for their leavening effect in the mass of text.
TABLE OF CONTENTS
LIST OF MAPS
LIST OF PLATES
II. THE SEGREGATION OF THE GENUS CHAMAESYCE
Key to the species of Chamaesyce in the Caribbean
1. Chamaesyce vaginulata
2. Chamaesyce articulata
3. Chamaesyce buxifolia
4. Chamaesyce lecheoldes
5. Chamaesyce cayensis
6. Chamaesyce porteriana
7. Chamaesyce myrtillifolia
8. Chamaesyce nutans
9. Chamaesyce hypericifolia
TABLE OF CONTENTS, Continued
SYSTEMATIC TREATMENT, Continued Page
10. Chamaesyce hyssopifolia 76.
11. Chamaesyce lasiocarpa 86.
12. Chamaesyce parciflora 88.
13. Chamaesyce brittonii 90.
14. Chamaesyce cowellii 93.
15. Chamaesyce blodqettii 97.
16. Chamaesyce serpens 107.
17. Chamaesyce centunculoides 111.
18. Chamaesyce torralbasii 117.
19. Chamaesyce camaguayensis 121.
20. Chamaesyce cordifolia 124.
21. Chamaesyce polygonifolia 128.
22. Chamaesyce ammannioides 130.
23. Chamaesyce cumulicola 134.
24. Chamaesyce prostrata 138.
25. Chamaesyce mendezii 144.
26. Chamaesyce hepatica 148.
27. Chamaesyce helwiqii 150.
28. Chamaesyce hirta 152.
29. Chamaesyce opthalmica 160.
30. Chamaesyce berteriana 165.
31. Chamaesyce maculata 168.
TABLE OF CONTENTS, Continued
SYSTEMATIC TREATMENT, Continued Page
32. Chamaesyce thymifolia 175.
33. Chamaesyce adenoptera 180.
34. Chamaesyce conferta 190.
35. Chamaesyce garberi 193.
36. Chamaesyce multinodis 203.
37. Chamaesyce minutula 205.
38. Chamaesyce turpinii 207.
39. Chamaesyce paredonensis 210.
40. Chamaesyce pinetorum 213.
41. Chamaesyce deltoidea 216.
42. Chamaesyce tumistyla 224.
43. Chamaesyce leonardii 226.
DOUBTFUL SPECIES 229.
BIOGRAPHICAL SKETCH 244.
LIST OF MAPS
Distribution of C.
Distribution of C.
Distribution of C.
Distribution of C.
Distribution of C.
Distribution of C.
Distribution of C.
lecheoides var. wilsonii
lecheoides var. lecheoides
lecheoides var. exumensis
porteriana var. keyensis
porteriana var. porteriana
porteriana var. scoparia
articulata and C. myrtillifolia
cowel 1 ii
LIST OF MAPS, Continued
of C. cordifolia
of C. polygonifolia
of C. ammannioides
of C. cumulicola
of C. prostrata
of C. mendezii
of C. hirta
of C. opthalmica
of C. berteriana
of C. maculata
of C. helwiqii and C. paredonensis
of C. thymifolia
of C. adenoptera ssp. adenoptera
of C. adenoptera ssp. perqamena
of C. adenoptera ssp. qundlachii
of C. conferta
of C. garberi
of C. turpinii
of C. pinetorum
of C. deltoidea ssp. serpyllum
of C. deltoidea ssp. deltoidea
of C. deltoidea ssp. deltoidea
LIST OF PLATES
Plate 1. CHAMAESYCE VAGINULATA (Griseb.) Millsp. 31.
Plate 2. CHAMAESYCE ARTICULATA (Aubl.) Britton 35.
Plate 3. CHAMAESYCE BUXIFOLIA (Lam.) Small 38.
Plate 4. C. LECHEOIDES (Millsp.) Millsp. var. WILSONII
(Millsp.) ined. 46.
Plate 5. CHAMAESYCE LECHEOIDES (Millsp.) Millsp. var.
Plate 6. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS
(Millsp.) ined. 48.
Plate 7. CHAMAESYCE CAYENSIS (Millsp.) Millsp. 53.
Plate 8. CHAMAESYCE PORTERIANA Small var. KEYENSIS (Small)
Plate 9. CHAMAESYCE PORTERIANA Small var. PORTERIANA 59.
Plate 10. CHAMAESYCE PORTERIANA Small var. SCOPARIA (Small)
Plate 11. CHAMAESYCE MYRTILLIFOLIA (L.) Millsp. 64.
Plate 12. CHAMAESYCE NUTANS (Lag.) Small 67.
Plate 13. CHAMAESYCE HYPERICIFOLIA (L.) Millsp. 73.
Plate 14. CHAMAESYCE HYSSOPIFOLIA (L.) Small 79.
Plate 15. CHAMAESYCE HYSSOPIFOLIA (L.) Small 80.
Plate 16. CHAMAESYCE HYSSOPIFOLIA (L.) Small 81.
Plate 17. CHAMAESYCE BRITTONII (Millsp.) Millsp. 92.
Plate 18. CHAMAESYCE COWELLII Millsp; in Britton 95.
Plate 19. CHAMAESYCE BLODGETTII (Engelm. ex Hitchc.) Small 99.
LIST OF PLATES, Continued
Plate 39. CHAMAESYCE
BLODGETTII (Engelm. ex Hitchc.) Small
BLODGETTII (Engelm. ex Hitchc.) Small
SERPENS (HBK.) Small
CENTUNCULOIDES (HBK.) Millsp.
CENTUNCULOIDES (HBK.) Millsp.
TORRALBASII (Urb.) Millsp.
CORDIFOLIA (Ell.) Small
POLYGONIFOLIA (L.) Small
AMMANNIOIDES (HBK.) Small
PROSTRATA (Ait.) Small
MENDEZII (Boiss.) Millsp.
HEPATICA (Urb. & Ekm.) ined.
HELWIGII (Urb. & Ekm.) ined.
HIRTA (L.) Millsp.
OPTHALMICA (Pers.) ined.
MACULATA (L.) Small
MACULATA (L.) Small
THYMIFOLIA (L.) Millsp.
Plate 40. C. ADENOPTERA (Bertol.) Small var. PERGAMENA
Plate 41. C. ADENOPTERA (Bertol.) Small var. PERGAMENA
LIST OF PLATES, Continued
Plate 42. C. ADENOPTERA (Bertol.) Small var. GUNDLACHII
(Urban) ined. 187.
Plate 43. CHAMAESYCE CONFERTA Small 192.
Plate 44. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 196.
Plate 45. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 197.
Plate 46. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 198.
Plate 47. CHAMAESYCE GARBERI (Engelm. ex Chapm.) Small 199.
Plate 48. CHAMAESYCE MULTINODIS (Urb.) Millsp. 204.
Plate 49. CHAMAESYCE TURPINII (Boiss.) Millsp. 209.
Plate 50. CHAMAESYCE PAREDONENSIS Millsp. 212.
Plate 51. CHAMAESYCE PINETORUM Small 214.
Plate 52. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small
ssp. SERPYLLUM (Small) ined. 219.
Plate 53. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small
ssp. DELTOIDEA var. DELTOIDEA 221.
Plate 54. CHAMAESYCE DELTOIDEA (Engelm. ex Chapm.) Small
ssp. DELTOIDEA var. ADHAERENS (Small) ined. 222.
Plate 55. EVOLUTION OF BRANCHING PATTERN IN CHAMAESYCE 234.
The genus Chamaesyce (Euphorbiaceae) is represented on all
continents, and, while many species are widespread in temperate areas,
the genus reaches its greatest diversity in tropical and subtropical
In 1941 Wheeler revised the group in North America exclusive of
Mexico and southern Florida, treating it as a subgenus of the Linnaean
genus Euphorbia. He found areas with high concentrations of species
centered in Arizona and in Texas, and suggested that a similar concen-
tration might exist in south Florida involving species with a Caribbean
affinity. The most recent treatment of the genus in Florida by Small
(1933) indicates the same thing, since eighteen species confined to
Florida are included in the twenty-nine that he lists for the state.
Many of these are from the pinelands around Miami or on the lower Keys,
areas with very distinctive plant associations, but not generally rich
in endemic species. The plants of south Florida are known to have strong
floristic ties both with Cuba and the Yucatan peninsula, and with the
Bahama Islands, and it was clear that the Florida species could not be
dealt with effectively without considering those of the surrounding area.
The Caribbean as a whole here considered to include southern Florida
as well as the Bahamas and the Greater and Lesser Antilles, but excluding
Trinidad and the islands of the Dutch West Indies off the coast of
Venezuela does, however, make up a contained unit floristically, and
is a convenient area for study. It has the added advantage that no
synthesis of the genus for this area has been attempted since Boissier's
treatment of Euphorbia sensu lato on a worldwide basis in 1862, although
certain regions have been extensively collected since that time, and the
genus revised in these localities.
Regional floras with relevant material include The Bahama Flora
by Britton and Millspaugh (1920), which draws on Millspaugh's strong
interest in Chamaesyce, Fawcett and Rendle's Flora of Jamaica (1920),
Scientific Survey of Porto Rico and the Virgin Islands by Britton and
Wilson (1924), and Flora de Cuba by Alain (1953). There is also material
relating to Hispaniola and to Cuba in Urban's Symbolae Antillanae (1898-
1928), and in the reports of Ekman's collecting trips in Haiti (Urban
and Ekman, 1929).
The consequence of this publication of works covering only small
geographic areas has been the description of the same plant under
different names from different parts of its range. It is fortunate
that much of the type material for these names is still accessible, and
it has proved possible in the course of the study to reduce many of them
to synonymy. Material from twenty-two herbaria has been examined, and
the large number of specimens available of many species has given
valuable indications of the extent of morphological variation to be
expected and taken into account when considering the validity of names
about which there is doubt.
Some 4500 sheets have been studied and additional material, as
well as valuable type specimens, examined during visits to other insti-
tutions. The herbaria from which material has been seen are listed below,
together with the abbreviations (most from Lanjouw and Stafleu, 1964),
by which they will be referred to in the text.
A Arnold Arboretum, Cambridge, Massachusetts
BM British Museum (Natural History), London
BRASS Herbarium of Archbold Biological Station, Lake Placid, Florida
BUS University of Miami, Coral Gables, Florida
DUKE Duke University, Durham, North Carolina
E Royal Botanic Garden, Edinburgh, Great Britain
F Chicago Natural History Museum, Chicago, Illinois
FLAS University of Florida, Gainesville, Florida
FSU Florida State University, Tallahassee, Florida
GH Gray Herbarium of Harvard University, Cambridge, Massachusetts
IJ Institute of Jamaica, Kingston, Jamaica
K Royal Botanic Gardens, Kew, Surrey, Great Britain
LINN Linnaean Society Herbarium, London
MO Missouri Botanical Garden, St. Louis, Missouri
NCU University of North Carolina, Chapel Hill, North Carolina
NSC North Carolina State University, Raleigh, North Carolina
NY New York Botanical Garden, New York
PUR Purdue University, Lafayette, Indiana
RUS Louisiana Technical College, Ruston, Louisiana
UCWI University of the West Indies, St. Andrews, Jamaica
US U. S. National Museum, Washington, D. C.
USF University of South Florida, Tampa, Florida
Many taxonomists have treated Chamaesyce as no more than a
subdivision of the large, diverse genus Euphorbia L. To a certain
extent the choice of which of these generic names to use is an
academic matter, but, insofar as the schemes of biological classifi-
cation are intended to have a natural basis, the collection of species
into a genus should reflect a close degree of common ancestry, and
the thesis of workers who would split Euphorbia into several genera
is that such a division represents this close relationship better than
would the maintenance of the broad genus.
The genus Euphorbia was described by Linnaeus in Genera Plantarum
ed. 5 (1754), and was delimited on the basis of its floral characters.
Reference was made to the earlier names Euphorbium proposed by D'Isnard
in 1720 for much the same group, and to Tithymalus and Tithymaloides,
both of which were used by Tournefort (1700), one of the first botanists
to make such formal definitions of groups rather than simply listing
The "sexual system" of classification followed by Linnaeus
divided plants into Classes according to the number of stamens present
in the flower, with these Classes then divided into Orders, usually
formed by reference to the number of styles. The genus Euphorbia was
placed in the third Order of the eleventh Class, DODECANDRIA TRIGYNIA,
since Linnaeus had found that twelve or more stamens were usually present,
and that there were three styles. The description in Genera Plantarum
stresses the characteristic calyx of the genus, with its segments joined
to form a four-or five-toothed cup and petals set on the margin alter-
nating with the teeth, and also the tricoccate capsule with a single
seed in each locule. Some of the variety to be found in the shape of
the petals and in the form of the stem is also mentioned. Linnaeus was
familiar with some fifty-six species at this time, and his descriptions
in Species Plantarum (1753) indicate that he felt the characteristic
flower to be a strong enough factor to hold together in one genus such
diverse morphological forms as the cactus-like E. mammillaris, the small,
prostrate E. maculata, and the erect E. lathyrus with its umbellate
branches and crescent-shaped petals.
The genus, characterized by the flower type, was accepted by most
authors of the period, although Wheeler (1943) points out that as early
as 1754 Miller had proposed that part of the genus, homogeneous for cer-
tain morphological characters, be split off under another name.
In 1818 Robert Brown, an Edinburgh surgeon and keen botanist,
published the results of anatomical work which showed that the flower
which Linnaeus described was, in fact, an inflorescence. The tricoccate
ovary was actually a terminal female flower with the perianth parts re-
duced to a vestigial calyx, and below this in the inflorescence were
groups of male flowers, each reduced to a single stamen. The cup-like
calyx which Linnaeus described was an involucre of fused bracts, while
the petals were glands and associated appendages set on the upper rim
of the involucre. The name cyathium was given to this type of inflorescence.
This work of Brown weakened the case for maintaining the diverse
forms under one genus because it is inconsistent with usage in the rest
of the plant kingdom, and because Linnaeus would presumably not have
defined a genus on this basis of a shared inflorescence type. The
parallel has been drawn with the situation in the Compositae, which
are defined as a family by the capitate inflorescence and by reduction
in some flower parts, and which would form a genus of some 20,000 species
if these were considered generic rather than family characters. This
obviously represents neither a realistic appraisal of the close degree
of common ancestry which is usually sought for a genus, nor an efficient
means of data retrieval which is one of the tacit aims in setting up
such a system of classification.
In spite of this or perhaps because Brown's results were not
universally accepted at first (Haber, 1925) many of the diverse species
described in the hundred years after the publication of Species Plantarum
were named using Euphorbia in its broad sense, and the stamp of approval
was given to the practice by Boissier in a series of publications cul-
minating in his monumental treatment of the family with J. Muller in
de Candolle's Prodromus (1862). The characters of the inflorescence
were still used to define the genus, but by this time the number of species
known had swollen to the point at which internal subdivision of the genus
was standard practice. Persoon (1806), Rafinesque (1817), and Reichen-
bach (1841) had all set up subgenera, while Haworth (1827) and Roeper
(1828) had divided the genus into sections. Boissier himself used sec-
tions and subsections, but was apparently dissatisfied with the bases for
his division, since he comments in his notes on the genus (1862) "The
characters of the sections are often trifling and open to exception
but better ones cannot be found" (translation by Croizat, 1936a). We
may assume from this that Boissier maintained the genus Euphorbia in
its widest sense not only because of the common inflorescence type, but
also because he could find no consistent characters on which to divide
Many more species have been added to the genus since the time
of Boissier, until today Scholz (1964) estimates that it includes some
The retention of the genus in its broad sense has been opposed
on a number of grounds, and attempts have been made from time to time
to justify the segregation of certain groups. Croizat (1936a) has
pointed out that the possession of a cyathium is not restricted to this
genus, but occurs in several others recognized by even the most con-
servative taxonomists. Among these Pax (who had written the account
of the family for Engler and Prantl's Pflanzenfamilien in 1890) is
quoted as saying in 1924 that the cyathium was the main character of
the Tribe Euphorbieae rather than the genus Euphorbia. Croizat also
suggests that the species in Euphorbia are more variable in their degree
of evolution than would be expected in a single genus, and that the
cyathium is not an anatomic constant, and concludes that the use of the
single character of the presence of a broadly similar inflorescence,
cutting across other indications of relationship as it does, is unsound
from a taxonomic point of view.
Evidence from new areas of taxonomic research confirms the lack
of homogeneity in the genus. Perry (1943) reports that a survey of
somatic chromosomes in the Euphorbiaceae showed a considerable degree
of uniformity in size, shape, and basic number within all genera except
Euphorbia, which seemed to include three distinct groups.
Studies on the electrophoresis of latex by Moyer (1934) gave
groupings within the genus corresponding roughly to the sections of
Boissier. The work on pollen morphology by Erdtman (1952) and Punt
(1962), however, gave no clear divisions, although some of the patterns
of exine surface showed differences through the genus. A difficulty
with interpreting these studies lies in the fact that no voucher speci-
mens are available, so that any conclusions drawn must be tempered by
the uncertainty as to the exact taxonomic entities represented by the
Haber (1925), while not favoring a division of the genus, found
that the sections described by Pax in Engler and Prantl's Pflanzen-
familien (1890) showed little affinity in terms of anatomy and morphology
of the cyathium.
For these reasons, together with the diversity of habit and form
in the group, it is hard to accept that Euphorbia fulfills the condition
of relatively close ancestry which is usually imposed when judging the
soundness of a genus.
It is unfortunate that no comprehensive evaluation has been made
since Boissier's work more than a century ago, since many of the proposals
for segregating groups which have seemed inconsistent with usage in the
rest of the unchanged genus, might well have been perfectly acceptable in
the light of an overall revision. One such segregation to which this
criticism of inconsistency might apply is the separation of Chamaesyce
as a genus without considering the equally valid genera Tithymalus,
Agaloma and Poinsettia. It is beyond the scope of the present work
to examine all these genera, but Chamaesyce is such a discrete group
in the area under consideration, and so easily distinguishable from
other members of Euphorbia sensu lato, that it will be accepted here
as a distinct genus. A case will be presented for this division below,
but final judgment of its validity must wait until a revision of the
complete Tribe Euphorbieae of the Family Euphorbiaceae has been under-
II. The Seqregation of the Genus Chamaesyce
Chamaesyce was first proposed as a segregate genus by S. F. Gray
in 1821 with a single species which would appear automatically to become
the type species. Wheeler (1943), however, supports Millspaugh (1909)
in designating Euphorbia peplis L. as type since Gray was clearly describ-
ing this species under his name C. maritima and, in fact, gave E. peplis
in synonymy, thus making the later name illegitimate. The type specimen
of E. peplis L. in the Linnaean Herbarium becomes the type of C. maritime
S. F. Gray and of the genus Chamaesyce S. F. Gray.
Wheeler (1943) points out that the name had been validly pub-
lished as a subgenus, and suggested provisionally as a genus, by Rafinesque
in 1817, but such provisional publication is not acceptable. Gray was
justified in selecting this generic name even though Anisophyllum had
been proposed for the same group at this taxonomic level by Haworth in
1812, since that name was preoccupied by Jacquin's use of it in another
sense in 1763.
Gray took the name Chamaesyce from Dioscorides, a Greek herbalist
of the first century A.D., who compiled lists of plant names and uses
from medical works and from herb-gatherers and peasants (Gunther, 1959).
The word is a transliteration of two Greek components, chamae, of the
ground or low-growing, and Syce which was one of the names used for the
fig, and the association is probably due to the presence in members of
the genus Euphorbia of a milky latex similar to that found in fig trees.
The name was listed as a synonym for the genus Euphorbia by Wiman (1752)
with a similar comment that it was a type of low-growing fig (quasi
Gray's name was not widely adopted for a number of years after
publication, but the cohesion of the species which belong in the genus
is indicated by their treatment by subsequent authors. Roeper pub-
lished the group as Euphorbia section Anisophyllum in 1828, while
Rafinesque changed his mind after his publication of the subgenus
Chamaesyce in 1817 to treat them as a set of related genera in 1838.
They were put back into a single category as Euphorbia subgenus
Anisophyllum Caesalp. ex Reichenbach in 1841, treated as the genus
Anisophyllum by Klotzsch and Garcke (1859), and finally the definitive
work by Boissier in 1862 followed Roeper by using section Anisophyllum
for the group.
Croizat (1936a, 1936b, 1937) has examined these species which
have been combined so consistently by authorities on the family. They
share a number of morphological characters, which have been used by the
various authors in an attempt to delimit the group at the particular
taxonomic level which they favored. Boissier (1862), for example, used
leaf arrangement and shape, stipule shape, inflorescence type, number
and form of glands and the ecarunculate nature of the seed in his com-
parisons with the other twenty-six sections which he recognized, while
Wheeler (1941). considering only New World species, separated four sub-
genera on the basis of number and form of glands and their appendages,
and on leaf arrangement and stipule development. Boissier's dissatis-
faction with his separation has already been mentioned, and Wheeler is
equally pessimistic about his success in defining his divisions. There
is general agreement in the literature, however, about the characters
shared by this group of species. They all have opposite, petiolate
leaves with oblique bases and interpetiolar stipules. All have an
interesting distribution of chlorophyll-bearing tissue in the leaves,
where it surrounds the veinlets leaving clear areas between them, and
all have cyathia with four glands whose appendages, if present, are
petalloid, never lunate. The fifth gland is either obsolete or much
reduced and set in a sinus through which the capsule stalk often droops
at maturity, and, finally, the seeds are ecarunculate.
None of these characters is restricted to this group of species,
but Croizat (1936a, 1936b) has demonstrated that a particular type of
branching, in which the main axis aborts early and is replaced by one
or more secondary axes, is peculiar to the group, and serves as a
diagnostic feature. Here, then, is a discrete character of a fundamental
nature the presence or absence of a type of branching which gives
an absolute separation of a group of species from the heterogeneous
Euphorbia sensu lato. What is more, the species showing this type of
branching share many other morphological features, and have been accepted
as a cohesive group by most authorities on the family. On these grounds
their segregation as the distinct genus Chamaesyce appears well justified.
This separation has been adopted in the present work, and is
particularly appropriate in the New World where the species of Chamaesyce
are easily distinguishable from other members of the family on even a
casual examination. The usage is by no means novel in this area in that,
of the floras mentioned in the Introduction, only those by Urban and by
Fawcett and Rendle maintain Euphorbia sensu lato. The split has not been
as widely adopted in other regions, however, and an assessment of its
value on a worldwide basis must await a more comprehensive study.
The largest members of the genus Chamaesyce are found in the
Hawaiian Islands where some species have become arborescent. In the
Caribbean no such development is found, although one species, C.
vaqinulata (Griseb.) Millsp., has strong affinities with species of
the Galapagos Islands, and exhibits some of the peculiarities of
structure otherwise found only in endemics of that group. The remarks
in the following sections are intended to be limited to species grow-
ing under natural conditions in the Caribbean, which include annual and
perennial herbs of prostrate, ascending or erect habit, and shrubs and
subshrubs of open form reaching a height of one to six meters.
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.
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
only one secondary axis develops its thickening may obscure and eventually
conceal its origin, but in species in which a number of branches grow
(particularly those in which they lie prostrate) this apex is well marked.
On these secondary axes laterals (tertiary axes) may develop,
either throughout the length, or only near the lower or upper end. A
common form is the production of short, rather congested laterals from
the upper nodes on which most of the cyathia develop. Branching appears
to be alternate, with occasional production of other weaker branches at
the node, often becoming dichotomous near the apex, and may be at an
obtuse or acute angle with the stem. In the herbaceous species, stems
are wiry and brittle, or wider and of softer texture, often becoming
woody at the base. Rooting at the nodes is common in some species.
Prostrate species form tight or loose mats to 5 dm. diameter, the erect
herbaceous species may reach 6-8 dm.
Croizat (1936b) describes the growth of the secondary axes as
sympodial with each node terminated by a bud, a cyathium or a capitulum,
and the tertiary axes developed lateral to these. This is in agreement
with Roeper (1824) who explains the apparently normal lateral branching
as actually a reduction from the potential dichotomy which is expressed
at times at the tips of branches (Wheeler, 1941). The strongly alternate
branching which many species exhibit merely requires that the reduction
of the branches of the dichotomy alternates at the nodes. The same
pattern of branching may develop on tertiary axes, and rarely on those
of higher orders.
In the systematic part of this study the word "stem" will be used
to indicate the secondary axes, and lateralss" or "side branches" will
be used for tertiary and higher order axes. The aborted apex of the
stem, from which the secondary axes develop, will be called the crown
of the root for convenience in making comparisons with descriptions by
Leaves are opposite on the stem, and more or less two-ranked.
The blades are simple, membranous or sometimes coriaceous or sometimes
somewhat fleshy, and broad or rarely almost linear, 1 x I to 35 x 15 mm.
The base is inequilateral, usually markedly so, the margin entire or
variously toothed and plane or somewhat revolute, and the apex round
to acute or sometimes minutely apiculate. The veins are rarely promi-
nent, but in thin leaves the concentration of chlorophyll-bearing cells
around the conducting tissue with clear spaces between marks out the
reticulation of the veinlets.
Petioles are short, and the interpetiolar stipules may be
separate, or those on either side of the stem may be joined to the
corresponding one from the paired leaf. Their texture is usually mem-
branous, and the margins may be entire or variously toothed or parted.
In some species the pairs of leaflets droop at night in well-marked
These are solitary and terminal, but often appear axillary when
the growth of a lateral continues the axis. They are often borne on
congested leafy laterals or in short-stalked lateral and terminal
dichasia, and sometimes, by reduction, in leafless dichasial glomerules.
Each cyathium is itself a compound inflorescence, but strongly
simulates a single flower. A single terminal female flower with, at
most, a vestigial calyx to represent the perianth, is surrounded by
groups of male flowers, each flower reduced to a single stamen and
interspersed by bracteoles. These are enclosed in a cup-like involucre
made up of five foliaceous structures, united at their margins but with
free tips. Between these, glands are borne on the rim of the cup,
supplied by a trace from each of the adjacent foliar structures (Haber,
1925) and often with a petalloid appendage, which may be as long as,
and up to four times as wide as, the subcircular or transversely ellip-
tic gland. The fifth gland is obsolete or reduced to a linear structure,
and the space between the two lobes which it would occupy becomes a
sinus into which the stalk of the capsule often droops at maturity. The
cyathia are short-peduncled or rarely sessile. The term lobe is used
here and in the systematic section to indicate the free portion of the
foliar structures which are fused to make up the involucre.
The glands appear moist under magnification, and may produce a
substance attractive to insects. A large number of ants are often
found on and around the plants, and may be the agent for cross-pollination.
An articulation between the pedicel and stamen indicates that
these monandrous structures are each a reduced flower rather than simply
floral parts. Each is subtended by a bracteole which may grow up between
the flowers. The staminate flowers form in five fascicles, arranged
opposite the lobes of the involucre, and in each fascicle the flowers
develop centrifugally with respect to the cyathium axis. In our species
five, or rarely fewer, to about thirty is the usual number of staminate
flowers to each cyathium. The number is relatively constant in a
given species. The stamens are exserted at maturity, carried on
elongated glabrous pedicels (Wheeler's term "androphore" is adopted
for this pedicel in the systematic section). Dehiscence of the
anthers is transverse, and the serial maturation of the flowers spans
the time during which the female flower is developing.
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.
This is a trilocular capsule with one seed in each chamber,
usually roundly lobed or triangular but sometimes subspherical.
Dehiscence is by longitudinal splitting at the sutures, and to a
smaller extent at the angles. The base of the capsule is released
first, and the seed may be thrown a short distance from the flower by
the parting of the capsule wall.
The caruncle, which is so well developed in other species of
Eunhorbia sensu lato, is completely absent in Chamaesyce. The embryo
and endosperm are surrounded by a membrane and the testa, which may in
turn be covered by a persistent cellulose layer. This layer and the
testa may be patterned or sculpted.
The shape of the seed follows that of the trilocular capsule in
which it develops. In the systematic section the faces to the outside
of the capsule will be referred to as the dorsal faces, subtending the
dorsal angle, and the other two as the ventral faces subtending the
ventral angle. The seed is irregularly ovoid in longitudinal section
with the ventral side flattened. In cross section it is four-angled,
often obscurely so, but with the dorsal angle usually more acute and
pronounced than the ventral, and the lateral angles intermediate
between the two.
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.
All species have a milky latex to which various caustic properties
have been ascribed.
Morphological characters have proved to be very plastic within
many of the species examined. The habit of the herbaceous species
often varies widely, with occasional erect specimens in species which
are usually prostrate to decumbent, and with sprawling plants from species
which are normally completely upright to ascending. The number of stems
growing from the root is a more constant feature, with prostrate plants
more likely to have multiple stems than erect species, but the number
often changes with the change in habit in the case of erect specimens
of prostrate species.
The pubescence of the plant is usually reliable as a character
when contrasted with the absence of hairs (there are exceptions to this
among the south Florida species), but the degree of pubescence and even
the form of the hairs present is likely to vary over a wide range within
Leaf shape is variable, as might be expected, but in most cases
there is also a strong likelihood of toothed margins on occasional
leaves (or whole plants) of species which normally have entire leaves.
Cyathial characters in general show less plasticity, but the size
and form of appendages is rarely reliable, and vestiture may differ
widely from plant to plant or branch to branch. The same is true of
capsular characters, but the seed has proved to be stable enough that
characters relating to its morphology are often sufficient to key plants
out, and certainly to confirm doubtful identifications.
CHAMAESYCE S. F. Gray, Nat. Arr. Brit. PI., 2: 260. 1821.
Type: Chamaesyce maritima S. F. Gray. This is a superfluous name for
Euphorbia pepiis L., and the type specimen of this species in the
Linnaean Herbarium (sheet 630-18) is thus the type for C. maritima and
for the genus. Euphorbia subgenus Chamaesyce (S. F. Gray) House, Bull.
N. Y. State Mus. 254: 470. 1924.
Anisophyllum Haworth, Syn. PI. Succ., 159. 1812. Type:
Anisophyllum peplis (L.) Haw., designated by Wheeler, Rhodora 43: 110.
1941. Euphorbia section Anisophyllum (Haw.) Pax in Engler & Prantl,
Nat. Pflanzenfam., 3(5): 104. 1891.
Euphorbia subgenus Chamaesyce Raf., Amer. Monthly Mag. 2: 119.
1817. Type: Euphorbia supina Raf.
Euphorbia section Anisophyllum Roeper in Duby, A. P. de Candolle,
Bot. Gall. ed 2, 1: 412. 1828. Type: Euphorbia peplis L., designated
by Wheeler, Rhodora 43: 111. 1941.
Euphorbia subgenus Chamaesyce Caesalp. ex Reichenbach, Fl. Germ.
Excurs., 755. 1832. Type: Euphorbia chamaesyce L., designated by
Wheeler, Rhodora 43: 111. 1941.
Ditritra Raf., Fl. Tell., 4: 114. 1838. Type: Ditritra hirta
Raf., designated by Wheeler, Am. Mid. Nat. 30: 465. 1943.
Endoisila Raf., Fl. Tell., 4: 114. 1838. Type: Endoisila
Aplarina Raf., New Fl. N. Amer., 99. 1838. Type: Aplarina
prostrata Raf., designated by Wheeler, Rhodora 43: 111. 1941.
Annual or perennial herbs, often suffrutescent, or shrubs;
prostrate to ascending or erect; glabrous or variously pubescent; milky
latex in all parts. Main axis aborting, secondary axes few to many,
wiry or thicker, sometimes rooting at nodes. Leaves opposite, petiolate
with inter-petiolar stipules; blades expanded, bases inequilateral.
Cyathia terminal but appearing axillary, solitary at nodes or clustered
in cymules; lobes five, approximating glands; glands four or with fifth
vestigial, alternating with lobes, with or without petalloid appendages.
Staminate flowers monandrous, naked, few to many, maturing serially,-
exserted when mature, anther dehiscence transverse. Pistillate flower
solitary, terminal; perianth reduced to pad of tissue or vestigial
calyx; very three-celled, each with a single ovule; styles three, free
or joined at base, usually bifid for part of length. Capsule dehiscent
at sutures. Seed ecarunculate, coat smooth or variously sculptured.
Plants of disturbed habitats, probably poor competitors in
many cases. Many species favor sandy soils, either coastal or inland
and often unstable, while others are restricted to pinelands whose
characteristic flora is maintained by periodic burning. The most common
species, C. maculata in Florida and C. hyssopifolia in the New World
tropics as a whole, follow man, and are frequent as weeds in paved areas
and cultivated or abandoned ground. Many species have a wide geographic
range, passing unfavorable periods of the year as seed or with growth
almost quiescent and quickly resumed when conditions improve. Seed pro-
duction is abundant in most species.
The use of the taxonomic categories "species", subspecies", and
"variety" in this work follows conventional lines (see, for example,
Davis and Heywood, 1963). They are considered to constitute an hier-
archy, and the increasing degree of morphological homogeneity as the
rank of the group decreases is taken to imply a greater freedom of
gene interchange within the group. The use of these three ranks is
a compromise between the naming as a distinct taxon of any population
which is prevented by any means from exchanging genes with other popu-
lations, whether or not morphological differences have become established,
and the more classical approach of applying names only to taxa which are
discrete by a set number of morphological characteristics.
Key to the Species of Chamaesyce in the Caribbean
1. Much-branched shrub, low-growing or 2-6 m. tall ..... 2.
I. Annual or perennial herbs, often woody at the base
and suffrutescent, but rarely reaching 1 m. ..... 3.
2. Leaves fleshy, subglobose or linear; closely branched
with stems often almost prostrate .... C. vaqinulata.
2. Leaves membranous, ovate to ligulate; branching
divaricate, stems upright ..... 2. C. articulata.
3. Capsule glabrous ..... 4.
3. Capsule pubescent ..... 24.
4. Plants erect or ascending ..... 5.
4. Plants prostrate to decumbent ..... 12.
5. Leaf margins manifestly toothed, or, if entire,
leaves ligulate ..... 6.
5. Leaf margins entire or obscurely toothed below apex,
leaves never ligulate ..... 10.
6. Stems several to many from the crown of a heavy root-
stock, wiry, rarely more than 1 mm. diameter ..... 13. C. brittonii.
6. Stems few to several, thick, sometimes
reaching 4 mm. diameter ..... 7.
7. Cyathia few, borne singly or rarely in groups of two
or three; plant perennial ..... 12. C. parciflora.
7. Cyathia many, grouped on short leafy laterals or in
glomerules; annual but sometimes perennating
8. Capsule less than 1.4 mm. long; cyathia usually in
peduncled leafless glomerules ..... 9
8. Capsule more than 1.6 mm. long; cyathia on
9. Tips of shoots pubescent, at least in lines on
the side of the stem; leaves pilose beneath; seed
with a rippled surface
9. Tips glabrous; leaves glabrous or sparsely pilose
beneath; seed with 2-4 lateral ridges
on each face .....
. C. hypericifolia.
..... 8. C. nutans.
10. C. hyssopifolia.
10. (5) Leaves and young stems fleshy; stipules conspicuous,
membranous, ca. 1 mm. long, white ..... 3. C. buxifolia.
10. (5) Leaves membranous or somewhat coriaceous, young
stems not fleshy; stipules not conspicuous unless
because of ciliate inner surface, coriaceous,
ca. 0.5 mm. long, brown
11. Stipules densely white ciliate on surface nearest stem;
leaf and young stem surface minutely white-papillose;
glands dark, appendages obsolete .... 4. C. lecheoides.
11. Stipules not ciliate; stem surface smooth, somewhat waxy,
leaf surface with raised markings but not papillose;
glands green or red, appendages small or to twice
width of gland ..... 6. C. porteriana.
12. (4) Stems wiry and stiff, rarely exceeding 0.5 mm.
diameter, usually many from root stock
12. (4) Stems flexible rather than wiry, few to several
from rootstock, either reaching 2-3 mm. diameter or
13. Leaf margins entire
Leaf margins serrate, at least in basal leaves
14. Leaves deltoid, margin somewhat revolute; glands fleshy,
appendages obsolete or minute ..... 41. C. deltoidea ssp. deltoidea
14. Leaves orbicular to elliptic, plane; glands cupped,
appendages prominent, to three times
as wide as gland ..... 19. C. camaguayensis.
15. All leaves serrate, most 4-8 mm. long; plants usually
decumbent ..... 18. C. torralbasii.
15. Only basal leaves serrate, most 2-3 mm. (rarely
to 5 mm.) long; plants usually forming a
lax, prostrate mat ..... 17. C.
16. (12) Leaves and stems pubescent ..... 39. C
16. (12) Leaves and stems glabrous
17. Stems minute, threadlike, tomentose on upper surface;
cyathia subsessile; styles undivided at apex; capsule
less than 0.7 mm. long ..... 42
17. Stems often reaching 2-3 mm. diameter, glabrous;
cyathia with an obvious peduncle, styles bifid for
part of length; capsule more than 1 mm. long
18. Stipules not united, or minutely so at base, deeply
parted or laciniate
18. Stipules joined, at least on upper or lower surface of
tips of branches, fringed or entire, never laciniate
19. Seeds strongly angled; appendages conspicuous; leaves
ovate to ovate-oblong, almost as wide as long..... 20.
19. Seeds terete to obscurely angled; appendages incon-
spicuous or absent; leaves narrowly elliptic
to oblong, much longer than wide
. C. tumistyla.
20. Capsule more than 2.5 mm. long; seed cuneiform,
more than 2 mm. long ..... 21. C. polygonifolia.
20. Capsule less than 2.5 mm. long; seed ovoid, less
than 2 mm. long
21. Seed 1.4-1.9 mm. long; leaves usually fleshy, size
often widely different on main stem
and laterals ..... 2:
2. C. ammannioides.
21. Seed 1-1.4 mm. long; leaves not fleshy, all
similar in size ..... 23. C. cumulicola.
22. (18) Stipules toothed at apex, white, prominent
although only ca. 0.5 mm. long; faces of seed smooth;
glands brown or deep purple, appendages white ..... 16. C. serpens.
22. (18) Stipules toothed or bifid almost to base, green or
tan colored, not conspicuous although often more than
1 mm. long; faces of seed obscurely transversely
ridged; glands greenish, appendages white, yellor or pink ..... 23.
23. Stipules short, to 0.4 mm.; texture of all parts of
plant subcoriaceous; leaf and stem surface
somewhat papillose ..... 14. C. cowellii.
23. Stipules long, often more than 1 mm.; leaf texture only
rarely firmer than membranous; leaf and stem surface
smooth ..... 15. C. blodqettii.
24. (3) Leaf margins manifestly serrate ..... 25.
24. (3) Leaf margins entire or obscurely toothed ..... 36.
25. Plants erect with one or few thick stems ..... 11. C. lasiocarpa.
25. Plants prostrate to decumbent with several to many
stems which rarely exceed 2 mm. diameter ..... 26.
26. Cyathia solitary at leafy nodes, appearing clustered
if on congested laterals, but not in glomerules ..... 27.
26. Cyathia in peduncled glomerules ..... 34.
27. Appendages of glands subequal in size, often much
reduced, or if two appendages much longer than other
two, capsule not fully exserted, splitting one side
of cyathium at maturity, glands subcircular or some-
what transversely elongated ..... 28.
27. Appendages of glands greatly unequal in size, one pair
longer than the other pair; capsule fully exserted and
nutant at maturity, often concealed by long appendages;
glands enlarged in length, to extend much of the
distance around rim of cyathium ..... 33.
28. Capsule pubescent only along the angles ..... 29.
28. Capsule pubescent all over ..... 30.
29. Stem short-pubescent in lines at sides and so
upper surface; faces of seed with deep,
29. Stem long-hirsute, at least in lines at sides;
surface of seed rippled
30. Capsule not completely exserted, splitting one
side of cyathium at maturity
Capsule completely exserted at maturity
..... 24. C. prostrata.
..... 25. C. mendezii.
.... 32. C. thymifolia.
Leaf margin and coarsely serrate, often incised
1/2-1/3 distance to midrib; stems wiry, scarcely reaching
1 mm, diameter, many from heavy root stock ..... 26. C. hepatica.
31. Leaf margin serrate but not deeply incised; stems not
wiry, usually 1 mm. or more in diameter, few to
several from annual or perennating root stock
32. Stem deliquescent through dichotomous branching in upper
part; leaf texture heavy; seeds with two broad faces
and one angle acute, almost triangular in
cross-section ..... 27. C
32. Stem excurrent, forming only congested leafy laterals in
upper part; leaf texture membranous; seeds with all
faces subequal, angles obtuse, almost square
in cross-section ..... 31. C. maculata.
33. (27) Stems long-pilose on upper surface; leaf apex acute;
cyathia congested on short laterals ..... 34. C. conferta.
33. (27) Stems short-tomentose or strigose on upper surface;
leaf apex obtuse or rounded; cyathia borne singly or
in small groups at upper nodes, if on laterals
not congested ..... 33. C. adenoptera.
34. (26) Leaf margins crenate-dentate or roundly
serrate, apex obtuse
34. (26) Leaf margins sharply serrate, apex acute
.... 30. C. berteriana.
35. Cymules terminal and lateral on leafless peduncles; stem
branching at base but only infrequently near tips (when
plant may appear to have cymules on laterals with a pair
of subtending leaves); mostly robust, ascending,
large-leaved plants ..... 28. C. hirta.
35. Cymules terminal and on leafy laterals; stem branching
freely; mostly low, small-leaved plants ..... 29. C. opthalmica.
36. (24) Plants suffrutescent; branches to 5 dm. long ..... 37.
36. (24) Plants herbaceous, or, if woody at the base, of small
stature, branches rarely exceeding 1 dm. long ..... 39.
37. Appendages obsolete; glands deep purple ..... 5. C. cayensis.
37. Appendages prominent; glands yellow or green ..... 38.
38. Open shrubs, branched from base; capsules more
than 2 mm. long ..... 7. C. myrtillifolia.
38. Suffrutescent, base mostly unbranched; capsules
1.3-1.5 mm. long ..... 6. C. porteriana var. keyensis.
39. (36) Stems pubescent on only one surface ..... 40.
39. (36) Stems pubescent on all surfaces ..... 41.
40. Stems tufted, upright, usually less than 0.5 mm.
diameter; leaf surface glabrous ..... 37. C. minutula.
40. Stems prostrate, usually ca. I mm. diameter;
scattered white hairs on
surface of leaf ..... 36. C. multinodis.
41. Cyathia in groups of 2-5 at nodes ..... 43. C. leonardii.
41. Cyathia solitary at nodes ..... 42.
42. Plants robust, stems 1-3 mm. diameter, to 3 dm.
long, leaves 4-9 mm. long ..... 35. C. garberi.
42. Plants delicate, stems wiry, scarcely reaching 1 mmn.
diameter even in old plants, to 2 dm. long,
leaves 2-5 mm. long ..... 43.
43. Plants closely appressed forming a dense mat, sometimes
becoming diffuse with age ..... 41. C. deltoidea.
43. Plants decumbent at tips or erect ..... 44.
44. Stems almost unbranched, villous-hirsute, tips canescent;
capsule reniform, sharply three-lobed,
angles acute ..... 40. C. pinetorum.
44. Stems branched freely, particularly in upper part of stem,
sparsely or densely short-pilose, tips only occasionally
canescent; capsule ovoid, roundly lobed,
angles obtuse ..... 38. C. turpinii.
1. CHAMAESYCE VAGINULATA (Griseb.) Millsp., Field Mus. Bot. 2:
302. 1909. Euphorbia vaqinulata Griseb., Fl. Brit. W. I. Islands, 52.
1859. Type: Turk Islands, Hjalm. (Lectotype: K !, fragment NY !).
Designation of the lectotype is made here. The specimen at Kew was
chosen because there is a strong probability that it was examined by
Grisebach. Many of Hjalmarsson's collections were deposited at Systema-
tisch-Geobotanisches Institute, Goettingen, and an isotype may exist
there. The photograph of a Hjalm. sheet from Kew at IJ can have no
standing as type material since it is not of this species but of C. ?
lecheoides (Millsp.) Millsp.
Shrub; prostrate or ascending to 2 m. Stem to 1.5 cm. diameter,
internodes to 3 cm. or congested; branching from base and throughout
length, laterals often lying in one plane, not rooting at nodes, glabrous,
becoming dark brown with age. Leaves fleshy; blades subglobular or
linear-elliptic, 1.5 mm. diameter or 5 x 1-1.5 mm., base subequal, sub-
cordate or rounded, margin entire, apex rounded, subarticulate from
petiole, glabrous, yellow-green to purple-brown; petiole 0.3-1 mm. long;
stipules sheathing stem and persistent, short, ciliate, white. Cyathia
solitary at nodes, often on vestigial branches; peduncle short, somewhat
expanded into obconical involucre, to 1 mm. diameter at mouth, lobes
fleshy, equalling glands, triangular, glabrous on outside, ciliate within,
glands transversely elliptic, 0.5 x 0.3 mm., fleshy, deep purple, append-
ages absent, fifth gland triangular, large, equalling lobes or rarely of
same form as other glands, sinus shallow. Staminate flowers 3-5 per
cyathium or absent, androphores glabrous, scarcely exserted. Gynophore
glabrous, to 1.5 mm. long at maturity, exserted, usually upright;
calyx a triangular pad of tissue; ovary glabrous; styles spreading,
o.4 mm. long, joined at base, bifid for 1/2 length. Capsule glabrous,
globose, 1.5 mm. long, 1.8-2 mm. at equator, roundly three-lobed,
angles rounded. Seed ovoid, 1.5 x 1.2 mm., scarcely angled, usually
Restricted to a few islands of the southern Bahamas, but
locally frequent on these in rocky and sandy areas behind the shore
Representative specimens seen: BAHAMAS. Inagua, 6 Mar. 1962,
Dunbar 107 (GH); Turtle Cove, Inagua, 19 Oct. 1904, Nash & Taylor 1170,
(NY); Caicos Islands: North Caicos, 24 Apr. 1954, Lewis s.n. (GH, IJ);
Bambarra, Middle Caicos, 2 July 1954, Proctor 9096 (IJ); South Caicos,
14-16 Dec. 1907, Wilson 7681 (GH, NY). Turks Islands: Waterloo, Grand
Turk, 20 Feb.-24 Mar. 1911, Millspaugh & Millspaugh 9055 (GH, NY);
Grand Turk, 27 Aug.-l Sept. 1905, Nash & Taylor 3764 (NY); Long Cay,
12 June 1954, Proctor 8793 (GH, IJ). Watling's Island: Southeast End,
27, 28 Nov. 1907, Wilson 7259 (NY). Castle Island, 22 Dec. 1907, Wilson
The fleshy leaves and involucres, and the shrubby habit with the
closely branched stems often held almost prostrate, make this quite
different from any other species in the area. Boissier (1862) grouped
it with endemics of the Galapagos Islands on the basis of its appearance,
fleshiness, and the persistent, ciliate stipules that ring the stem,
but this morphological similarity is probably fortuitous and not an indi-
cation of close common ancestry. Many species of other families grow in
I NEW YORK OTif (l .Iiln
S9- ? TH-E C AICO
plate 1. CHAMAESYCE VAGINULATA (Griseb.) Millsp.
C. va inulata (Griseb.) Millsp., Long Cay, Caicos Group, Bahamas, 7 Mar.
1911, Millspaugh & Millspaugh 9237 (NY).
'"> --- ------
Map 1. Distribution of C. vaqinulata
'.-- --. -
Map 2. Distribution of buxif
Map 2. Distribution of C. buxifolia
both the Bahamas and the Galapagos, but most of these are common through-
out the intervening tropics, and give no hint of a disjunction of range
such as this species would show from those which it resembles.
2. CHAMAESYCE ARTICULATA (Aubl.) Britton, Mem. N.Y. Bot. Card. 6:
574. 1916. Euphorbia articulata Aubl., P1. Guian., 1: 480. 1775. Type
in Herb. Brit. Mus. if extant.
Euphorbia linearis Retz., Observationes Botanicae, 3: 32. 1783.
Type: Bot. Mus., Lund, Sweden, if extant. E. linearls heterophylla
Kuntze, Rev.Gen. Pl., 605. 1891. C. linearis (Retz.) Millsp., Field
Mus. Bot. 2: 410. 1916.
Anisophyllum vahlii K1. & Gke., Monats. Akad. Berlin 1859: 36.
1859. Type: Specimen by Willdenow, in Bot. Mus. Berlin, if extant.
Euphorbia vahlii Willd. ex K1. & Gke., loc. cit. 37. 1859. Chamaesyce
vahlil (K1. & Gke.) P. Wilson, Bull. N.Y. Bot. Gard. 8: 395. 1917.
Shrub; to 6 m., erect. Stems to 2 cm. at base, internodes to
5 cm. long; divaricate-branched throughout length, glabrous or canescent
at tips, light brown becoming black with age. Leaves membranous; blades
elliptic, broadly so to linear, 20-60 x 2-12 mm., base subequal, rounded,
margin entire, slightly thickened, apex obtuse, mucronate, midrib promi-
nent on underside, glabrous or short pubescent, light green; petiole
1 to 1.5 mm. long; stipules joined, deltoid, to 0.5 mm. long, densely
ciliate on inner surface, somewhat persistent. Cyathia solitary; peduncles
5 to 7 mm. long, involucre obconical, to 2 mm. diameter at mouth, lobes
scarcely equalling glands, deltoid, glabrous on outside, densely ciliate
within, glands broadly transversely elliptic, to 1 mm. long, broadly
cupped, appendages absent, fifth gland deltoid, short, sinus broad, shallow.
Staminate flowers 8 20 per cyathium, androphores glabrous, to 2.5 mm.
long at maturity. Gynophore glabrous, to 3 mm. long at maturity, ex-
serted, nutant; calyx vestigial; ovary glabrous; styles erect, to 3 mm.
long, joined for half length, bifid in apical 1/3. Capsule glabrous or
minutely pubescent, ovoid or conical, 2.3 mm. long, 2.3 mm. wide above
base, roundly three-lobed, angles rounded. Seed elliptic-ovoid, 1.8 2
mm. long, 1.2 1.4 mm. radial width at equator, four-angled, the dorsal
faces longer and dorsal angle more acute than others, surface pitted and
wrinkled, brown or ashen.
Frequent from Puerto Rico south through the Lesser Antilles to the
northern coast of South America. No collections have been seen from the
Bahamas since those of Millspaugh in 1911 from East Caicos, but these
specimens are undoubtedly of the species (Map 10).
Representative specimens seen: BAHAMAS. East Caicos: Rocky plain,
Jacksonville and vicinity, 26-27 Feb. 1911, Millspaugh & Millspaugh
9086 (F, NY); Low scrub, Jacksonville and vicinity, 26-27 Feb. 1911,
Millspaugh & Millspaugh 9102 (F, NY). PUERTO RICO. Susua State Forest,
28 June 1961, Alain 9265 (IJ); Yauco, 2 Apr. 1880, Garber s.n. (GH);
Seashore, Fajardo, 17 Feb. 1900, Heller 4608 (F, GH, MO, NY); Santurce,
21 Feb. 1900, Heller 4678 (F, MO, NY); Rio Piedras, Fajardo, 4 July 1913,
Johnston 792 (NY); Guayanilla to Tallahoa, 13 Mar. 1913, Shafer 1993 (F,
NY); Fajardo, 19 May 1885, Sintenis 1612 (MO, NY). LESSER ANTILLES.
Culebra, 3-12 Mar. 1906, Britton & Wheeler 4 (F, NY); Culebras Island,
19-20 Jan. 1899, Millspaugh 1781 (F); Racoon Bay, St. Thomas, 31 Jan.-
4 Feb. 1913, Britton, Britton & Shafer 167 (F, NY); St. Thomas, Jan. 1887,
Eggers s.n. (F); Tortola, 16 Nov. 1918, Fishlock 263 (F, GH, NY); Anagada,
19-20 Feb. 1913, Britton & Fishlock 978 (NY); Barbuda, 6 Apr. 1956, Smith
10466 (GH, IJ). St. Kitts: Canada Estate, 8 Sept.-5 Oct. 1901, Britton &
E. linearis Retz., St. Thomas, ? Bertero (MO).
E. linearis Retz., St. Thomas, ? Bertero (MO).
Cowell 263 (NY); Sir Timothy's Hill, 24 Dec. 1958, Proctor 18418 (IJ);
Nag's Head, 27 Dec. 1958, Proctor 18514 (IJ). Antigua: Santa Maria Hill,
25 Apr. 1937, Box 756 (MO); Nook Hill, 3 Oct. 1937, Box 1134 (MO).
Montserrat, 8 Feb. 1959, Proctor 19079 (IJ); Guadeloupe, 1892, Duss
2451 (NY); Martinique, Sieber 392 (F); St. Lucia, 22 Apr.-18 May 1950,
Howard 11417 (IJ). St. Vincent: Mt. Pleasant, 22 Mar. 1962, Cooley
8534 (GH); Bequia, 26-31 Mar. 1950, Howard 11269 (GH).
The affinities of this species are with C. myrtillifolia (L.)
Millsp. and with coastal species of the Bahamas and Florida such as
C. lecheoides (Millsp.) Millsp. and C. porteriana Small. These, however,
differ in size, habit, cyathial and stipular characters.
A number of pubescent collections of C. articulata have been
compared with the more common glabrous plants, but these prove to be
only the extremes of a range, with many intermediates showing occasional
shoots with minute velvety pubescence.
The name is accepted although no type material has been seen.
There is general agreement in the literature with this application, and
that E. linearis is a synonym.
3. CHAMAESYCE BUXIFOLIA (Lam.) Small, Fl. SE. US., 712. 1903.
Euphorbia buxifolia Lam., Encyc. 2: 421. 1786. Type: plate in manuscript
of Botanicum Americanum t.4, f.2 by Plumier, in the Bibliotheque General,
Museum D'Histoire Naturelle, Paris.
E. qlabrata Sw., Prodr., 76. 1788. Type: unknown. A specimen
from Swartz with this name is reported from Herb. Stockholm, Fawcett &
Rendle, Fl. Jam., 4(2): 336. 1920.
E. litoralis HBK., Nov. Gen. et Sp., 2: 44. 1817. Type: Cumana,
prope Bordones (Herb. Mus. Paris, not seen).
E. flexuosa HBK., Nov. Gen. et Sp., 2: 44. 1817. Type: Cumana,
Bonpland (Herb. Mus. Paris, not seen). E. buxifolia Lam. var. flexuosa
(HBK.) Soiss. in DC., Prod., 15(2): 15. 1862.
Perennial shrub or subshrub; erect to ascending, or nearly
decumbent in shifting sand, to 6 dm. Stems to 8 mm. diameter at naked
woody base, internodes to 3 cm.; branching sparse at base, more frequent
near shoot apex, glabrous, green becoming darker. Leaves somewhat
fleshy; blades ovate to elliptic, 5-12 x 3-6 mm., base slightly unequal,
truncate to cordate and obscuring stem, margin entire, apex obtuse, mid-
rib sometimes prominent on underside of folded leaf, glabrous, yellowish
to dark green and glaucous; petiole 0.5 to 1 mm. long; stipules united,
to 1 mm. long, ligulate or deltoid, short cleft or fringed, white.
Cyathia solitary at upper nodes; peduncles 0.5-1 mm. long, cyathia campanu-
late, to 0.7 mm. diameter at mouth, lobes equalling glands, triangular
glabrous outside, densely ciliate in throat, glands transversely elliptic,
to 0.5 mm. long, brown, fleshy, appendages slightly longer than, and as
wide as gland, white, margin entire or undulate, fifth gland linear,
shorter than lobes, sinus shallow, broad. Staminate flowers 12-20 per
cyathium, androphores glabrous, to 1 mm. long at maturity. Gynophore
glabrous, to 3 mm. long at maturity, exserted, nutant; calyx a triangular
pad of tissue; ovary glabrous; styles spreading, 0.3-0.4 mm. long,
joined at base, bifid for 1/2 length. Capsule glabrous, subspherical
1.5-2 mm. long, 2.2-5 mm. wide just below equator, deeply three-lobed,
angles rounded. Seed broadly ovoid, 1.3 mm. long, 1.2 mm. wide just below
equator, angles mere ridges except near apex, faces plump, convex, surface
obscurely pitted, ashen.
A frequent component of the vegetation of sandy and rocky shores,
.I I I II I. I
Plate 3. CHAMAESYCE BUXIFOLIA (Lam.) Small
C. buxifolia (Lam.) Small, Grassy Key, Monroe County, Florida, 9 Aug.
1963, Burch 153 (FLAS).
and of their associated beach-scrub; Bermuda, Florida in all coastal
areas south of Pinellas and Volusia counties, Bahamas, Yucatan, Cuba,
Jamaica, Hispaniola, Puerto Rico, Lesser Antilles and the northern
coast of Venezuela (Map 2).
Representative specimens seen: FLORIDA. Brevard: Cape Canaveral,
2-5 Apr. 1904, Burgess 668 (F, NY); Merritt Island, 18 Apr. 1946,
Foster & Smith 1364 (DUKE, F, FLAS, IJ, MO, NSC, NY); Malabar Cape, 17
Oct. 1964, Lakela (FLAS, USF). Broward: Port Everglades, 3 Aug. 1955,
Godfrey & Kral 53854 (FSU, NSC); Lauderdale Beach, 17 Dec. 1929, Moldenke
263a (DUKE, MO, NY). Collier: Vanderbilt Beach near Naples, 13 Aug. 1963,
Burch 182 (FLAS); Marco Island, 19 Mar. 1938, Standley 57622 (F). Dade:
Soldier Key, 30 Mar. 1904, Britton 337 (F, NY); Miami, May 1877, Garber
s.n., (FLAS, NY); Key Biscayne, 17 Jan. 1937, McAllister 210a (DUKE);
Lemon City, 6 Nov. 1903, Small & Carter 1116 (NY). Flagler: Beach near
Marineland, 18 Nov. 1961, Godfrey 61675 (FSU). Indian River: Vero Beach,
7 Aug. 1963, Burch 112 (FLAS). Lee: Sanibel Island, 19 Dec. 1953,
Cooley 2401 (FLAS, NY); Punta Rassa, July-Aug. 1900, Hitchcock 325 (F,
MO); Captiva Island, 21 May 1901, Tracy 7226 (F, MO, NY). Manatee:
Anna Maria Key, 12 July 1917, Cuthbert 1578 (FLAS, NY); Sneed's Island,
9 Oct. 1899, Tracy 6377 (F, MO, NY). Martin: Atlantic side of Jupiter
Island, 7 Aug. 1963, Burch 125 (FLAS). Monroe: Cape Sable, Middle Cape,
18 Apr. 1964, Burch & Ward 304 (BM, BUS, E, F, FLAS, GH, IJ, MO, NSC, NY);
New Found Harbor Key, 13-14 Feb. 1935, Killip 31395 (NY); Marquesas Keys,
10-12 Mar. 1904, Lansing, Jr. 2225 (F, NY); Key Largo, 12 Jan. 1930,
Moldenke 402 (DUKE, MO); Key West, 1874, Palmer 499 (MO, NY); Big Pine
Key, 27 June 1960, Webster & Samuel 10210 (DUKE, IJ, PUR). Palm Beach:
Beach between Palm Beach and Lake Worth, 7 Aug. 1963, Burch 118 (FLAS);
Palm Beach, 24 May 1895, Curtiss 5404 (NY); Juno Beach, 20 Apr. 1962,
Lakela 24955 (FLAS, USF); Lake Worth Inlet, 28 Dec. 1895, Webber 236
(F, MO). Pinellas: St. Petersburg, Williamson s.n. (NY). St. Lucie:
North of Ft. Pierce, 7 Aug. 1963, Burch 120 (FLAS). Sarasota: Longboat
Key, 20 Nov. 1964, Godfrey 65236 (FSU); South end of Longboat Key,
6 Oct. 1964, Long & Lakela 27547 (FLAS, USF); Casey Keys, Spring 1887,
Rothrock s.n. (F, NY). Volusia: Turtle Mound, 27 Apr. 1961, Ray, Wood,
Smith & Eaton 10821 (FSU, NCU). BAHAMAS. Great Bahama, 5-13 Feb. 1905,
Britton & Millspaugh 2365 (F, NY); Bimini, 27 July 1964, Stimson 776
(DUKE, FLAS); Riding Rocks, Great Bahama Bank, 13 May 1909, Wilson 7914
(F, MO, NY); New Providence, Nov. 1890, Hitchcock 26 (F, MO); Salt Key,
21 May 1909, Wilson 8089 (F, MO, NY); Watling's Island, Graham's Harbor,
26 Nov. 1907, Wilson 7228 (F, NY); Hog Island, Jan. 1905, Wight 55a
(F, NY); Inagua, 15-25 Aug. 1960, Williams 7 (DUKE, PUR); South Caicos,
27 June 1954, Proctor 8981 (IJ); Grand Turk, 22-27 Apr. 1954, Lewis s.n.
(IJ). CUBA. Isles of Pines: Caleta Cocodrilos, 8 Mar. 1916, Britton,
Wilson & Leon 15295 (F, NY). Havana: Havana, 6 May 1905, Curtiss 752
(F, MO, NY). Oriente: Santiago, Siboney, 1917, Ekman 8719 (F). Camaguey:
Cayo Sabinal, 9 Mar. 1909, Shafer 699 (F, NY); Cayo Palo, 24 Oct. 1909,
Shafer 2733 (F, NY). Pinar del Rio: Las Martinas to the coast, 19 Dec.
1911, Shafer 11105, (MO, NY). CAYMAN ISLANDS. Cayman Brac., South
Shore, 10 Feb. 1899, Millspaugh 1233 (F); Grand Cayman, Grape Tree Point,
14 Apr. 1956, Proctor 11973 (IJ). JAMAICA. Portland: Port Antonio, 21
June 1897, Fredholm 3038 (NY). St. Catherine: Healthshire Hills, 3 May
1913, Harris 11627 (F, MO, NY, UCWI). St. Elizabeth: Long Acre Point,
13 Sept. 1907, Harris 9967 (F, NY, UCWI). St. James: Montego Bay,
June 1951, Robbins 423 (UCWI). St. Mary: Galina Point, 7 Dec. 1952,
Proctor 7452 (IJ). St. Thomas: Morant Point, 1-13 Mar. 1909, Britton
4098 (F, NY). Westmoreland: Negril area, Homer's Cove, 23 June 1959,
Webster, Ellis & Miller 8501 (PUR). HAITI. Bayeux, 28 Nov. 1924,
Ekman H2676 (IJ); Port de Paix, 28 Apr. 1925, Ekman H3957 (IJ); Gonave
Island, Aug. 1927, Eyerdam 213 (F, NY); Tortue Island, 3-10 May 1929,
Leonard & Leonard 15328 (MO). DOMINICAN REPUBLIC. Monte Cristi: Monte
Grande, 12 July 1929, Ekman H13165 (F, PUR). Barahona: San Domingo,
May 1911, Fuertes 1111 (F, NY); Trail between Pedernales and Aceitial,
8-12 Aug. 1946, Howard 8106 (NY). PUERTO RICO. Condado, San Juan,
16 Sept. 1941, Blomquist 11789 (DUKE); Fajardo, 4 July 1913, Johnston
791 (NY); Ratones Island, 21 Jan. 1899, Millspaugh 658 (F); Salinas de
CaboRojo, 6 Feb. 1885, Sintenis 559 (UCWI); Beach at Santurce, 21 Feb.
1900, Heller 4681 (F, MO, NY). LESSER ANTILLES. Soldier Bay, St.
Thomas, Dec. 1880, Egger s.n. (F); St. Croix, I Nov. 1895, Ricksecker
36 (F, MO, NY); Anguilla, 1908, Boldingh 3561 (NY); Barbuda, 27 Apr.
1959, Cowan 1656 (IJ, NY); Guadeloupe, 1894, Duss 2741 (MO, NY);
Martinique, 25 June 1939, Egler 173 (NY); St. Lucia, I Aug. 1958, Sauer
2174 (F); St. Vincent, Smith & Smith, 208 (NY); Grenada, Pointe Saline,
23 Sept. 1945, Beard 1222 (MO, NY). Extra-territorial specimens: BERMUDA.
Ducking Stool, 31 Aug.-20 Sept. 1905, Brown & Britton 6 (F, NY); Agar's
Island, 22 July 1913, Collins 219 (F, NY); Rocky shore, 18 Mar. 1910,
Flynn 101 (MO); Hamilton Bay Islets,. 29 Dec. 1898, Millspaugh 10 (F).
The species named by Kunth are no more than growth forms of this
taxon, which is just as plastic in its response to soil and wind conditions
as the completely herbaceous species.
Euphorbia mesembrianthemifolia Jacq., Enum. PI. Carib., 22.
1760, and Sel. Stirp. Amer., 150. 1763, was put into synonymy for
this species by Boissier (1862) on the basis of Jacquin's description.
This is unacceptable, since, if the name can be linked to the species
beyond reasonable doubt, it is much earlier than Lamarck's and would
have to replace it. Jacquin's description suggests this species in
many respects, but has no character which is diagnostic. However, if
it could be shown that specimens of this entity were in Jacquin's
possession at the time that his description was written, this would
support the acceptance of his name. Mr. W. T. Steam of the British
Museum (Natural History) was kind enough to search their herbarium, and
Dr. K. H. Rechinger checked that of the Naturhistorisches Museum, Vienna,
but in neither case were any specimens found which were known to have
passed through Jacquin's hands at this time. Many of Jacquin's specimens
were lodged in these two herbaria, and the absence of any relevant to
this problem, together with the fact that Boissier did not see any
specimens to support the name, suggest that none have survived. With
nothing more than Jacquin's description to typify the name E. mesembri-
anthemifolia, it would be irresponsible to accept it and upset the well-
understood use of C. buxifolia, which will be maintained here until some
firm evidence supports a change.
4. CHAMAESYCE LECHEOIDES (Millsp.) Millsp., Field Mus. Bot. 2:
Shrub; erect or ascending to 4 dm. Stem to 8 mm. diameter, inter-
nodes to 2 cm.; branched either from crown of root and throughout length
or from only the upper half of the single or few stems from base,
branching open or strict, old stem ringed by stipule scars, glabrous,
brown. Leaves membranous, somewhat coriaceous; blades ovate to linear
elliptic, 3-7 x 1.5 5 mm., midrib sometimes prominent on under side
of leaf, marked by depression on upper surface, glabrous, the surface
minutely white-papillose, base inequilateral, rounded to subcordate,
margin entire, somewhat revolute, rarely crenulate or serrate toward
apex, apex obtuse or rounded, gray-green or pink; petiole 0.5 I mm.
long; stipules distinct or joined, deltoid to 1 mm., densely ciliate
on inner surface, brown, cilia white. Cyathia in upper nodes; peduncles
to 1 mm. long, involucre campanulate, 1-1.6 mm. diameter at mouth, lobes
equalling glands, triangular, glabrous on outside, densely pilose within,
glands broadly transversely elliptic, 0.4 0.7 by 0.2 mm., often dark
purple, rarely much reduced, appendages obsolete, fifth gland vestigial
or absent, sinus V-shaped, to 1/4 depth of cyathia. Staminate flowers
to 30 per cyathium, androphores glabrous, to 0.7 mm. at maturity.
Gynophores glabrous, to 2 mm. long at maturity, exserted, nutant or
upright; calyx a triangular pad of tissue; ovary glabrous; styles spread-
ing, 0.3 mm. long, joined at base, subentire to 1/3 bifid, somewhat
clavate. Capsule glabrous, ovoid, 1.5 mm. long, 2 mm. wide between equator
and base, three-lobed, angles obtuse. Aberrant inflorescences are quite
frequent, producing a thick (to 0.6 mm.) gynophore and sterile capsule to
2.5 mm. long. Seed angular-ovoid, 1 mm. long, to 0.7 mm. diameter at
equator, ventral angle somewhat obscure, others well marked, faces flat,
somewhat pitted, gray-brown.
Key to varieties of C. lecheoides
1. Plant branching from crown of root and throughout length of
stem; branches decumbent ..... a. var. wilsonii.
1. Plant with few stems from ground level, branching mostly in
upper part of stem; branches erect, strict ..... 2.
2. Leaves linear-ovate, much longer than wide, apex
acute ..... b. var. lecheoides.
2. Leaves ovate-elliptic, scarcely longer than
wide, apex obtuse ..... c. var. exumensis.
4a. C. LECHEOIDES (Millsp.) Millsp. var. WILSONII (Millsp.) comb.
nov. C. wilsonii Millsp., Field Mus. Bot. 2: 301. 1909. Type: Castle
Island, Bahamas, 22 Dec. 1907, P. Wilson 7795 (Holotype: F !; Isotype:
Restricted to the whitelands of islands of the Inagua, Turks
and Caicos groups at the southern end of the Bahamas (Map 3 ).
Representative specimens seen: BAHAMAS. Great Inagua, near
lighthouse, 20 Feb. 1963, Dunbar 239 (GH); Inagua, 14 Oct. 1904, Nash
& Taylor 1034 (NY); South Caicos, near East Bay, 17 June 1954, Proctor
8828 (IJ); Turks Islands, Salt Cay, 10 Mar. 1911, Millspaugh & Millspaugh
Much branched from a heavy rootstock, with the branches lying
almost prostrate. Distinguished from var. lecheoides only on the basis
of its many wiry stems, much-branched throughout length.
4b. C. LECHEOIDES (Millsp.) Millsp. var. LECHEOIDES. Euphorbia
lecheoides Millsp., Field Mus. Bot. 2: 163. 1906. Type: Grand Turk,
Turks Islands, Bahamas, 27 Aug.-l Sept. 1905, G. V. Nash & N. Taylor 3888
(Lectotype: F !; Isotype: NY !). Selection of type made here.
Restricted to scrub areas and rocky shores on islands in the
southeast of the Bahamas chain from Watling's Island to the Turks group
Representative specimens seen: BAHAMAS. Watling's Island,
S. W. End, 30 Nov. 1907, Wilson 7347 (NY); Long Cay, 7-17 Dec. 1905,
Brace 4179 (NY); Crooked Island, 9-23 Jan. 1906, Brace 4588 (NY);
Mariguana, 6-8 Dec. 1907, Wilson 7534 (NY); Inagua, 21 Jan. 1962, Dunbar
53 (GH). South Caicos: Northern Hills, 19 June 1954, Proctor 8887 (GH,
IJ); Cockburn Harbor, 28 Mar. 1958, Proctor 17497 (IJ). Between
Commissioner's House and airfield, Grand Turk, 13 June 1954, Proctor
8799 (GH, IJ).
Usually rather few, stout stems from ground level which remain
simple for about half their length. Branching is often, but not invariably,
rather strict. The leaves are smaller and narrower than those of var.
4c. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS (Millsp.) comb.
nov. C. exumensis Millsp., Field Mus. Bot. 2: 301. 1909. Type: Stocking
Island, Great Exuma, Bahamas, N. L. Britton & C. F. Millspaugh 3071
(Holotype: F 1; Isotype: NY !).
Whitelands and coastal sands of islands in the Bahama group from
Eleuthera south to Great Ragged Island (Map 5).
Representative specimens seen: BAHAMAS. Eleuthera, S. E. Point,
27 July 1960, Webster & Williams 10659 (DUKE, IJ, PUR); Little San Salva-
dor, 25-26 Feb. 1907, Britton & Millspaugh 5666 (NY); Cat Island, Bight
and vicinity, 1-6 Mar. 1907, Britton & Millspaugh 5804 (NY); Watling's
1 $ 0 0 f11 12 II 14 .
I: r T
Plate 4. C. LECHEOIDES (Millsp.) Millsp. var. wILSONII (Millsp.) ined.
Isotype: C. wilsonil Millsp., Castle Island, Bahamas, 22 Dec. 1907,
WiIson 7795 (NY).
1905, Nash & Taylor 3888 (NY).
Plate 6. C. LECHEOIDES (Millsp.) Millsp. var. EXUMENSIS (Millsp.) ined.
Isotype: E. exumensis Millsp., Stocking Island, Bahamas, 22-28 Feb. 1905,
Millspaugh 3071 (NY).
Map 3. Distribution of C. lecheoides var. wilsonii
Map 4. Distribution of C. lecheoides var. lecheoides
'-^ 0 Y -r.
Map 5. Distribution of C. lecheoides var. exumensis
Map 6. Distribution of C. cayensis
Island, 12-13 Mar. 1907, Britton & Millspeugh 6143 (NY); Conception
Island, 5 Jan. 1932, Fairchild 16 (GH); Rum Cay, 4 Dec. 1905, Brace
3927 (NY); Long Island, Clarence Town, 16-19 Mar. 1907, Britton &
Millspaugh 6323 (NY); Great Ragged Island, 24-25 Dec. 1907, Wilson
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
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:
Subshrub; ascending to 3 dm. Stems to 3 mm. wide, internodes to
2 cm.; branching sparingly throughout, tomentose, sometimes canescent at
tips, somewhat glabrate, brown or gray. Leaves membranous or sub-
coriaceous; blades ovate, sometimes orbicular or lanceolate, 5-12 x 3-9
mm., base inequilateral, rounded or cuneate, margin entire, slightly
thickened, apex obtuse or rounded, midrib sunken on underside of leaf,
both surfaces sparsely to densely tomentose, or woolly, the surface
minutely white-papillose, yellowish green or pink, often ashen; petiole
to 1 mm. long; stipules united, triangular, to 1 mm. Cyathia solitary
at uppermost nodes; peduncle to I mm. long, involucre broadly obconical,
1.5 2 mm. diameter at mouth, lobes scarcely equalling glands, triangular,
strigose on outside, hairy within, glands broadly transversely elliptic,
0.7 x 0.5 mm., appendages obsolete, fifth gland vestigial or absent,
sinus shallow. Staminate flowers to 30 per cyathium, androphores glab-
rous, to 1 mm. long at maturity, exserted, somewhat nutant; calyx obso-
lete; ovary pilose; styles spreading, 0.3 0.4 mm. long, joined at base,
subentire to 1/3 bifid, somewhat clavate. Capsule strigose, broadly
ovoid, 1.7 mm. long, 1.8 2.1 mm. wide between base and equator, three-
lobed, angles rounded. Seed cuneiform, to 1.2 mm. long, 0.6 mm. wide
above base, strongly four-angled with ventral angle somewhat rounder than
others, faces flat, wrinkled, brown.
Restricted to the coastal whitelands and pine and palmetto-scrub
areas of the northern and western islands of the Bahamas chain (Map 6).
Representative specimens seen: BAHAMAS. Grand Bahama: Freeport,
30 Oct. 1964, McDaniel & Schmiederer 5395 (FSU); Freeport, 8 Aug. 1960,
Webster 10896 (DUKE, PUR); Abaco, 8 Dec. 1904, Brace 1571 (NY). Berry
Islands: Whale Cay, 29 Jan. 1905, Britton & Millspaugh 2178 (NY); Great
Sturrup Cay, 1 Feb. 1905, Britton & Millspaugh 2266 (NY); Great Harbor
Cay,2-3Feb. 1905, Britton & Millspaugh 2334 (NY). Salt Key Bank: North
End, Anguilla Isles, 15-18 May 1909, Wilson 7961 (MO, NY); South End,
Anguilla Isles, 15-18 May 1909, Wilson 8027 (MO, NY); Water Key, 22 May
1909, Wilson 8139 (MO, NY).
Plate 7. CHAMAESYCE CAYENSIS (Millsp.)
Isotype: E. cayensis Millsp., Joulter's
Millspaugh 2295 (NY).
Cays, Bahamas, 11 April 1904,
Recent collections have extended the geographical range, and
also the habitats, from which this species is known. The whitelands,
from which the type collections came, are sandy areas with a very limited
flora, but the collections from Grand Bahama, from the pinelands and from
pine/palm scrub, suggest that the plant is able to tolerate quite strong
competition and shading.
C. bracei Millsp. is no more than a trivial variation of this
species. Of the two specimens cited by Millspaugh with his description,
one is also cited for C. cayensis, and the other differs from the usual
appearance of that species only in the degree of pubescence.
6. CHAMAESYCE PORTERIANA Small, Fl. SE. US., 711. 1903.
Annual or perennating; erect or ascending to 5 dm. Stems to
4 mm. thick at somewhat woody base, internodes to 2 cm.; branching sparse,
mostly from upper nodes or, rarely, much branched, branches at times at
a very acute angle to stem, broom-like, more usually lax, glabrous or
short-tomentose, green, often with a purple cast, or red. Leaves
membranous or somewhat chartaceous; blades elliptic or ovate, in some
suborbicular, in others almost lanceolate, 4-10 x 1-5 mm., base inequi-
lateral, rounded or subcordate, margin entire or rarely somewhat serrate,
apex obtuse to acute, sometimes mucronate, glabrous or sparsely pubescent,
rarely tomentose, green, often suffused purple or red particularly on
underside; petiole 0.5 1 mm. long; stipules distinct or joined at base,
triangular, to 0.4 mm. long, entire or 2-3 parted. Cyathia solitary at
upper nodes; peduncle 0.5 1 mm. long, involucre campanulate, 0.8 1.2
mm. diameter, lobes scarcely equalling glands, triangular, entire or
short-cleft, glabrous or strigose on outside, pilose at mouth within,
glands fleshy, somewhat concave, transversely elliptic, 0.3 0.5 x
0.1 mm., appendages a mere rim, to twice as wide as gland, white, pink,
or dark red, margin entire or crenulate, fifth gland linear, shorter
than lobes or obsolete, sinus broad and shallow. Staminate flowers 8-14
or more per cyathium, androphores glabrous, ca. I mm. long at maturity.
Gynophore glabrous or sparsely pilose, 1-1.5 mm. long at maturity, ex-
serted, nutant; calyx a triangular pad of tissue; ovary glabrous or
strigose; styles spreading, 0.4 mm. long, minutely joined at base, bifid
for 1/2 length. Capsule glabrous or strigose, ovoid, 1.5 mm. long,
1.8 2.1 mm. wide close to base, three-lobed, angles obtuse. Seed
oblong-ellipsoid, 0.7 0.9 mm. long, 0.6 mm. diameter at equator, four-
angled, with ventral angle somewhat rounded, faces flat, obscurely
wrinkled, red-brown or gray.
Key to varieties of C. porteriana
1. Plant pubescent, sometimes glabrate; branching usually rather
lax; leaves membranous, usually deep green; appendages white, margin
entire or crenulate ..... a. var. keyensis.
1. Plant glabrous or, rarely, sparsely short-pubescent; branching
usually strict; leaves parchment-like, often flushed with red or
purple; appendages white or pink, margins entire .....2.
2. Leaves ovate-elliptic, most almost as wide as long, apex obtuse to
rounded; usually rather sparingly branched ..... b. var. porteriana.
2. Leaves linear-elliptic, all except oldest much longer than wide,
apex acute; freely branched, often strict and broom-like
..... c. var. scoparia.
6a. CHAMAESYCE PORTERIANA Small var. KEYENSIS (Small) comb. nov.
Chamaesyce kevensis Small, Torreya 28: 6. 1928. Type: No Name Key,
Monroe County, Florida, 4-5 Feb. 1916, J. K. Small 7439 (Holotype NY 9).
Restricted to coastal scrub and open hammocks of the Florida
Keys, as far north as Lignum Vitae Key (Map 7).
Representative specimens seen: FLORIDA. Monroe County: Lignum
Vitae Key, 13 Dec. 1921, Small, Small & Matthews 10260 (NY); Long Key,
25 Aug. 1949, Woodbury & Jackson s.n. (BUS); Greyhound Key, 21 Nov. 1964,
BLrch 562 (BM, BUS, DUKE, E, F, FLAS, GH, IJ, MO, NY); Ohio Key, 19 Nov.
1964, Burch 538 (FLAS, FSU, MO, NSC, PUR, US); Bahia Honda Key, 9 May
1919, Small, Cuthbert & Mattaus 9139 (FLAS, NY, US); Big Pine Key, 27
June 1960, Webster & Samuel 10216 (DUKE, PUR).
Strongly erect unless growing through dense grass, and evenly
branched, particularly in the upper third of the stem. Distinguished
from the other two varieties by its pubescence and the membranous leaves
which rarely have the suffusion of red or purple that the others show.
6b. CHAMAESYCE PORTERIANA Small var. PORTERIANA. Type: Miami,
Florida, 1877, A. P. Garber s.n. (Holotype: NY !).
Locally frequent in Dade and Monroe counties, Florida. In pine-
land on oolitic limestone from Miami south through the Everglades Keys,
and on the lower Florida Keys. (Map 8).
Representative specimens seen: FLORIDA. Dade County: Pinelands
between Cutler and Black Point Creek, 15 Nov. 1906, Small & Carter 2504
(NY); Pinelands east of Naranja, 14 Jan. 1909, Small & Carter 3066 (NY);
Camp Longview, Everglades, 11 Nov. 1912, Small 3705 (DUKE, FLAS); 6 mi.
west of entrance, Everglades National Park, 17 April 1964, Burch & Ward 298
(BM, BUS, E, F, FLAS, GH, IJ, MO, NSC, NY, PUR, US, USF). Monroe County:
Vacca Keys, 28-29 Jan. 1909, Small & Carter 3048 (NY); Boot Key, 7-12
April 1909, Britton 540 (NY); Big Pine Key, 14 Dec. 1951, Killip 41547
(NY); Key West, 19 Sept. 1954, Killip 44474 (US).
The leaves are broader than those of var. scoparia, and the plant
is usually more sparsely branched. Occasional specimens show sparse
pubescence on the underside of leaves and on the capsules, but the hairs
are shorter and stiffer than is usual in var. keyensis.
6c. CHAMAESYCE PORTERIANA Small var. SCOPARIA (Small) comb. nov.
Chamaesyce scoparia Small, Fl. Florida Keys, 81. 1913. Type: Big Pine
Key, Monroe County, Florida, 17 Nov. 1912, J. K. Small 3819 (Holotype:
Restricted to the lower Florida Keys, usually in pineland on
oolitic limestone, but occasionally in sand or in open hammocks (Map 9).
Representative specimens seen: FLORIDA. Monroe County: Pinelands,
Big Pine Key, 17 Nov. 1912, Small 3770 (DUKE, FLAS, NY); Pinelands, Big
Pine Key, 17 Dec. 1913, Small & Small 5030 (MO, NY); Pinelands, Big Pine
Key, 19 April 1942, Eyles & Eyles 8186 (GH); Pine woods, Big Pine Key, 10
Aug. 1963, Burch 140 (BM, E, F, FLAS, IJ, MO, NY, US); No Name Key, no
date, Curtiss s.n. (GH, NY); Edge of airport, Key West, 26 Dec. 1964,
G. Avery s.n. (FLAS, MO).
There is some intergradation of leaf form between this taxon and
var. porteriana, but the extremes are quite distinct. Plants showing
the strict branching that Small described as "broom-like" when describing
the species have so far only been found on Big Pine Key.
The differences between the type specimens of Small's species do
S'' 2 :I 7 1 Ii, I I II I
Plate 8. CHAMAESYCE PORTERIANA Small var. KEYENSIS (Small) ined.
Holotype: C. keyensis Small, No Name Key, Monroe County, Florida, 4-5
Feb. 1916, Small 7439 (NY).
, ,, ', ,
Plate 9. CHAMAESYCE PORTERIANA Small var. PORTERIANA
Holotype: C. porteriana Small, Miami, Florida, 1877, Garber s.n. (NY).
I Y' -T
~~~ ,-- *\i-. **
*~L~ A 1^-^'Y: -"
2 1 I 7 A 9 1o 11 125 11 It 15
Plate 10. CHAMAESYCE PORTERIANA Small var. SCOPARIA (Small) ined.
Holotype: C. scoparia Small, Big Pine Key, Monroe County, Florida, 17
Nov. 1912, Small 3819 (NY).
Map 7. Distribution of C. porteriana var. keyensis
Map 8. Distribution of C. porteriana var. porteriana
Map 9. Distribution of C. porteriana var. scoparia
-_- _- ---- ---_ _-
1< _____ ___ ..... -_ __
--. ----. -.- --
Map 10. Distribution of C. articulata and C. myrtillifolia
not hold up on examination of a range of collections. Pubescence
seems to be, in general, a very plastic character in the genus, and
can rarely be trusted as a major character for the differentiation of
species. The texture of the leaves and the coloring of the plant
appear to be correlated with habitat, and the branching pattern changes
radically with the accompanying vegetation and the period of time since
the area was exposed to fire. In consideration of these factors it
was deemed more realistic to reduce the species to varietal status until
some clear distinction between them can be demonstrated.
7. CHAMAESYCE MYRTILLIFOLIA (L.) Millsp., Field Mus. Bot. 2:
410. 1916. Euphorbia myrtillifolia L. Syst. Nat. ed. 10, 1048. 1759.
Shrub, erect, to 1 m. Stems to 5 mm. diameter, internodes to
4 cm. long; branching open, throughout length, short-pubescent, brown.
Leaves membranous; blades elliptic or obovate, 7-15 x 5-10 mm., base
oblique, obtuse or cuneate, margin serrate, often only in upper half,
apex emarginate, obscurely mucronate, midrib prominent on lower surface,
upper surface glabrous or sparsely pilose, lower pilose-pubescent, yellow-
green; petiole to 1 mm. long, short-pubescent; stipules joined for half
length, free part narrow, ciliate or toothed. Cyathia solitary at upper
nodes; peduncle to 2 mm. long, involucre obconical, to 1.5 mm. diameter
at mouth, lobes equalling glands, deltoid, toothed, pilose on outside and
within, glands transversely elliptic, 0.5 x 0.2 mm., dark brown, appendages
longer than glands and as wide, margin somewhat dentate, creamy-yellow,
fifth gland linear, short or obsolete, sinus shallow. Staminate flowers
18-30 per cyathium, androphores glabrous, to 1.5 mm. long at maturity.
Gynophore short-pubescent, to 3 mm. long at maturity, exserted, upright;
A I I II I
Plate II. CHAMAESYCE MYRTILLIFOLIA (L.) Millsp.
E. myrtillifolia L., Gordon Town, St. Andrew, Jamaica, 8 Dec. 1954,
Proctor 9595 (NY).
calyx a pad of tissue or obsolete; ovary sparsely pilose; styles spread-
ing, to I mm. long, joined at base, bifid for one-third length. Capsule
sparsely pubescent, glabrate, ovoid with truncate or cordiform base, 2.2
mm. long, 2.3 mm. wide above base, broadly three-lobed, angles rounded.
Seed long-conical, to 1.8 mm. long, triangular with ventral angle obso-
lete, others acute, dorsal faces convex, strongly wrinkled, base truncate,
Reported only from the Gordon Town and Guava Ridge area of the
Blue Mountains of Jamaica (Map 10).
Specimens seen: JAMAICA. St. Andrew: Near Gordon Town, 23 May
1894, Harris 5196 (NY); Valley of Hog Hole River above Gordon Town,
8 Dec. 1954, Proctor 9595 (IJ, NY); 1/4 mile northeast of Gordon Town,
20 Mar. 1956, Proctor 11874 (GH, IJ).
The affinities of this species are with the shrubby C. articulata,
but it differs from that species in.being smaller and having elliptic
leaves which are pilose on the under surface. The involucre of C. myr-
tillifolia is smaller, with less prominent glandular appendages, and the
style and other floral parts are reduced proportionally.
8. CHAMAESYCE NUTANS (Lag.) Small, Fl. SE. US. 712. 1903.
Euphorbia nutans Lag., Gen. et Spec. Pl. 17. 1816. Type: "Habit.[at]
in N.[ova] H.[ispania]. Perhaps at Madrid ." (Wheeler, 1941).
E. preslii Guss., Fl. Sic. Prod. 1: 539. 1827. Type: Palermo,
Italia, Todaro (fragment F seen by Wheeler). C. preslii (Guss.) Arthur,
Torreya 11: 260. 1912.
E. hypericifolia L. var. communis Engelm. in Emory, U.S. & Mex.
Bound. Surv. 2: 188. 1859. Type: New Mexico, 1851-52, C. Wright 1842
(fragment F seen by Wheeler).
E. hypericifolia L. sensu Michaux, Fl. Bor.-Am., 1803; Torrey,
Fl. State New York, 1843.
E. maculata L. sensu Wheeler, Contr. Gray Herb. 127: 74. 1939.
Annual; erect or ascending to 8 dm. Stem to 4 mm. thick at some-
what woody base, internodes to 3 cm. long; branched mostly from upper
nodes, young tips and a line on older stem densely tomentose, glabrate,
straw-colored or darker brown. Leaves membranous; blades ovate-
lanceolate, often somewhat oblong or falcate, 10-30 x 6-10 mm., base
inequilateral, rounded, margin serrate, apex acute to obtuse, glabrous
or sparsely long-pilose on upper surface, more generally long-pilose
below, both surfaces mid-green often with a red spot on upper surface
or the whole suffused with red; petiole 1-2 mm. long; stipules joined or
early distinct near tip of shoot, triangular to I mm. long, margin
toothed and ciliate, thin textured, brown. Cyathia borne singly in
lateral and terminal short-stalked compound dichasia; peduncle 0.5 2
mm. long, involucre campanulate, 0.7 I mm. diameter, lobes triangular
exceeding glands, subentire or deeply laciniate, glabrous outside,
slightly hairy within, glands broadly transversely elliptic, 0.1 0.3
mm. long, appendages obsolete or to three times width of gland, white
or pink, fifth gland linear, shorter than lobes, sinus shallow. Staminate
flowers 5-14 per cyathium, androphores glabrous, 1 mm. long at maturity.
Gynophore glabrous, to 2 mm. long at maturity, exserted, upright or nutant;
calyx obsolete; ovary glabrous; styles spreading, 0.4 1 mm. long, minutely
joined at base, bifid for 1/3 1/2 length. Capsule glabrous, broadly
ovoid with somewhat flattened base, 1.8 2.3 mm. long, 2 2.3 mm. diameter
! r '-"
I I ., I. Il I:.
Plate 12. CHAMAESYCE NUTANS (Lag.) Small
C. nutans (Lag.) Small, between Marianna and Campbellton, Jackson County,
Florida, 7 Nov. 1964, Godfrey 64932 (FLAS).
below equator, strongly three-lobed, angles subacute. Seed oblong-ovoid,
to 1.3 mm. long, 0.8 mm. wide at equator, ventral angle rounded and
obscure, others well marked, faces convex, rippled, black, often with
A common weed of the eastern United States reaching its southern
limit in north Florida. It is possible that it was present as a weed
of cultivated ground in Puerto Rico and many of the Caribbean islands
as reported by Britton and Wilson (1924), but almost all of the speci-
mens in NY and F on which these records might have been based prove to
be misidentifications of plants of E. hyssopifolia L. and E. hypericifolia
L. The collections by Fredholm and by Hitchcock from central Florida are
correctly named, but no more recent records have been obtained from this
area in spite of heavy collecting, and it is doubtful that the species is
present as more than a short-lived, casual introduction. (Map 11).
Specimens seen: FLORIDA. Alachua: June-July 1898, Hitchcock 1697
(F). Brevard: Okeechobee region, 17 Sept. 1903, Fredholm 6014 (GH, NY).
Citrus: June-July 1898, Hitchcock 1699 (F, MO). Gadsden: Jim Woodruff
Dam, Chattahoochee, 19 Oct. 1963, Burch 207 (FLAS, IJ, US); River Junction,
7 Aug. 1927, Wiegand & Manning 1818 (GH); Chattahoochee, 7 Aug. 1927,
Wiegand & Manning 1819 (GH); Chattahoochee, 8 Sept. 1931, West s.n. (FLAS).
Jackson: Marianna, 19 Oct. 1963, Burch 210 (FLAS); near Cottondale, 26 Oct.
1964, Burch 504 (FLAS, MO, NCU, NY); between Marianna and Campbelltown,
7 Nov. 1964, Godfrey 64932 (FLAS, FSU); Marianna State Park, 15 Sept. 1960,
Mitchell 839 (FSU); Marianna Caverns State Park, 19 Nov. 1960, Mitchell
903 (FSU). Leon: Tallahassee, 11 Sept. 1942, Henry & Baggs s.n. (FLAS);
18 Sept. 1942, Kurz s.n. (FLAS); Tallahassee, 2-3 Sept. 1895, Nash 2519
(F, GH, NY). Levy: June-July 1898, Hitchcock 1698 (F). Wakulla: Newport,
13 Sept. 1964, Godfrey 64618 (FLAS). Walton: Northwest section, 9 July
1964, Hume s.n. (FLAS). Washington: west side of Careyville, 26 Oct. 1964,
Burch 499 (FLAS). Extra-territorial specimens: BERMUDA. North Shore, 31
Aug.-20 Sept. 1905, Brown & Britton 8 (F, NY); Devonshire, 31 Aug.-20 Sept.
1905, Brown & Britton 79 (F, NY); Devonshire, 19 Aug. 1963, Manuel 265 (A).
This species bears a strong resemblance to C. hypericifolia (L.)
Millsp., C. hyssopifolia (L.) Small and C. lasiocarpa (Klotzsch) Arth.,
but may be distinguished by its stem pubescence, limited to the tips and
lines at the side of older parts; by the cyathia carried in a leafy
dichasium; by the large capsules and by the black, plump seed with a
The application by Wheeler (1939) of the name E. maculata L. to
this species, rather than to the prostrate plant for which it has been
widely used, deserves comment. His case, based on the selection of a
specimen from Herb. Linn. as the type, was never proved, since there
was some doubt about the identification that Linnaeus made for this sheet.
The description of E. maculata in Species Plantarum is placed among the low-
growing rather than the erect species, suggesting that Linnaeus intended
it for a prostrate plant and that the traditional application is correct.
The use of the name E. hypericifolia L. for this species may be
traced to annotations on the Linnaean specimens made by Sir J. E. Smith
during the years when he owned the herbarium. Michaux (1803) may have
examined the material and arrived at the decision independently, but the
use in this fashion was confirmed by correspondence between Smith and
Torrey when his Flora of the State of New York (1843) was in preparation.
P-~ ~ ~
Map 1!. Distribution of C. nutans
- ...-- ------ ------ -
Map 12. Distribution of C. hypericifolia
---------I ---- ------0
The basis for Smith's actions was sound in that the description of
E. hypericifolia in Linnaeus' early works was written from this northern
plant. The specific limits were widened to include a tropical plant,
however, before the publication of Species Plantarum, and since the name
has been used most widely for this entity, the interests of stability of
nomenclature are best served by maintaining this application. Burch (1965)
deals more fully with the questions relating to the name of this species.
9. CHAMAESYCE HYPERICIFOLIA (L.) Millsp. Field Mus. Bot. 2: 302.
1909. Euphorbia hypericifolia L., Sp. Pl. ed. 1, 454. 1753. Type:
Sloane Hist. 1, t. 126. (lectotype, designated by Burch, Rhodora 67. 1965,
E. pilulifera L. sensu Wheeler, Contr. Gray Herb. 127: 76. 1939.
(But see disclaimer, loc. cit. 78).
C. qlomerifera Millsp. Field Mus. Bot. 2: 377. 1913. Type: El
Rancho, Dept. Jalapa, Guatemala, 20 Jan. 1908, W. A. Kellerman 8053 (F i).
E. glomerifera (Millsp.) L. C. Wheeler, Contr. Gray Herb. 127: 78. 1939.
Annual or perennating in tropics; erect to ascending, or sprawling
to 4 dm. Stem to 3 mm. diameter at somewhat woody base, internodes to
3 cm. long; branching throughout length particularly in upper part of stem,
glabrous, tan to dark brown. Leaves membranous, blades ovate-elliptic to
elliptic-obovate, at times somewhat falcate, 15-35 x 8-14 mm., base inequi-
lateral, rounded or cuneate, margin serrate, apex acute, glabrous, green,
sometimes purple-spotted above and lighter green on underside; petiole
to I mm. long; stipules joined, sheathing, I 1.5 mm. long, margin serrate,
inner surface pubescent at least at apex, light-brown, or, at upper nodes,
separate, to 0.5 mm. long, entire or deeply 2-4 cleft. Cyathia in
short-stalked, lateral and terminal, congested dichasia forming glomerules
with a few basal leaves; involucre short-peduncled with stipule-like
bracts at junction with pedicel, narrow campanulate or more flaring, to
0.8 mm. diameter at mouth, lobes equalling or exceeding glands, triangular,
toothed or cleft, glabrous on outside, sparsely long-pilose within, glands
subcircular to 0.1 mm. diameter, appendages obsolete or prominent, to
1 mm. wide, entire, white or pink, fifth gland obsolete, sinus shallow.
Staminate flowers 10-12 or rarely only 4 per cyathium, androphores glabrous,
to 0.6 mm. long at maturity. Gynophore glabrous, to 1 mm. long at matu-
rity, exserted, nutant; calyx a triangular pad of tissue; ovary glabrous;
styles 0.3 mm. long, spreading, minutely joined at base, bifid for 1/2
length. Capsule glabrous, subspherical, to 1.2 mm. diameter, roundly
three-lobed, angles rounded. Seed ovoid, to 0.8 mm. long, 0.5 mm. wide,
strongly four-sided, ventral angle more obtuse than others, faces flat
or convex, wrinkled, light brown.
A frequent plant of waste places in Old and New World tropics and
subtropics. Common in the southern part of Florida with isolated collec-
tions as far north as Sapelo Island, Georgia, and present throughout the
Greater and Lesser Antilles (Map 12).
Representative specimens seen: FLORIDA. Alachua: University of
Florida campus, Gainesville, 30 Oct. 1964, Burch 521 (FLAS). Broward:
Fort Lauderdale, 19-25 Nov. 1903, Small & Carter 644 (NY). Collier: road
to Marco Island, 15 Apr. 1954, West, Arnold & Cooley 147 (FLAS). Dade:
vicinity of Homestead, 22 Nov. 1964, Godfrey 65583 (FLAS, FSU); Buena
Vista, I Jan. 1930, Moldenke 330 (DUKE, NY); Miami, 14 May 1904, Tracy
9127 (F, MO, NY). Glades: Caloosahatchee River bank, 7 Nov. 1964,
a I a 3 t 6 7 p u a ,J ll II 1 II IS
Plate 13. CHAMAESYCE HYPERICIFOLIA (L.) Millsp.
C. hypericifolia (L.) Millsp., University of Florida campus, Gainesville,
Alachua County, Florida, 30 Oct. 1964, Burch 521 (FLAS).
Hodgson 230 (FLAS). Highlands: Rd. 25, south of Sebring, 22 June 1948,
Garrett s.n. (FLAS). Lee: Sanibel Island, 13 Apr. 1954, West & Cooley
2630 (USF). Manatee: U.S. 41 & U.S. 19, 29 Nov. 1962, Ward & Burch 3163
(FLAS). Monroe: Upper Mateconbe Key, June, ?? Curtiss 2486 (F, MO);
No Name Key, 21 Mar. 1898, Pollard, Collins & Morris 126 (F, NY); Key
Vaca, 3 Sept. 1954, Thorne 15177 (IJ); Key Largo, north end, 27 June 1960,
Webster & Samuel 10239 (DUKE, PUR). Palm Beach: Jupiter Inlet, west of
DuBoise Park, 27 Aug. 1964, Christensen RC-44a (FLAS, FSU). Pinellas:
Don Caesar Hotel, St. Petersburg Beach, 24 May 1964, Taylor s.n. (FLAS).
Polk: Haines City, 25 June 1963, Conard s.n. (FSU). Sarasota: Laurel,
21 Nov. 1964, Godfrey 65273 (FLAS, FSU). Sumter: Wildwood exit, 1-75,
18 Nov. 1964, Burch 524 (FLAS). Volusia: Pierson, 17 Sept. 1943, West
& Arnold s.n. (FLAS). BAHAMAS. Great Bahama, 5-13 Feb. 1905, Britton &
Millspaugh 2476 (F, NY); North Bimini, 16 Apr. 1904, Millspaugh 2399 (F,
NY). Andros: Mastic Point, 19-28 Mar. 1907, Brace 7087 (F, NY); Nicholl's
Town, 4-5 Feb. 1910, Small & Carter 8937 (F, NY). New Providence, Nassau,
Nov. 1890, Hitchcock s.n. (F, MO). Eleuthera: Gregory Town, 6 July 1903,
Coker 371 (NY); Governor's Harbor, 14 Nov. 1890, Hitchcock s.n. (F). Cat
Island, the Bight & vicinity, 1-6 Mar. 1907, Britton & Millspaugh 5816
(F, NY); Watling's Island, Cockburn Town, 12-13 Mar. 1907, Britton &
Millspaugh 6053 (F, NY); Great Exuma, 22-28 Feb. 1905, Britton & Millspaugh
2945 (F, NY); Long Cay, 7-17 Dec. 1905, Brace 4036 (F, NY); Inagua, 3 Dec.
1890, Hitchcock s.n. (F, MO). CUBA. Camaguey: Atalaya, 20 Mar. 1909,
Shafer 983 (F, NY). Havana: Havana, 5 May 1905, Curtiss 743 (F, MO, NY).
Oriented: Monte Verdi, Jan.-July 1859, Wright 1422 (F, MO, NY). Santa Clara:
Cienequita, Cienfuegos, 2 May 1895, Combs 4 (F, MO, NY); Limones, Soledad,
Cienfuegos, 27 Sept. 1924, Jack 6334 (MO). GRAND CAYMAN. Georgetown,
2 June 1963, Crosby, Hespenheide, & Anderson 50 (DUKE, UCWI); Spot Bay,
13-14 Feb. 1899, Millspaugh 1304 (F). JAMAICA. Portland: Kildare, 21
Aug. 1962, Adams 11555 (UCWI); Port Antonio, 4 Jan. 1906, Wight 2 (F,
NY). St. Andrew: Mount Pleasant, Stony Hill, 21 July 1912, Harris
11134 (F, NY, UCWI); Long Mountain, June 1953, Robbins 434 (UCWI).
St. Ann: Mosely Hill Cave, 1 mile west of Blackstonedge, 12 Dec. 1952,
Proctor 7474 (IJ). St. James: Ironshore, May 1953, Asprey 432 (UCWI).
Trelawny: Cowie Park near Troy, 19 Oct. 1917, Harris 12646 (F, MO, NY,
UCWI). Westmoreland: Negril, 29 Mar. 1962, Adams 10980 (UCWI). HAITI.
Jeremie, 13 July 1941, Seibert 1725 (MO); Port au Prince, 21-23 Feb. 1920,
Leonard 2778 (NY); St. Michel de 1'Abalaye, 1927, Ekman 8394 (F, IJ).
DOMINICAN REPUBLIC. Barahona, June 1910, Fuertes 255 (F, MO, NY); San
Pedro de Macoris, 26 Mar. 1913, Rose, Fitch & Russell 4154 (NY); Santo
Domingo City, 9-10 Mar. 1913, Rose, Fitch & Russell 3769 (NY). PUERTO
RICO. Camuey, 14 June-22 July 1901, Underwood & Griggs 204 (NY);
Fajardo Light House, 17 Apr. 1899, Heller 1179 (F, NY); Guanica, 22
Jan. 1899, Millspaugh 753 (F); Guayama, May 1874, Kuntze 554a (NY);
Penuelas, 16 July 1886, Sintenis 4823 (MO, NY); Rio Piedras, 9 Apr. 1912,
Johnston 216 (NY); San Juan, Dec. 1913, Hiorani s.n. (NY). LESSER ANTILLES.
St. Thomas: Charlotte Amalie, 17-18 Jan. 1899, Millspaugh 382 (F); Forts
Square, Jan. 1877, Eggers s.n. (MO, NY). St. Croix: Bassin Yard, 15 Nov.
1895, Ricksecker 75 (F, MO, NY); Peter's Rest, 2 June 1924, Thompson
772 (NY). Tortola, 18 Sept. 1918, Fishlock 144 (NY). St. Kitts, 8 Sept.-
5 Oct. 1901, Britton & Cowell (NY). Guadeloupe, Pointe-a-Pitre, 19 May
1936, Stehle 1190 (NY); Dominica, Soufriere, 1903, Lloyd 490 (NY).
Martinique: Case Pilote, 1879, Duss 488 (F, MO, NY); Ste. Anne village,
29 July 1939, Egler 39-208 (NY). Barbados: Bathsheba, Apr.-June, 1895,
Waby 101 (F); Waterford, St. Michael, 15 Aug. 1906, Dash 166 (F).
Grenada, St. George's, 29 Sept. 1906, Broadway s.n. (F). Extra-territorial
specimens: BERMUDA. St. David's Isle, 22 May-6 June 1914, Brown,
Britton & Bisset 2093 (F, NY); neglected field, 7 July 1921, Degener 1003
(MO). GEORGIA. S.W. section, Sapelo Island, 13 Oct. 1956, Duncan 20624
(DUKE, NCU, NSC).
The Linnaean name E. hypericifolia has been applied to this species
and to the one known here as C. nutans. Both elements were represented
in the diagnosis and in the synonymy of the original publication in Species
Plantarum, and the choice of this species to carry the name was made on
the basis of its widespread application in this way dating from Miller's
Gardener's Dictionary edition 8 (1768). This matter, and Wheeler's appli-
cation of the name to the plant known here as C. lasiocarpa, are discussed
in more detail by Burch (1965).
The use of the name E. pilulifera L. for this species cannot be
accepted if the sheet (630-8) in the Linnaean Herbarium bearing this name
is allowed any status as type material. The identity of the specimen is
not clear, but the small part of a seed visible in one capsule showed none
of the surface markings characteristic of seed of C. hypericifolia, and
the general aspect of the plant was closer to that of C. hirta.
10. CHAMAESYCE HYSSOPIFOLIA (L.) Small,J. N.Y. Bot. Gard. 3: 429.
1905. Euphorbia hyssopifolia L. Syst. Nat. ed 10, 1048. 1759. Type: in
Herb. Linn. sheet 630-9 Anisophyllum hyssopifolium (L.) Haw., Syn. PI.
Succ., 161. 1812. E. hypericifolia L."P hyssopifolia L." Griseb. Fl. Br.
W. Ind. Isl., 54. 1859. E. brasiliensis Lam. var. hyssopifolia Boiss.
in DC. Prod. 15(2): 24. 1862.
E. brasiliensis Lam., Encyc. 2: 423. 1786. Type: Mus. Nat.,
Paris, if extant. C. brasiliensis (Lam.) Small, Fl. SE, US., 712. 1903.
C. nirurioides Millsp., Field Mus. Bot. 2: 394. 1914. Type: Malvern,
Santa Cruz Mountains, Jamaica, 5 Sept. 1907, Britton 1186 holotypee: F !,
isotype: NY !). E. nirurioides (Millsp.) Fawcett & Rendle, Fl. Jam., 4(2):
C. jenningsii Millsp. in Britton, Bull. Torr. Bot. Club 43; 465.
1916. Type: near Vivijagua, Isle of Pines, Cuba, 7 May 1910, Jennings 621
(Holotype: Herb. Carnegie Mus., portion of holotype F !; Isotype: NY 1).
Annual, occasionally perennating; erect or ascending or rarely
decumbent, to 6 dm. Stem to 4 mm. thick at somewhat woody base, internodes
to 3 cm. long; branched at all levels but most in upper part of plant,
glabrous or sparingly long hairy near base, rarely strongly short-pubescent
at lower nodes particularly in plants from more tropical areas, straw-
colored or darker brown. Leaves membranous; blades very variable, linear-
lanceolate, 15-35 x 3-6 mm., with subcordate inequilateral base, margin
serrate in upper half, apex acute, to ovate-elliptic, occasionally some-
what falcate, 10-30 x 6-10 mm., with rounded to subcordate inequilateral
base, margin generally serrate, apex.obtuse, glabrous or sparsely long
pilose, on shoots near base of plant, both surfaces mid-green; petiole 1-2
mm. long; stipules joined or nearly free at upper nodes, almost obsolete to
deltoid, 0.8 mm. long, margin short-fringed. Cyathia borne singly,
terminal and axilary in short-stalked, diffuse or compact, leafy dichasia;
peduncle short, involucre commonly narrowly tubular, 0.7 1 mm. diameter,
sometimes flaring-campanulate, lobes triangular, deep cleft, exceeding
glands, glabrous outside, sparsely pilose within, glands transversely
elliptic or almost circular, 0.1 0.3 mm. long, appendages obsolete to
prominent, elliptic, twice as long and three times as wide as gland,
white or pink, fifth gland obsolete or tiny, linear, sinus shallow.
Staminate flowers 9-20 per cyathium, androphores glabrous, 0.6 1 mm.
long. Gynophore glabrous, ca. 1 mm. long at maturity, exserted, usually
nutant; calyx a triangular pad; ovary glabrous; styles upright or spread-
ing, to 0.4 mm. minutely joined at base, bifid for 1/3 1/2 length.
Capsule glabrous, truncate-cuneiform to broadly ovoid, 1.5 2 mm. long,
1.6 1.8 mm. wide just above base, strongly three-lobed, angles subacute.
Seed oblong-ovate, to 1 mm. long, 0.5 mm. wide, four-angled with ventral
angle less acute than others, faces flat, marked with transverse ridges,
a central longitudinal ridge sometimes on dorsal faces, brown or black,
angles sometimes light.
Widespread throughout Florida and sporadically to South Carolina,
common in all parts of the Caribbean except the Bahamas, where the reports
for New Providence and Eleuthera (Britton and Millspaugh, 1920) are
probably based on collections by Britton and Coker (NY !) which prove to
be misidentifications of plants of C. hypericifolia (L.) Millsp. (Map 13).
Representative specimens seen: FLORIDA. Alachua: Pineland, Gaines-
ville, 15 June 1910, Hood 24465 (FLAS, MO); University of Florida campus,
Gainesville, 10 Sept. 1956, Jackson 707 (FLAS). Baker: Olustee National
Forest, 28 Nov. 1955, Kral 1572 (FSU). Brevard: Melbourne, 22 Aug. 1958,
Kral 7971 (FLAS, GH, NY, USF). Broward: Pinelands, Ft. Lauderdale, 19 &
25 Nov. 1903, Small & Carter 1141 (F, NY); U.S. 27, north of Andytown,
7T I10 II 12 13 14 IA
r 1 I
I'. I F
Plate 14. CHAMAESYCE HYSSOPIFOLIA (L.) Small
C. hyssopifolia (L.) Small, Fairchild Tropical Gardens, Dade County,
Florida, 31 December 1962, Burch 52 (FLAS).
." I eT'I sp.r.cv.
es~ c j
9. c! ii..i ii' ii]j~l I'ii ~iLL
Plate 15. CHAMAESYCE HYSSOPIFOLIA (L.) Small
Isotype: C. nirurioides Millsp., Malvern, Santa Cruz Mountains, Jamaica;
5 Sept. 1907, Britton 1186 (NY).
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PLANTS OF ISI.I: o01 PINE:
Plate 16. CHAMAESYCE HYSSOPIFOLIA (L.) Small
Isotype: C. ienninqsii Millsp. in Britton, near Vivijagua, Isle of Pines
Cuba, 7 May 1910, Jennings 621 (NY).
26 Jan. 1963, Ward & Burch 3338 (FLAS). Calhoun: N.W. of Blountstown,
25 Oct. 1964, Burch 506 (FLAS). Charlotte: Charlotte Harbor, 21 Nov.
1964, Godfrey 65360 (FLAS); Peace River, 13 Aug. 1963, Burch 185 (FLAS).
Citrus: Dunnellon, 16 June 1958, Kral 6901 (FLAS, FSU, USF). Collier:
Immokalee, 26 Sept. 1964, Lakela 27492 (USF). Dade: Buena Vista, 1 Jan.
1930, Moldenke (DUKE, F, MO, NY); Miami-Kendall Station, 5 Nov. 1906,
Small & Carter 2521 (NY). Ouval: Jacksonville, 17 Nov. 1929, Moldenke
5232 (NY). Franklin: Apalachicola, 2 Nov. 1875, Chapman 3844a (MO, NY).
Gadsden: Greensboro, 9 July 1955, Godfrey 53614 (DUKE, FSU, NY). Hendry:
La Belle, 19 Apr. 1930, Moldenke 5911 (NY). Hernando: Brooksville,
3 Sept. 1959, Ray 9461 (USF). Highlands: Childs, 3 Mar. 1945, Brass
14744 (US); Sebring, 25 Apr. 1960, Ray, Gleason & Eaton 9728 (USF).
Hillsborough: Tampa, 20 Mar. 1923, Churchill 546 (MO, US); Oldsmar,
12 Aug. 1960, Ray, Lakela & Patman 10006 (USF). Jackson: between Mari-
anna & Campbellton, 7 Nov. 1964, Godfrey 64933 (FSU). Lake: Eustis,
16-30 June 1894, Nash 1076 (F, MO, NY). Lee: Ft. Myers, July-Aug. 1900,
Hitchcock 324 (F, MO, NY); 5 miles east of Ft. Myers Beach, 1 Aug. 1958,
Kral 7539 (GH, USF). Leon: Tallahassee, 30 June 1955, Godfrey 53571
(DUKE, FSU, NY). Marion: Silver Springs, 1 Oct. 1963, Godfrey 63145 (FSU).
Manatee: Bradenton, 13 June 1916, Cuthbert 1312 (NY); Cortez, 19 July 1958,
Jackson 732 (FLAS). Martin: Stuart, 3 Sept. 1944, West & Arnold s.n.
(FLAS). Monroe: Key Largo, 26 Jan. 1963, Ward & Burch 3332 (FLAS). Nassau:
Fort Clinch, 12 Oct. 1963, Burch 195 (FLAS). Okaloosa: Ft. Walton Beach,
24 Oct. 1964, Burch 470 (FLAS). Okeechobee, 27 Dec. 1964, Burch 589 (FLAS).
Orange: Orlando, 21 June 1900, Curtiss 6670 (MO, NY). Osceola: St. Cloud,
27 Feb. 1952, Cooley 1032 (USF). Palm Beach: between Palm Beach & Lake
Worth, 7 Aug. 1963, Burch 119 (FLAS). Pasco: Land O'Lakes, 10 July
1958, (FLAS, GH, NY). Pinellas: St. Petersburg, 23 Sept. 1907, Deam
2758 (MO). Polk: Lake Hamilton, 30 Sept. 1962, Conard s.n. (FLAS).
Putnam: Welaka, 5 Aug. 1940, Laessle s.n. (FLAS). St. Lucie: Ft.
Pierce, 7 Aug. 1963, Burch 122 (FLAS). St. Johns: Ponte Vedra, 11
Oct. 1964, Burch 437 (FLAS). Sarasota: Osprey, 10 Mar. 1904, Smith
712 (DUKE). Sumter: Withlacoochee River bridge, 8 Nov. 1946, West &
Arnold s.n. (FLAS). Suwannee: Live Oak, 4 Oct. 1964, Elam s.n. (FLAS).
Volusia: Daytona Beach Shores, 3 Aug. 1962, Ray 11159 (USF). Wakulla:
3 Nov. 1964, Godfrey 64887 (FLAS, FSU). Washington: Chipley, 31 July
1954, Ford & Arnold 3860 (FLAS). CUBA. Camaguey: La Gloria, Feb. 1909,
Shafer 285 (F, NY). Cienfuegos: Colonia Limones, Ingenio Soledad, 24 Jan.
1903, Pringle 59 (F, MO, NY); 2 May 1895, Combs 3 (F, MO, NY). Havana:
Guanabacoa, 27 Jan. 1905, Curtiss 618 (F, MO, NY); Havana, Apr. 1903,
Shafer 91 (F). Isle of Pines: Nueva'Gerona, 19 Dec. 1903, Curtiss 244
(F, NY). Oriented: Bayate, 18 July 1914, Ekman 2012 (NY). Pinar del Rio:
Herradura, 26-30 Aug. 1910, Britton, Britton, Earle & Gager 6332 (F, NY);
between Vinales and Pinar del Rio, 16 June 1953, Webster 4670 (PUR). San
Pedro: banks of Ariguanaba lagoon, 5 Apr. 1923, Leon 11417 (NY). Santa
Clara: Hoyo de Manicaragua, 26-28 Feb. 1910, Britton, Britton & Wilson
4660 (F, NY); Rio Hondo Plains, 5 May 1903, Cook & Doyle 528 (F). JAMAICA.
Clarendon: Alley, 11 Feb. 1947, von der Porten s.n. (IJ); Kupuis, 23 June
1955, Proctor 10330 (IJ). Hanover: Negril to Bloody Bay, 12 Sept. 1962,
Adams 11663 (UCWI). Manchester: Lititz, 5 Nov. 1961, Adams 9864 (UCWI).
Portland: Port Antonio, Dec. 1890, Hitchcock s.n. (MO). St. Andrew: Con-
stant Spring to Bardowie, 2 Aug. 1915, Harris 12112 (F, MO, NY, UCWI);
Hope, 18 Aug. 1897, Harris 6803 (F, NY, UCWI); road to Constitution
Hill, 20 Oct. 1905, Harris 9035 (F, NY, UCWI); University of West Indies
campus, 4 June 1963, Crosby, Hespenheide & Anderson 60 (F). St. Catherine:
Bog Walk, 17 Dec. 1890, Hitchcock s.n. (MO). St. Thomas: Morant Bay, 6
Dec. 1959, Adams 5678 (UCWI); Port Morant, 24 Dec. 1890, Hitchcock s.n.
(MO). Westmoreland: Bluefields & vicinity, 6-7 Mar. 1908, Britton &
Hollick 1943 (F, NY). HAITI. Gonaives, Dec. 1899, Buch 278 (IJ); La
Gonave, 9 Aug. 1927, Ekman 8862 (A); Mt. La Porte, near Pilate, 21 Aug.
1903, Nash 628 (NY); Pointe-a-pitre, 12 July 1936, Stehle 497 (NY); Port
de Paix, 9 Jan. 1929, Leonard & Leonard 13543 (NY); St. Michel de l'Atalaye,
1 Jan. 1926, Leonard 8501 (MO); San Michel, 5 Aug. 1905, Nash & Taylor
1417 (NY). DOMINICAN REPUBLIC. Barahona, Feb. 1910, Fuertes 149 (F, NY).
San Juan: Rio Arriba, 9-14 Sept. 1946, Howard & Howard 8821 (GH, NY).
Santo Domingo: Villa Allagracia, 19 Jan. 1929, Ekman H11229 (PUR). PUERTO
RICO. Guayanilla-Tallahoa, 13 Mar. 1913, Shafer 1976 (F, MO); Martin
Pena Station, 20 Feb. 1900, Heller 4665 (F, MO); Punta Arenas, 15 Feb.
1914, Shafer 2896 (F, NY); Rio Piedras, Station Grounds, 2 Oct. 1937,
Otero 123 (MO); Utuado, 15-20 Mar. 1906, Britton & Cowell 409 (F, NY).
LESSER ANTILLES. Charlotte Amalie, 17-18 Jan. 1899, Millspaugh 398 (F);
Antigua, Belvidere, 15 Aug. 1937, Box 964 (MO); Montserrat, 5 Aug. 1939,
Friend 51 (NY); Guadeloupe, Manive, 1893, Duss 2920 (NY); Martinique,
Case-Pilote, 1901, Duss 4641 (F, NY); St. Lucia, 7 Sept. 1935, Potter
5508 (GH); St. Vincent, Kingstown, 2 Dec. 1945, Beard 1397 (MO, NY);
Grenada, St. George's, 30 Oct.-ll Dec. 1957, Proctor 16812 (IJ).
C. hyssopifolia (L.) Small may be distinguished from similar species
by its combination of usually glabrous stems, cyathia in a leafy dichasium,
-0- -- --
Map 13. Distribution of C. hvssopifolia
Map 14. Distribution of C. lasiocarpa
large capsules and seed with prominent transverse ridges on at least
the tangential faces.
The specimen in the Linnaean Herbarium is of a plant with linear
leaves and rather few cyathia in a diffuse dichasium. This form occurs
in most of the islands of the Greater Antilles, and is particularly well-
marked on Hispaniola, but grades into the broader leaf form (often on the
same plant), and the type specimen represents only the extreme leaf shape
of this very variable species.
11. CHAMAESYCE LASIOCARPA (Klotzsch) Arthur, Torreya 11: 260.
1911. Euphorbia lasiocarpa Klotzsch, Nov. Act. Nat. Cur. 19, supp. 1:
414. 1843. Type: Bot. Mus., Berlin, if extant. E. hypericifolia L.
" lasiocarpa K1." Griseb., Fl. Brit. W. I. Isl. 54. 1859. (excl. syn.
E. hypericifolia L. sensu Wheeler, Contr. Gray Herb. 127: 73. 1939.
Annual; erect or ascending to 1 m. Stem to 5 mm. thick at somewhat
woody base, internodes to 3 cm. long; branched throughout, densely tomen-
tose, straw-colored becoming darker with age. Leaves membranous; blades
ovate-lanceolate to obovate, somewhat falcate, 15-40 x 8-20 mm., base
inequilateral, rounded to obtuse, margin sharp-serrate, apex acute to ob-
tuse, densely gray pubescent, rarely glabrate, green; petioles 2 to 3 mm.
long; stipules joined, deltoid to 1 mm., toothed or ciliate, sometimes
deeply bifid near tips of shoot. Cythia solitary in lateral and terminal
short-stalked compound dichasia; peduncle ca. 1 mm. long, involucre
campanulate, 1.2 mm. diameter at mouth, lobes triangular equalling glands,
toothed or cleft, densely pubescent on outside, pilose within, glands trans-
versely elliptic 0.3 mm. x 0.1 mm., appendages lunate, to three times as
long and broad as gland, white, fifth gland deltoid, short or obsolete,
sinus to one fourth depth of involucre. Staminate flowers ten to eighteen
per cyathium, androphores glabrous, to 1 mm. long at maturity. Gynophore
pilose to 1.5 mm. long at maturity, exserted, nutant; calyx a triangular
pad of tissue; ovary strigose; styles upright, to 0.7 mm. long, joined at
base, bifid for 1/3 length. Capsule densely pilose, broadly ovoid or
conical, 1.5 2 mm. long, 2 mm. wide below equator, broadly three-lobed,
angles acute. Seed oblong-ovoid, to 1 mm. long, 0.6 mm. wide, strongly
four-angled, ventral angle more obtuse and rounded than other three, faces
flat, with 2-4 deep transverse ridges, dorsal faces with an additional
central longitudinal ridge, black, often with light angles.
A common roadside weed in Jamaica and Haiti, less common in Puerto
Rico. Collections have been seen from Barbados and Tobago, and the plant
is reported from Central America and northern South America (Wheeler,
1939) (Map 14).
Specimens seen: JAMAICA. St. Andrew: Kingston, 29 Aug. 1908,
Britton 2978 (NY); Irish Town, 17 June 1963, Crosby, Hespenheide & Anderson
186 (DUKE, PUR); Liguanea, 7 Feb. 1960, Proctor 20564 (IJ, NY, PUR); Stony
Hill, 21 July 1912, Harris 11136 (NY, UCWI). St. Thomas: Easington
Bridge, 23 Mar. 1954, Proctor 8512 (IJ). Manchester: Warwick, 8 Nov. 1962,
Proctor 22886 (IJ). St. Elizabeth: Unity Hill, Aug. 1961, Robertson 9136
(UCWI). HAITI. Port-au-Prince, Sept. 1919, Buch 1713 (IJ); St. Marc,
25-28 Feb. 1920, Leonard 2954 (NY); Petionville, 15-28 June 1920, Leonard
4967 (NY); St. Michel de l'Atalaye, 26 Dec. 1925, Leonard 8483 (MO); Jean
Rabel, 27 Jan.-9 Feb. 1929, Leonard & Leonard 12789 (NY). PUERTO RICO. Coamo,
23 Nov. 1899, Goll 756 (NY); Juana Diaz, 3 Nov. 1913, Hess 4002 (NY).
LESSER ANTILLES. Barbados, 30 July 1901, Freeman s.n. (UCWI); Tobago,
6 Feb. 1913, Broadway 4415 (GH, MO).
This species is distinguishable by the dense pubescence of its
stem and leaves, which even the most heavily pubescent specimens of C.
hyssopifolia do not approach. In most other characters there is a
close resemblance between the two.
Wheeler's application of the name Euphorbia hypericifolia L. to
this species was based on a misidentified specimen in the Linnaean
Herbarium, which was not seen or annotated by Linnaeus until long after
the publication of the name.
12. CHAMAESYCE PARCIFLORA (URB.) comb. nov. Euphorbia parciflora
Urban in Fedde, Repert. 15: 411. 1919. Type: prope Marmelade, Haiti,
1-2 Aug. 1905, G. V. Nash & N. Taylor 1269 (in Herb. Bot. Mus. Berlin,
if extant; Isotype: NY 1).
Perennial; erect or ascending to 3 dm. Stem to 2 mm. diameter,
internodes to 3 mm. long; branched sparsely throughout length, glabrous
or sparingly short-tomentose on one surface, dark brown. Leaves membranous;
blades ovate, 6-10 x 3-5 mm., base inequilateral, rounded, margin sharply
serrate, apex acute to obtuse, midrib distinct, glabrous, olive-green;
petiole 1 mm. long; stipules joined, triangular to 1 mm. long, toothed
or deeply cleft, sometimes bifid and almost distinct. Cyathia solitary
at upper nodes; peduncle slender, to 3 mm. long, involucre campanulate,
to 1.8 mm. diameter at mouth, lobes barely exceeding glands, attenuate,
glabrous on outside, densely ciliate within, glands transversely elliptic,
0.6 x 0.1 mm., cupped, appendages as long as gland and up to four times
as wide, creamy white, margin entire or crenulate, fifth gland linear,
.M.a.p 52.-... -.: f C -- -
ap ^ ] st i ut on of C p rc fl r
Map 16. Distribution of C. brittonii
almost equalling lobes, sinus shallow. Staminate flowers 4-12 per
cyathium, androphores glabrous, to 1.3 mm. long at maturity. Gynophore
glabrous, to 2.5 mm. long at maturity, exserted, nutant; calyx obsolete
or a subcircular pad of tissue; ovary glabrous; styles spreading, 0.5 mm.
long, joined at base, bifid for 1/2 length. Capsules glabrous, broadly
ovoid, 2 2.2 mm. long, 2.5 2.8 mm. wide above base, narrowly three-
lobed, angles acute. Seed ovate, to 1.3 mm. long, 1 mm. diameter just
above base, four-angled, dorsal faces larger than ventral, dorsal angle
acute, ventral angle flattened, faces flat, wrinkled, white.
Reported only from the mountains of Haiti, at elevations of 500-
2000 m. in pineland or open woods (Map 15).
Specimens seen: HAITI. Badeau, 21 Feb. 1927, Ekman H4968 (IJ);
Gonaives, June 1901, Buch 634 (IJ); M. Bonfiere, 29 Sept. 1925, Ekman
H7627 (IJ); Petionville, 24 Aug. 1924, Ekman H1593 (IJ); San Michel to
Marmelade, 6 Aug. 1905, Nash & Taylor 1466 (NY).
The affinities of this species are not clear. Some of the extreme
forms of C. hyssopifolia in Hispaniola have similar cyathia but are narrow-
leaved, and plants growing strongly usually develop lateral dichasia of
cyathia as they get older.
13. CHAMAESYCE BRITTONII (Millsp.) Millsp., Field Mus. Bot. 2:
303. 1909. Euphorbia brittonii Millsp., Field Mus. Bot. 2: 159. 1906.
Type: Race course, New Providence, 26 Aug. 1904, Britton & Brace 839
(Lectotype: F !; Isotype: NY .).
Chamaesyce paucipila (Urb.) Millsp., Field Mus. Bot. 2: 410. 1916.
Euphorbia paucipila Urban, Symb. Ant., 5: 389. 1908. Type: Prope
Guanabacoa, Habana, Cuba, August, Baker and Hasselbring 7199 (Holotype: