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Full text of "Flora of Peru : Euphorbiaceae, spurge family"




-L I B RARY 

OF THL 

U N I VERS ITY 
OF ILLINOIS 

56O.5 

FB 
Y.13 
pt. 3- 



Return this book on or before the 
Latest Date stamped below. 

University of Illinois Library 



L161 H41 



FLORA OF PERU 



BY 

J. FRANCIS MACBRIDE 

CURATOR, PERUVIAN BOTANY 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART IIIA, NUMBER 1 

OCTOBER 17, 1961 

PUBLICATION 680 



HIST. 



FLORA OF PERU 



BY 

J. FRANCIS MACBRIDE 

CURATOR, PERUVIAN BOTANY 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART IIIA, NUMBER 1 

OCTOBER 17, 1951 

PUBLICATION 680 



PRINTED WITH THE ASSISTANCE OF 

The Frederick R. and Abby K. Babcock Fund 



PRINTED IN THE UNITED STATES OF AMERICA 
BY FIELD MUSEUM PRESS 



V.I 



FLORA OF PERU 



J. FRANCIS MACBRIDE 



EUPHORBIAGEAE. Spurge Family 

Herbs, shrubs or trees, sometimes vines, rarely in Peru fleshy 

o or simulating in form some cactus, often more or less lactescent. 

A Leaves simple, entire or dentate but lobed only in a few genera, 

sometimes deeply and even tri-foliate. Flowers monoecious or 

dioecious, ordinarily small and apetalous or also asepalous, some- 

times enclosed in a calyx-like involucre. Stamens 1-many. Ovary 

cr^uperior, 3(l-4)-celled with 1 or 2 pendulous ovules in each cell. 



or stigmas as many or twice as many as ovary cells. Capsules 
c-jcommonly 3-lobed, 3-valved. Embryo straight, the flat cotyledons 
^.almost as wide as the fleshy oily endosperm. 

^ The genus Hevea in this large family is the most important source 

of natural rubber, Manihot of the starch cassava (tapioca), Ricinus 

of castor oil, all plants known in Peru either native or in cultivation. 

The Asian Aleurites Fordii and other species yielding tung oil, could, 

r as in other warm regions, become valuable cultivated trees. Notable 

^ as the family is in an economic way it is scarcely less exploited in 

., horticulture, an outstanding example in Peru being the widely 

grown and sometimes adventive foliage ornamental "Croton," its 

leaves usually variously mottled with yellow and red; it is Codiaeum 

^ : variegatum (L.) Blume from Malay and the islands of the Pacific 

and is quite distinct from the genus Croton so common in Peru. 

[ J As yet less familiar in Peru is the "Pascua" of Mexico (Euphorbia 

pulcherrima Willd.), cultivated in northern countries as Christmas 

r Flower or Poinsettia, named after Poinsett, a minister to Mexico 

from the United States who promoted the plant in horticulture 

about 1828. 

This compilation is taken by and large from Pax & Hoffmann's 
revision in Pflanzenreich as cited for each genus; I acknowledge 
my indebtedness to their careful treatment. However, it is my 
impression that they have not succeeded in advancing the taxonomy 
of the group but have accepted and proposed far too many mono- 
typic or small genera, apparently on the basis of tradition or logic, 
and without the comparative consideration of their general relation- 
ships that probably would result in their inclusion within better- 

3 



4 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

defined groups at most as sections. Even some generic groupings 
seem less satisfactory than those of Mueller; one example is their 
emphasis on the presence of a rudimentary ovary, a character of 
course often developed in degree and not "holding," according to 
descriptions. As elsewhere in this work the terms "male" and 
"female," for convenience and to avoid repetition, have been asso- 
ciated not alone with the word "flower." 

The key is based largely on the male flowers, more generally 
present in herbaria. 

Flowers (male and female) surrounded by a calyx-like involucre; 
perianth none or minute. 

Involucre regular, calyx-like or cupulate 1. Euphorbia. 

Involucre irregular, oblique, appendaged 2. Pedilanthus. 

Flowers not involucrate or rarely and then the involucre, unless in 
Per a, containing only one sex; perianth usually present. 

Ovules geminate; petals if present usually scale-like; flowers com- 
monly fasciculate or solitary in the leaf -axils (spicate or race- 
mose in Richeria, paniculate in Hyeronima). 

Fruit capsular; pubescence not lepidote. (Also see genera men- 
tioned under Dry petes.) 

Petals none; plants often ligneous, at least in Peru eglandular. 
Petioles short or wanting; herbs, shrubs, rarely trees. 

Flowers solitary or in open clusters; herbs or shrubs. 

3. Phyllanihus. 

Flowers in axillary glomerules; trees 4. Securinega. 

Petioles elongate; trees 5. Richeria. 

Petals 5; herb or half shrub, in Peru glandular. .6. Andrachne. 
Fruit drupaceous; petals none; pubescence often lepidote. 

Flowers paniculate; pubescence lepidote; petioles long. 

7. Hyeronima. 

Flowers fasciculate; pubescence not lepidote; petioles short. 

8. Dry petes. 

Ovules 1 in each cell; flowers not fascicled or solitary (bracteately 
involucrate in Dalechampia and Per a], inflorescences some- 
times conoid or short and glomerulate. 

Stamens inflexed in bud; indument either stellate or lepidote, 
usually conspicuous; rudimentary ovary none. . .9. Croton. 



FLORA OF PERU 5 

Stamens straight in bud, sometimes sessile, anther tips directed 

upward or (and) rudimentary ovary obvious. 
Segments of male calyx valvate in bud, rarely wanting (in 
Peru, Pera); male disk or receptacle often present (as 
noted); petals developed only in first three genera 
(male flowers unknown in Conceveibastrum). 
Petals of male flowers well developed. 

Ovary rudiment none or obsolete; disk glandular; tri- 
chomes in part malpighiaceous. 

Petals entire 10. Argythamnia. 

Petals parted 11. Chiropetalum. 

Ovary rudiment obvious; disk none; trichomes simple. 

12. Caperonia. 

Petals wanting in flowers of both sexes. 
Inflorescence partly involucrate by small, calyciform or 
even foliaceous bracts; disk none. 

Inflorescence bisexual; bracts geminate, often large. 

13. Dalechampia. 

Inflorescence unisexual; bracts calyciform. . .14. Pera. 
Inflorescence not bracteately involucrate. 
Leaves divided or deeply lobed. 

Leaves trifoliate; disk glandular 15. Hevea. 

Leaves deeply lobed; disk none 16. Ricinus. 

Leaves entire or not lobed. 
Plants twining, scandent or if erect pubescence in 

part stinging, unless Megalostylis. 
Capsules 4-celled, except Megalostylis; ligneous 
plants without stinging trichomes; stamen 
receptacle more or less developed. 
Stamens free or mixed with glands; leaves 

palmately 3-nerved from base. 
Female flowers at base of male racemes; 

stamens free 17. Plukenetia. 

Female flowers in axillary racemes; filaments 

mixed with glands 18. Elaeophora. 

Stamens united into a columnar or globular 
receptacle; leaves pinnately nerved or 5- 
nerved from base. 



6 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Receptacle globose, the sessile anthers many; 
leaves pinnate-nerved ... 19. Apodandra. 

Receptacle columnar, the anthers 6; leaves 
basally 5-nerved 20. Megalostylis. 

Capsules 3-celled; plants sometimes half-shrubs 
but usually with stinging trichomes; disk 
none or glandular 21. Tragia. 

Plants not twining or scandent. 

Anther cells elongate and narrow, often flexuose; 
flowers usually in slender spikes or racemes, 
the male, especially, dense. . . .22. Acalypha. 

Anther cells globose or oblong; flower clusters 
more or less remote or spikes short; disk or 
receptacle present. 

Male flowers panicled, sometimes narrowly (un- 
known in Conceveibastrum; genera doubtful). 

Styles slender, usually long and soon con- 
spicuous; stamens 8 or fewer. 

23. Akhornea. 

Styles rather stout, short, papillose. 
Leaves cordate; stamens (known) 3-5. 

Leaves glandular; female sepals 4-6. 

24. Aparisthmium. 

Leaves eglandular; female sepals 10-12. 
25. Conceveibastrum. 

Leaves oblong-elliptic; stamens 16. 

26. Conceveiba. 

Male flowers racemose or spicate, the inflores- 
cence elongate (Peru). 

Disk thick, sometimes 1 or 4-6 stamens cen- 
tral 27. Caryodendron. 

Disk none or glandular, the stamens then 

4-22, free. 

Disk none; styles wanting or elongate. 
Stamens 2-3; stigmas sessile or nearly. 
28. Adenophaedra. 

Stamens many; styles elongate, bifid. 

29. Cleidion. 



FLORA OF PERU 7 

Disk usually glandular; styles short. 

30. Bernardia. 

Segments of male calyx imbricate or open in bud or before 

anthesis, or subunited and cupulate, often reduced or 

wanting; disk none unless in first five genera; petals 

none except Jatropha, Pausandra, Pogonophora. 

Flowers in dichotomous cymes or congested cymules; leaves 

usually lobed. 
Leaves usually lobed; cymes dichotomous. 

Petals present; pubescence innocuous 31. Jatropha. 

Petals lacking; pubescence stinging. . .32. Cnidoscolus. 

Leaves entire; cymules congested 33. Cunuria. 

Flowers variously disposed, not dichotomously cymose, 

nor in congested cymules. 
Leaves deeply lobed, at least some, or linear. 

34. Manihot. 

Leaves neither lobed nor linear. 
Petals present. 

Leaves dentate; flowers in spicately arranged 

glomerules 35. Pausandra. 

Leaves entire. 

Flowers in short axillary panicles. 

36. Pogonophora. 
Flowers in elongate panicles in upper axils. 

37. Dodecastigma. 
Petals lacking. 

Petioles biglandular at or toward the tip. 
Erect; glands small; foliose bracts lacking; fruit 

capsular. 
Spikes commonly terminal; approximate leaf 

nerves glabrous. 

Pericarp base persisting as a 3-radiate 
receptacle; petioles short. 44. Stillingia. 
Pericarp base not as above; petioles (Peru) 
mostly or all 1.5-3 cm. long. 

45. Sapium. 

Racemes or panicles axillary; remote leaf 
nerves early pubescent. 

38. Tetrorchidium. 



8 FIELD MUSEUM OP NATURAL HISTORY BOTANY, VOL. XIII 

Scandent; glands 2 mm. long; bracts in part 
foliose; fruit fleshy 39. Omphalea. 

Petioles eglandular (leaf blades sometimes glan- 
dular at or above base). 

Male flowers in racemes or panicles, these some- 
times narrow or panicled spikes. 

Petioles of subelliptic leaves apically tumidu- 
lous, slender; stamens usually 5-12; 
styles nearly free. 

Male flowers panicled; sepals 4; ovary 
2-celled 40. Nealchornea. 

Male flowers in panicled spikes; sepals 3-5; 
ovary usually 3-celled.41. Senefeldera. 

Petioles of oblongish leaves not tumid at apex; 
stamens 10-70 or 2-3; styles connate. 

42. Mabea. 

Male flowers in very slender or often multi- 
bracted, rather ovoid, simple spikes. 

Leaves small, serrulate; spikes slender, ter- 
minal, opposite leaves or on leafy branch- 
lets. 

Pericarp not persisting; male calyx usually 

3-lobed or -parted. . . .43. Sebastiania. 

Pericarp persisting, 2-3-radiate; male calyx 

shortly bilobed 44. Stillingia. 

Leaves entire or ample; spikes subovoid or 
short and axillary. 

Spikes axillary 46. Actinostemon. 

Spikes terminal. 

Leaves not cordate 47. Maprounea. 

Leaves cordate 48. Hura. 

1. EUPHORBIA L. 

Reference: Boissier in DeCandolle Prodr. 15, pt. 2: 1-187. 1862. 

Variable vegetatively as commonly interpreted, but drawn to 
include plants with milky sap, simple leaves, and several male 
flowers enclosed with one centrally disposed female in a calyx-like, 
cup-shaped, sometimes oblique involucre called a cyathium, this with 



FLORA OF PERU 9 

some narrow bracts within, 4-5 lobes and often as many glands 
without and alternate with the lobes, the glands frequently with a 
white or colored appendage. Male flowers early deciduous from 
articulate pedicels, 1 stamen (the anther cells ordinarily globose), 
and usually without a calyx, this sometimes present in the female 
flower, minute, 3-lobed. Ovary sessile on pedicel tip, 3-celled, 3- 
ovuled, the 3 styles free or partly connate, often bifid. Capsules 3 
bivalved cocci finally separating from the persisting axis and opening 
ventrally. Seeds often ecarunculate. 

Louis Cutter Wheeler's careful and detailed revision of the sub- 
genus Chamaesyce in Canada and the United States, Rhodora 43: 
1941, has been freely drawn upon as regards the few species extend- 
ing to Peru; I record my indebtedness with appreciation. The page 
references after these species, indicated by W followed by a number, 
refer to his work; others of this subgenus, indicated by B followed 
by a number, refer to that of Boissier. Croizat, Journ. Arnold Arb. 
24: 182-184. 1943, summed up his arguments for the recognition of 
Chamaesyce S. Gray emend. Croizat as a distinct genus. Casual as 
is my knowledge of the family it seems to me that there is no clari- 
fication possible from any viewpoint by disturbing the status quo 
in the taxonomy of this group and its allies and that the constructive 
work in its taxonomy remains to be done within the other 
tribes. 

Involved nomenclature and characters, the relative significance 
of the latter apparently often still undetermined, combine to make 
this group a fascinating challenge to the student, a boresome annoy- 
ance to the compiler. However, supported by Wheeler's feeling that 
even the subgenus Chamaesyce is difficult of demarcation I have 
arranged the following key on the basis of expediency, but most of 
the species that grossly resemble one another are brought into con- 
trast. 

Poinsettia or Flor de Pascua (E. pukherrima Willd., native of 
Mexico) and Corona de Cristo, "Crown of Thorns" (E. Milii Des 
Moulins, the earliest name forE 1 . splendens Boj.; cf. Croizat, Journ. 
Arnold Arb. 21: 506. 1940) are the well-known cultivated examples 
of this large group. 

The general interest in the genus lies further in the fact that 
many herbaceous species may harbor the organisms that cause 
tropical leg-ulcers; Wheeler cites Mesnil, Ann. Sci. Nat. se>. 10. 3: 
xlii-lvii. 1921, for an interesting re"sum4 of the voluminous literature 
relating to flagellates in Euphorbia and other lactiferous plants. 



10 FIELD MUSEUM OP NATURAL HISTORY BOTANY, VOL. XIII 

Plants cactus-like, more or less succulent or at least thorny. 

Leaves rudimentary; bracts not showy E. Weberbaueri. 

Leaves developed; bracts showy, red E. Milii. 

Plants tall shrubs or small trees with crowded oblanceolate-obovate 

leaves much longer than broad, exstipulate. 
Leaves abruptly caudate; bractlets partly adnate. .E. Tessmannii. 
Leaves mucronate or short-acuminate; bractlets deciduous or 

apparently distinct. 
Cymes monocephalous; petioles to 5 mm. long.E 1 . apurimacensis. 

Cymes polycephalous; petioles to 2 cm. long E. laurifolia. 

Plants never cactus-like, rarely shrubs, the leaves then neither 

crowded nor much longer than broad. 
Involucres with 1-4 cornute or cupulate glands, sessile or crowded 

and terminal; annuals, the leaves mostly alternate. 
Leaves acute or acuminate; glands cupulate, usually 1. 

E. heterophylla. 

Leaves rounded-obtuse; glands 4, cornute E. Peplus. 

Involucre glands usually 4-5, not cornute or cupulate, sometimes 

fimbriate or petaloid appendaged. 
Petioles of mature leaves, and usually even the upper at least 

about a third as long as the blades. 
Involucre teeth as appendages notably setaceous or ciliate- 

fimbriate; stipules setaceous E. adiantoides. 

Involucre teeth rather broadly fimbriate, the petaloid ap- 
pendages entire or crenate-dentate; stipules glanduli- 
form or subulate. 

Shrubs, trees or at least clearly suffrutescent below. 
Leaves rounded at base, mostly 3-8 cm. long; petioles 

elongate E. cotinoides. 

Leaves usually acute at base, mostly or all 1-2 cm. long; 

petioles to 1 cm. long E. viridis. 

Annuals or scarcely more enduring (unless E. graminea), 
with tall or weak slender herbaceous stems, not 
suffrutescent even below. 

Involucres narrow, long-pedicellate in open cymes soon 

exceeding the little reduced leaves. . . .E. Spruceana. 

Involucres subcampanulate, short-pedicellate, mostly in 

small terminal cymes little if at all longer than the 

reduced leaves. 



FLORA OF PERU 11 

Leaves canescent beneath with a short indument. 

E. arenaria. 

Leaves glabrous both sides or indument more or less 

long-pilose. 

Seeds pitted, ovoid; leaves pubescent, glabrate in 
age E. Poeppigii. 

Seeds irregularly tubercled, broader than long; leaves 
glabrous or nearly E. graminea. 

Petioles short, sometimes even obsolete, in any case very much 
shorter than the blades. 

Erect, diffuse or spreading, if annual, leaves ordinarily 1 cm. 
long or longer, internodes soon usually longer, or plants 
obviously perennial, the often smaller leaves approximate, 
pubescence of annuals (if present) usually in part elongate, 
unequal in length. 

Annuals or rarely more enduring but the stems or branches 
characteristically (at least in age) erect, rarely prostrate 
but internodes mostly longer than leaves; stipules 
present unless in E. peruviana and E. tacnensis. 

Involucres unless in a variant in dense axillary and 
terminal leafless glomerules. 

Plants distinctly pubescent E. hirta. 

Plants glabrous or nearly E. glomerifera. 

Involucres solitary or in few-flowered axillary cymes or, 
if terminal, dichotomous. 

Gland appendages laciniate; stipules none.E". tacnensis. 

Gland appendages usually entire; stipules rarely ob- 
solete. 

Styles short, the stigmas not enlarged; leaves often 
oblongish, 1-3 cm. long; appendages entire or 
2-3-lobulate. 

Ovary typically glabrous (scarcely valid character). 

Capsules wider below middle, 1.6 mm. long; 
cymes not very dense E. hyssopifolia. 

Capsules widest at middle, to 1.4 mm. long; 
cymes dense E. glomerifera. 

Ovary as capsule more or less pubescent as to types. 
E. brasiliensis, E. hypericifolia. 



12 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Styles to 1.3 mm. long, stigmas clavate or ellipsoid; 
leaves about oval, 4-12 mm. long; appendages 
usually crenate or dentate. 
Stipules usually in 1-2 linear segments. 

E. peruviana. 

Stipules fimbriate; appendages crenate (always?). 

E. Meyeniana. 

Perennials, either low or prostrate herbs or suffrutescent 
at base and then especially often spreading-diffuse; 
leaves usually approximate. 
Plants glabrous or merely pulverulent-pruinose. 
Stipules undeveloped; glands unappendaged. 

Leaves coarsely mucronate-dentate .". Hinkleyorum. 

Leaves entire or the denticulation minute, obscure. 

Leaves obovate-subrotund, sessile or subsessile. 

E. chilensis, E. huanchahana. 
Leaves orbiculate, distinctly petioled. 

E. raphanorrhiza. 
Stipules developed; glands appendaged sometimes 

minutely or concolored. 
Leaves orbiculate, shortly petioled; seeds smooth. 

E. orbiculata. 

Leaves obovate or at least oblique at base, subsessile. 
Appendages white, showy, longer than glands. 

E. Ruiziana. 
Appendages nearly or quite concolor with glands, 

minute E. melanocarpa. 

Plants obviously pilose, stipulate. 

Involucres in part dichotomous; appendages broad, 

crenulate or rarely subentire E. Meyeniana. 

Involucres all solitary; appendages narrow, entire, the 

red edge undulate E. trancapatae. 

Depressed annuals (rarely more enduring) soon forming mats 
or cushions, the leaves ordinarily a few mm. long, the 
internodes usually about as long (1-2 or 3 mm.) unless 
in form of E. micromera and young plants of E. hyperici- 
folia; pubescence often short or lacking. 
Leaves entire or at least most of them, none denticulate 
unless obscurely at tip; seeds smooth or obsoletely 
and undulately rugulose. 



FLORA OF PERU 13 

Gland appendages obsolete or rudimentary; stipules dis- 
tinct or none E. micromera. 

Gland appendages obvious, often conspicuous; stipules 
a single membranous scale (except in mentioned 

species) E. serpens. 

Leaves denticulate, at least usually some of them; seeds 

usually rugose, at least granular; glands appendaged. 

Capsules glabrous or sparsely villous; seeds not ridged 

but sometimes low-rugulose; involucres usually in 

short leafy cymes. . . .E. brasiliensis, E. hypericifolia. 

Capsules strigose or, if trichomes spreading, the seeds 

ridged. 

Capsules strigose; seeds with low rounded concolored 
ridges; involucres 2-several on axillary branchlets. 

E. thymifolia. 

Capsules tomentose or the trichomes crisped-spreading; 
seeds transversely deeply or closely sulcate or 
ridged E. viatilis. 

Euphorbia adiantoides Lam. Encycl. 2: 426. 1788; B57. 

Erect striate glabrous stems attaining several dm. and shortly 
dichotomously branched, the very slender branchlets fragile, about 
1 dm. long; stipules setaceous; petioles elongate; leaves opposite, 
entire, thin-membranous, glabrescent, ovate-elliptic, subcuneate at 
base, obtuse, mucronulate, 2.5-3.5 cm. long; involucres in the upper 
axils and terminal, shortly pedicelled, turbinate, hirtellous with 
ovate fimbriate teeth, the ovate transverse glands a third narrower 
than the 5-6-setaceous-parted appendages; styles undivided; cap- 
sules hirtellous; seeds spheroid, tuberculate (ridges dentate in Weber- 
bauer collections). The Peruvian specimen may rather have come 
from Ecuador as now defined. This is a very delicate and singular 
plant with nearly the aspect of Adiantum of Montpellier (Lamarck). 
Croizat, suggesting that the affinity of E. sciadophila Boiss., B57, 
of southeastern Brazil with this species requires study, hints that 
the Peruvian plant may have a wide range. Standley referred the 
recent collections to E. Poeppigii. F.M. Neg. 32508 (Hartweg). 

Tumbez: Rainy-green formation, near Hacienda La Choza, 150 
meters, Weberbauer 7706. Piura: Chanro, 250 meters, Weberbauer 
6002. Without locality, Jussieu, type. Ecuador; Colombia; 
Argentina? 



14 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Euphorbia apurimacensis Croizat, Journ. Arnold Arb. 27: 289. 
1946. 

Small tree, 3-5 meters high, the pale brown bark lenticellate, the 
new shoots puberulent; leaves elliptic-obovate, shortly mucronate- 
acuminate, gradually narrowed to base, this more or less produced 
on the 5 mm. long petiole, 3-7 cm. long, 1-1.5 cm. wide, entire, 
glabrous except for the ciliolate slender spreading obscure veins; 
involucres single, puberulent, about 3 mm. long, 5 mm. wide at 
throat, glabrous, entire, erect; peduncle 5 mm. long, bractlets 
deciduous, sub triangular, 4-5 mm. long. Related to E. Latazi 
HBK., B106, of Ecuador and Colombia with longer petioles (-2 cm.), 
peduncles and smaller involucres (Croizat), but it seems probable 
that these differences are individual rather than specific and that 
the plant is a part of E. laurifolia Juss., to which Diels referred the 
plant of HBK. 

Apurimac: Quebrada de Matara, xerophytic slopes, 2-2,800 
meters, Prov. Abancay, Vargas 2290, type. 

Euphorbia arenaria HBK. Nov. Gen. & Sp. 2: 57. 1817; B56. 

Erect herb, crisp pubescent, the indument on the leaves ap- 
pressed and canescent beneath, the stems firm but fistulose as 
the opposite branches; petioles 8-16 mm. long; leaves opposite or 
ternate, oblong-elliptic or slightly ovate, entire, 3-5 cm. long, about 
half as wide, obtusish both ends, green and glabrate above, the 
floral petaloid, white, 3-4 mm. long, exceeding the involucres; cymes 
terminal with 5-7 heads, the peduncle -to 7 cm. long; involucres 
subcampanulate, shortly pedicellate, appressed hirsutulous, the lobes 
fimbriate-dentate, glands 4, yellowish, hirsute without, entire, 
rounded; styles biparted; capsule hirsute; seeds obscurely tubercu- 
late. Capsule unknown in type but ovary described as globose, 
smooth, rarely with a few trichomes; therefore the above description 
probably applies only to var. quitensis Boiss. I.e., defined as glabres- 
cent with smaller leaves, possibly distinct. F.M. Neg. 18039. 

Ancash: Trujillo above Casma on coast, Bonpland, type. Ecua- 
dor? Colombia? 

Euphorbia brasiliensis Lam. Encycl. 2: 423. 1788; B24. 

Annual, usually erect, glabrous or patently pubescent, often 
several dm. high and more or less branched; stipules small, ciliate; 
leaves sessile or subsessile, oblong to ovate or obovate, oblique at 
base, subacute or obtuse, serrulate, 1-3.5 cm. long, those of the 



FLORA OF PERU 15 

axillary inflorescences narrower; involucres few, mixed with linear 
bracts, campanulate, glabrous; glands oblong or orbicular with 
rather conspicuous transversely elliptic white appendages; capsules 
white pubescent at least on backs of carpels, 2 mm. in diameter; 
styles deeply bifid; seeds subquadrangular, with 2-3 transverse 
ridges. Lanjouw, in Pulle, Fl. Surinam 2, pt. 1: 95. 1932, noted 
that the inflorescence is not terminal as previously described. 
Wheeler, Rhodora 43: 143. 1941, has followed Mueller in suggesting, 
apparently with reason, that this is a variant of E. hyssopifolia, 
probably the only difference being the pubescent ovary and capsule. 
Cf. also E. hypericifolia. Illustrated, Miq. Stirp. Surin. Sel. pi. 28 
(asE. Klotzschiana). 

Peru: Material that could be placed here is cited for convenience 
under E. hypericifolia, the earliest name for these closely related 
forms. Generally distributed in the warmer parts of the Americas. 

Euphorbia chilensis Gay, Fl. Chile 5: 335. 1849; B103. E. 
portulacoides Spreng. Syst. 3: 792. 1826, not L. 

Glabrous herb (rarely puberulent), the several spreading-decum- 
bent stems ascending from elongate woody roots often 1-2 dm. long, 
branched above, densely leafy, with sessile obovate-subrotund some- 
times oblanceolate leaves attenuate to base, usually 7-15 mm. long, 
5-7 mm. wide, rounded at tip or, in var. acutifolia Boiss., apparently 
the typical state, oblong-lanceolate, acute; floral leaves like the 
lower, little smaller, terminating the dichotomous branches; involu- 
cres more or less pedicellate, hemispheric, the throat hirsute, as 
the oblong truncate 4-fibrillate lobes; glands purplish, transverse- 
oblong, subundulate, depressed punctate; capsules long-pedicellate, 
depressed, deeply trisulcate; seeds ovoid-subtetragonous, smooth, at 
least in some material so determined in herbaria, about 3 mm. long, 
light gray. Doubtfully in Peru in typical state but compare the 
apparently scarcely distinct E. huanchahana. Possibly E. caespitosa 
Lam., B103, of Uruguay, low from tuberous root, leaves 8-12 mm. 
long, capsules acutely trigonous, not depressed, little sulcate, 4 mm. 
long and broad, may prove to be the earliest name for a variable 
species; intermediate apparently is E. copiapina Phil., B103, the 
deeply sulcate capsules about 2 mm. long, 3 mm. wide. My number 
3071 placed for expediency under E. huanchahana matches closely 
material of E. caespitosa from Montevideo, except that it has the 
small capsules of E. copiapina, not, however, deeply sulcate! 

Peru (possibly). Chile; apparently Argentina and Bolivia. 



16 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Euphorbia cotinoides Miq. Stirp. Surin. Sel. 96. 1850; B60. 
Alectoroctonum cotinoides (Miq.) Kl. & Gke. Abh. Akad. Wiss. 
Berlin 1859: 40. 1860. 

A glabrous shrub sometimes several meters high, with articulated 
stems and ternate or opposite branches and leaves, the latter broadly 
ovate or elliptic, often subpeltate at the rounded base, apically 
rounded or usually obtuse or acute, 3-8 cm. long (petioles about 
as long), 1.5-5 cm. wide, rigid-membranous, glaucous beneath, entire, 
with 7-10 lateral nerves; stipules rounded, glanduliform or those of 
the floral leaves subulate; cymes axillary, lax, corymbose-paniculate, 
sometimes with peduncles 4-6 cm. long; involucres about 2 mm. 
long, sparsely hirtellous, the minute lobes fimbriate, the broad 
glands with larger white crenulate appendages; interfloral bractlets 
many, pubescent; ovary pedicellate; capsules apparently not yet 
described. My 2907 has glabrous rotund leaves rounded at both 
ends, sparsely pubescent capsules, ovoid seeds little narrowed at 
obtuse tip, 2 mm. long, about 1.5 mm. thick, closely blunt-tubercu- 
late, dark gray; the 4 appendages are green and subentire as in 
E. viridis, which it also resembles more in habit than E. cotinoides 
except for the broad leaves on elongate petioles, and perhaps it should 
be placed with E. viridis as a local variant. Someone has suggested 
it may be E. caracasana (Kl. & Gke.) M. Arg., but that at least 
typically has leaves hirsute on nerves beneath, 5 glands, oblong seeds 
irregularly foveolate and confluently scrobiculate. The leaves are 
used as fish "poison." 

Tumbez: East of Hacienda Chicama, 900 meters, Weberbauer 
7655. San Martin: Tarapoto, Spruce 4502 Cdet. Mueller); Williams 
5915 (det. Steyermark). San Roque, Williams 7303. Pongo de 
Cainarachi, Klug 2674 (det. Steyermark). Loreto: Yurimaguas, 
Poeppig. Lima: In slide rock at Matucana, 2907? To the Guianas. 
"Juquilla," "yuquillo" (Williams), "huarus" (Weberbauer). 

Euphorbia glomerifera (Millsp.) Wheeler, Contr. Gray Herb. 
127: 78. 1939. Chamaesyce glomerifera Millsp. Field Mus. Bot. 2: 
377. 1913. E. hypericijolia of auth., not L., accord, to Wheeler, 
I.e. 73 et seq. 

Slender, erect, usually glabrous (or nearly) annual, the often 
purplish stems commonly branched and attaining 1-several dm. in 
length, the axillary cymose peduncled inflorescences ordinarily dense 
and many-flowered; stipules ovate, dentate, ciliate; petioles very 
short; leaves oblong, oval or obovate, rounded or obtuse at the 



FLORA OF PERU 17 

strongly oblique base, serrate, paler beneath, mostly 1.5-3 cm. long; 
involucres very small, turbinate, hirtellous only within, the lance- 
triangular lobes lacerate-dentate; glands stipitate, suborbicular, 
with usually as wide, sometimes ovate white or reddish appendages; 
capsules glabrous, 1.5 mm. broad; seeds red, ovoid-trigonous, with 
conspicuous angles and irregularly rugose sides. After Standley 
and Steyermark, Fieldiana: Bot. 24, pt. 6: 101. 1949. Croizat, Bull. 
Torrey Club 75: 407-408. 1948, wrote: "I accept this binomial as a 
matter of convenience . . . but I fail to see how this form can be 
extricated as a separate species out of the Old World C. pilulifera 
(L.) Small, C. hypericifolia (L.) Millsp. and C. indica (Lam.) Croiz.," 
a remark which seems, from even my casual observations, apropos. 
Peru (probably). North to Mexico, southern United States and 
the West Indies. 

Euphorbia graminea Jacq. Sel. Stirp. Amer. Hist. 151. 1763; 
B54. 

Perennial, with erect or weak herbaceous, more or less procum- 
bent stems, somewhat pubescent with spreading or crisped trichomes; 
stipules minute, subulate; petioles slender, elongate, the lower 
alternate, the upper opposite (or verticillate?), the leaves thin- 
membranous, entire, variable in shape, the upper usually elliptic- 
lanceolate, more or less narrowed or acute or rounded both ends, 
often several cm. long; involucres small, pedicellate in forks of 
branches and in terminal cymes, rarely glabrous, the ovate lobes 
fimbriate; glands 2-4, transversely ovate, narrower than the obovate 
subentire white or yellowish appendage; capsules glabrous; seeds 
tuberculate. The type from Carthagena, Colombia; Svenson, Amer. 
Journ. Bot. 33: 460. 1946, described the seeds of a specimen from 
the coast of Ecuador as 1.5-2 mm. broad, 1 mm. long, terete, acute 
apex, gray with reddish dots, broad quadrate indentations, while 
another specimen det. as "affine" had seeds 2 x 1.5 mm. with wart- 
like excrescences; the Peruvian specimen, glabrous, woody below, 
leaves somewhat verticillate, had seeds 2.5 x 2 mm., ashy gray 
becoming black, with irregular obtuse ridges; evidently the group 
needs monographic study to determine the significance of these 
variations. Related species include E. Fraseri Boiss., B55, of Ecuador, 
with appendages narrower than the glands and E. delicatula Boiss., 
B55, herb. Pavon from Nova Hispana (maybe), the appendages 
much longer than the minute glands, nearly glabrous and involucres 
all long-pedicelled in axils. 



18 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Piura: Cerro Prieto, (Haught & Svenson 11620, det. Svenson, 
affine). To Mexico. 

Euphorbia heterophylla L. Sp. PI. 453. 1753; B72. E. geni- 
culata Ortega Decad. 18. 1797. E. elliptica Lam. Encycl. 2: 425. 
1788? Poinsettia Ruiziana Kl. & Gke. Abh. Akad. Wiss. Berlin 
1859: 102. 1860. 

Nearly glabrous or slightly pubescent erect annual, soon sparsely 
leafy or leafless below, simple or with few branches; petioles often 
long, slender but early short; leaves mostly alternate, upper opposite, 
varying greatly in shape, usually oblong-lanceolate to ovate but not 
rarely somewhat panduriform or even linear, entire or the irregularly 
dentate floral often red or pink at base; involucres commonly termin- 
ally congested, the lobes fimbriate, the normally solitary gland un- 
appendaged; capsules 6 mm. broad, glabrous; styles bifid; seeds 
ovoid, acute, grayish or darker, irregularly tuberculate. A linear- 
leaved variant, apparently in Peru, is var. graminifolia (Michx.) 
Engelm. The Lamarck plant may be distinct as thought by Croizat, 
who includes in it E. geniculata and E. prunifolia Jacq.; it is the 
elliptic-leaved form represented by my collections. The Mexican 
and widely cultivated Flor de Pascua (E. pulcherrima Willd.), well 
known for its bright red floral leaves and shrubby habit, is related 
to this weedy species. 

San Martin: Abandoned land, Tarapoto, Williams 6647 (det. 
Standley); 6285. Ancash: Lomas de la Choy, Goodspeed 9221 (det. 
Standley). Lima: In rock slide, San Geronimo, 5894 (det. Johnston, 
E. geniculata}. Matucana, cactus-boulder slope, 273 (det. Mill- 
spaugh, P. geniculata). Echarate (abandoned field), Goodspeed 
10471. Barranca, Beetle 9027 (det. Johnston). Huanuco: Field 
weed, Pozuzo, 4670. Junin: Near Quirimi Bridge, Killip & Smith 
23934 (det. Croizat var. graminifolia following Boissier, with remark 
that critical study may prove it not the same as plant of United 
States). La Merced, stream course and shrubby trail edge, 5234; 
5380 (det. Johnston). Cuzco: Valle del Apurimac, Limatambo, 
2,800 meters, (Vargas 295). Santa Ana, weed in cultivated land, 
Cook & Gilbert 1524 (var. graminifolia, in part). Widely distributed 
in warm America. "Pascua;" "flor de pascua de monte." 

Euphorbia Hinkleyorum Johnst. Contr. Gray Herb. 70: 72. 
1924. 

Glabrous perennial from a stout root, the many erect herbaceous 
stems laxly and dichotomously branched from about the middle; 



FLORA OF PERU 19 

stipules lacking; lower leaves alternate, few, upper and those of 
branches opposite, approximate, conspicuous, concolor, orbicular or 
obovate or subflabelliform, broadly affixed at the truncate base or 
cuneate dentate, 8-24 mm. broad and long, usually distinctly lacerate- 
dentate; involucres solitary in the axils, cyathiform, about 2 mm. 
long, peduncle about 1 mm. long, throat with subulate or narrowly 
deltoid lobes, hirsute, 0.5 mm. long; glands 5, 1.3 mm. long, some- 
times slightly hirsute beneath, laminate-rugulose above, lip 
retuse or truncate-undulate; styles about 1.5 mm. long, stigmas 
little enlarged; seeds oblong, obscurely quadrangular, 2 mm. long, 
1.3 mm. thick, ecarunculate, gray, delicately dark spotted. 
Evidently belongs to Boissier's group Ipecacuanhae, B101, . . . but 
has larger, apparently connate (but actually distinct), broader leaves 
and more loosely branched habit. The collectors noted the plant 
used in the treatment of fevers. 

Arequipa: Sandy pampa south slope, Chachani Mountain, 3,300 
meters, Mr. & Mrs. F. E. Hinkley 7, type. Argentina; Bolivia? 
"Esquera" (Hinkley). 

Euphorbia hirta L. Sp. PI. 454. 1753; W169. E. pilulifera of 
auth. not L., I.e. E. capitata Lam. Encycl. 2: 422. 1788. Chamaesyce 
hirta (L.) Millsp. Field Mus. Bot. 2: 303. 1909. C. pilulifera of auth. 
accord, to Croizat, not L. which centers in C. hypericifolia (Bull. 
Torrey Club 75: 408. 1948). 

Erect to decumbent or prostrate and often abundantly pubescent, 
with yellowish spreading trichomes or these sometimes lacking; 
subsessile or pedunculate cymose terminal and usually lateral heads 
of many cyathia and yellowish capsules characteristic for the species; 
internodes 1-4 (-7) cm. long; petioles 1-2 mm. long; stipules attenu- 
ate, distinct or nearly, often linear-divided at base; leaves usually 
ovate-lanceolate or rhombic, also narrower, oblique at base, serrate, 
commonly 1-3 cm. long, acutish or rarely acuminate, sparsely strigose 
above, sparsely appressed puberulent beneath; involucre lobes cilio- 
late; white appendages narrower to twice as wide as the small stiped 
glands; ovary minutely strigose; seeds reddish, acutely ovoid- 
trigonous, 0.7-0.9 mm. long, transverse rugulose. After Wheeler 
in part, who adopted the above synonymy, the reason for his action 
elucidated by Svenson, Amer. Journ. Bot. 33: 460. 1946. A well- 
marked variant, maybe distinct, of Piura and Ecuador, is var. 
destituta Wheeler, Contr. Gray Herb. 127: 70. pi. 4c, fig. 1. 1939, 
long internodes nearly glabrous, cyathia smaller, leaves acuminate, 



20 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

seeds 0.7 x 0.3 (Svenson); Croizat, Journ. Arnold Arb. 24: 188. 1943, 
suggested this might be E. microcephala Boiss., B1262, of Bolivia, 
but that has obtuse leaves. When the plants are low and freely 
branched the cymules according to Wheeler are all terminal or on 
leafy branchlets and then referable to var. procumbens (DC.) N. E. 
Brown, a disposition agreed upon by both Croizat and Moldenke 
but under Chamaesyce as a subspecies or as a variety. Very near 
and maybe in Peru but glabrous even to ovary is E. glomerifera 
(Millsp.) Wheeler, W168. Illustrated, Wheeler, pi. 657A, opposite 
page 168. 

This is one of the "herbas de la golondrina," used, according to 
Ruiz and Pavon, to destroy cataracts and as a purge. According 
to M. Martinez, whom Wheeler has thoughtfully quoted, the name 
"golondrina" is derived from the popular belief that the swallows 
which skim close to the ground upon which these spurges live are 
feeding on the seeds. 

Piura: Piura to Nomala, Weberbauer 5953 (var.). Cabo Blanco, 
Haught 150; 54. Lima: Surco, Ruiz & Pavdn. Near Lima, J. Ball. 
San Martin: San Roque, Williams 7681 ; 7327. Pongo de Cainarachi, 
King 2735. Junin: Chanchamayo, Isern 2345. Huanuco: Tingo 
Maria, Soukup 2270. Balsaplaya, Vargas 5345. Loreto: Yuri- 
maguas, Williams 3932; 4473; 5019. Rio Mazan, Williams 8165. 
Near Iquitos, King 346; 1240. Rio Nanay, Williams 278; 335. 
Rio Itaya, Williams 3334. La Victoria, Williams 2903. Cuzco: 
Hacienda Santa Rosa, Prov. Convention, Soukup 912. Warm and 
temperate Americas and widely introduced elsewhere. "Golondrina," 
"yerba Colorado," "urpai-mucuna" (Williams). 

Euphorbia huanchahana (Kl. & Gke.) Boiss. in DC. Prodr. 
15, pt. 2: 103. 1862. Tithymalus huanchahanus Kl. & Gke. Abh. 
Akad. Wiss. Berlin 1859: 71. 1860. T. penicillatus Millsp. Field. 
Mus. Bot. 4: 95. 1925. 

Glabrous (type from a fleshy cylindrical root branched at apex) ; 
stems low, dichotomously much-branched, squamose, the slender 
decumbent branches leafy; leaves subsessile, obovate, often minutely 
serrate, attenuate at base, obtuse, 4-6 mm. long, entire, the floral 
basally subcordate; peduncles equaling the campanulate involucres, 
throat of these with oblong lobes ciliate, glands dark red, ovate, 
subtruncate. The species proposed by Millspaugh seems to be the 
same and his observations may therefore be added to the description 
of Boissier: branches 7-14 cm. long, lower leaves 8-10 mm. long, 



FLORA OF PERU 21 

the upper ovate, apiculate, 12-15 mm. long, ciliate; glands orbicular, 
agaricoid, papillate, central stipe thick; styles about one-fourth 
bifid, penicillate; capsules depressed, trigonous, about 2 mm. high, 
2.5 mm. wide; seeds ovoid-quadrangular, bluish-white, finally choco- 
late-brown, 2 x 1.8 mm., smooth, the ventral facet with 1 sulcus 
and 2 protuberances (these not observed by me). But the seeds of 
other fruiting material in Peru seem to be smaller than those of 
specimens determined as C. chilensis. Croizat, Journ. Arnold Arb. 
24 : 179. 1943, has defined under T. huanchahanus two variants, one 
Argentinian, the other Bolivian and subsp. typica, the glabrous leaves 
serrate; presumably then E. chilensis, which compare, is restricted 
to the narrow-leaved plant; but the larger seeds noted by me were 
from Argentine specimens. 

Ruiz and Pavon noticed the roots (their sap viscid) used as a 
laxative and sold in the Lima market. 

Ancash: Huardz, Raimondi 256. Lima: Matucana, grassy slope, 
299 (type, T. penicillatits); Savatier 1356. Prov. Canta, Ruiz & 
Pavdn, type (E. tuberosa in herb.). Junin: Cerro de Pasco, grassy 
limestone slope, 3071 (det. Johnston, E. chilensis, reduced, 3 cm. 
high, root fusiform). Yauli, Weberbauer 3760. La Oroya, Weber- 
bauer 169. Cuzco: Paucartambo Valley, 3,500 meters, Herrera 
1371; 1389 (in herb, as E. penicillata) . Puno: Rocky siliceous 
slope, 4,200 meters, Pennell 18483. Tacna: Candarave, 2,900 
meters, Weberbauer 7387? (young). Bolivia; Argentina (Croizat). 
"Huachamccana," "huachancano," "chancano" (Ruiz & Pavon). 

Euphorbia hypericifolia L. Sp. PI. 1: 454. 1753; B23. E. lasio- 
carpa Klotzsch, Nov. Act. Acad. Leop.-Carol. 19, Suppl. 1: 414. 
1843. Chamaesyce hypericifolia (L.) Millsp. Field Mus. Bot. 2: 
302. 1909. E. boliviana Rusby, Bull. N. Y. Bot. Gard. 4: 442. 1907 
as to Peru. 

Apparently very similar to, if separable from E. brasiliensis and, 
if that is only a form of E. hyssopifolia with pubescent ovary, scarcely, 
of course, from the latter but as to plant of Klotzsch entire plant 
typically (but see below) crisply canescent, stems often divaricately 
branched, leaves oblong, rounded or subcordate at base, obtusish, 
or the narrower upper acute, usually all somewhat dentate; stipules 
shortly triangular; cymes rather densely corymbulose, floral leaves 
linear-lanceolate; involucres strongly hirtellous, glabrous within, the 
lobes triangular; appendages white, obovate, rather broader than 
glands, entire or 2-3-lobulate; capsules ovoid, typically appressed 



22 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

yellowish hirsute, carpels rounded; seeds ovate-quadrangular with 
a few transverse rugae. After Mueller. Wheeler, Proc. Biol. Soc. 
Wash. 53 : 10. 1940, confirms from fragments of type his earlier opinion 
that the plants of Linne" and Klotzsch are conspecific. Svenson, 
using the name of Klotzsch, Amer. Journ. Bot. 33: 460. 1946, "made 
a large number of collections to show the transition from pubescent 
plants with hairy capsules to plants perfectly glabrous throughout, 
erect to prostrate, from obvious annuals to perennials with thick 
roots; seeds usually elongate (1.1 x 0.45 mm.), the rather flat dorsal 
surface frequently white-coated when mature and usually with trans- 
verse ridges." These observations by so careful a student show that 
this species, if correctly identified as that of Klotzsch is variable and 
probably the plants maintained as species on these characters are 
variants of one. I have therefore cited all the Peruvian specimens 
that appear to belong here sens. lat. including at least E. brasilensis. 
Mathews 502, cited by Rusby as belonging to his species, no data 
and not seen, must have come from Peru; however, my 1298, deter- 
mined by Millspaugh as Rusby's species, seems, as determined by 
Johnston, to belong here. E. lasiocarpa has a strong resemblance to 
the North American E. Preslii, but has larger seeds and more 
prominent glands (Svenson). Diminutive young plants placed here 
could go to some other species as E. serpyllifolia Pers., W229, the 
involucres solitary, but apparently unknown south of Mexico. 

Piura: East of Cabo Blanco, Haught 107; 175. Cana Dulce, 
Haught 199. Talara, Beetle 26203. Cerro Prieto, Haught 151; 
(Haught & Svenson 11604}- Quebrada Mongollon, Amotape Hills, 
( Haught & Svenson 11519). Ancash: Lomas de Mongon, San Rafael, 
Goodspeed Exped. 9178. Lima: Beach near Supe, Beetle & Horton 
9084 (det. Johnston, E. nutans). San Lorenzo Island, Anderson. 
Canta, Pennell 14596. Atocongo, open sandy slopes, Pennell 14784. 
Lomas, Lurin, 5951. Chillon near Viscas, Pennell 14490 (det. 
Wheeler). Chosica, 505 (det. Millspaugh, C. brasiliensis). San 
Martin: San Roque, Williams 7009; 7443; 7798. Lamas, Williams 
6438. Chazuta, King 4015. Huanuco: Tingo Maria, Soukup 2265. 
Stony slopes, 2327 (det. Millspaugh, E. brasiliensis) ; 3216 (det. 
Wheeler; Johnston, E. brasiliensis); 3235 (det. Wheeler); 3218 (det. 
Johnston). Ambo, river flat, 3186; 3187 (det. Johnston). Rio 
Huallaga Canon, 4232 (det. Wheeler). Piedra Grande, 3679 (det. 
Wheeler; Johnston). Junin: Uspachaca, 1298 (det. Johnston). La 
Merced, Killip & Smith 23414 (det. Standley). Loreto: Yuri- 
maguas, Williams 4644 (det. Standley, E. brasiliensis). Apurimac: 



FLORA OF PERU 23 

Open sandy places, Goodspeed Exped. 10545 (det. Standley). 
Cuzco: Hacienda Lucre, Vargas 9758 (det. Standley). Near Cuzco, 
3,000-3,600 meters, Herrera. Ollantaytambo, Cook & Gilbert 4&*(f) 
Arequipa: Posco, Cook & Gilbert 57. Mollendo, Hitchcock 22405 
(orE. serpyllifolia?). South America to Mexico. 

Euphorbia hyssopifolia L. Syst. Nat. ed. 10. 1048. 1759; W140. 
Chamaesyce hyssopifolia (L.) Small, Bull. N. Y. Bot. Gard. 3: 429. 
1905. E. brasiliensis Lam. var. hyssopifolia (L.) Boiss. in DC. 
Prodr. 15, pt. 2: 24. 1862. 

Ordinarily erect glabrous-stemmed annual or sometimes more 
enduring, simple or branched from the base, a few cm. to several 
dm. high; stipules mostly united, usually as broad as high, with 
slightly lacerate margin; petioles 1-1.5 mm. long; leaves lanceolate, 
often falcate to oblong, sometimes a little pilose toward the oblique 
base, 5-30 mm. long, usually serrate; cyathia solitary in the upper 
bifurcations and in few-flowered leafy cymes; peduncles glabrous, 
to 2 mm. long; bractlets linear; involucres obconic, 0.7-0.9 mm. in 
diameter, glabrous unless within at base of attenuate lobes and on 
stipes; glands circular to elliptical, the white or reddish entire ap- 
pendages one-third as wide; flowers 4-15 per cyathium; ovary gla- 
brous; capsules broadly ovoid, strongly 3-lobed, 1.6-2.1 mm. long; 
seeds ovoid-subtriangular, 1-1.4 mm. long, chocolate-brown to gray- 
ish-white, low smooth ridges defining shallow depressions. After 
Wheeler. Probably should include E. brasiliensis as a variant with 
pubescent ovary; unfortunately Boissier, who did so, used the earlier 
name as the varietal. E. maculata L. has stems usually crisply 
pubescent at tip, smaller seeds finely rugose and is to be expected 
as widely distributed; but according to Wheeler (at one time) the 
true E. maculata could be E. hyssopifolia; see Fosberg, Bull. Torrey 
Club 74 : 332. 1947, however, for discussion and references pertaining 
to application of name, and Croizat, I.e. 75: 188. 1948, for a reply. 
E. ammannioides HBK., mentioned under E. serpens, might be 
sought here. Illustrated, Wheeler, pi. 656D, opp. p. 101. 

Peru: Scarcely in typical form; see note above. Temperate South 
and North America. 

Euphorbia laurifolia Juss. in Lam. Encycl. 2: 418. 1788; 
B106. E. Latazi HBK. Nov. Gen. & Sp. 2: 58. 1817 and E. cestri- 
folia HBK. I.e. 59, fide Diels, Bibl. Bot. 116: 104. 1937. E. Leh- 
manniana Pax, Bot. Jahrb. 26: 508. 1899, fide Diels, I.e. 



24 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Glabrous shrub 2-6 meters high, with terete stout cicatrose 
branches, apically foliate and axillary cymes, the upper peduncles 
with 3 heads; leaves crowded, shortly petioled, oblanceolate-oblong, 
entire, 1-1.5 dm. long, 2-3 cm. wide, at least the lower more or less 
spreading-deflexed; cyathia green, 5 mm. long, 8 mm. wide, primary 
lobes denticulate, hirtellous, 1.5-2 mm. long and wide, glands 
fleshy, 1-1.5 mm. long, 3.5 mm. wide, hirsute without, otherwise 
glabrous; stamens 25-30, filaments 4 mm. long, anthers yellow, 1-1.5 
mm. long; bracts 2-3 mm. long, lacerate-hirsute (after Diels). 

Cajamarca(?) : Near Jae"n(?), Weberbauer 6295. Apurimac: Near 
Curahuasi, rocky places, Vargas 9157; 9821 (det. Standley). Pincos, 
rainy-green shrubland, in rocks, 2,700 meters, Stork & Norton 10662 
(det. Standley). Puno: Ollachea, Prov. Carabaya, Vargas 6938. 
Without locality, Jussieu, type. To Colombia. "Latzi" (Bonpland, 
Colombian). 

Euphorbia melanocarpa Boiss. in DC. Prodr. 15, pt. 2: 41. 
1862. 

Completely glabrous with black ligneous stems; branches fili- 
form, decumbent, 7-15 cm. long, the internodes longer than the 
lower leaves, these in type 4-6 mm. long and wide, thickish, entire, 
ovate-orbicular, the lower petioled with obliquely truncate base, the 
gradually smaller upper cuneate at base; stipules triangular, coales- 
cent, shortly fissate; involucres solitary in the upper axils, typically 
purplish-black with oblong fissate lobes equaled by the pedicels, 
the transverse ovate porous-punctate glands black, the concolored 
(in Peru lighter) very narrowly appendaged margin repand; styles 
bifid; capsules glabrous, blackish-purple; seeds ovoid-tetragonous, 
obsoletely rugulose-tuberculate. The Peruvian material, my collec- 
tions determined by Millspaugh, has somewhat smaller leaves, 
shorter internodes and the dark coloring described for the type is 
less pronounced. 

Junin: Prostrate in heavy stony dry "wash," Tarma, 1010. 
Prostrate on firm stony river flat, La Oroya, 982. Near Huancayo, 
3,400 meters, open rocky hillside, leaves flat on ground, Killip & 
Smith 22023 (det. Killip). Ecuador. 

Euphorbia Meyeniana Klotzsch, Nov. Act. Acad. Leop.-Carol. 
19, Suppl. 1: 414. 1843; B42. Chamaesyce Meyeniana (Kl.) Croizat, 
Journ. Arnold Arb. 24: 184. 1943. 

Entire plant densely and crisply cinereous velutinous, the pro- 
cumbent stems divaricately branched; stipules minutely fimbriate; 



FLORA OF PERU 25 

leaves subsessile, cordate at base, ovate or oblong, to about 1 cm. 
long, obtuse, rather coarsely and acutely serrate; involucres dichot- 
omous and in upper axils, campanulate-turbinate, hirsute as the 
ovate lobes within and without, the ovate glands with white obovate 
much broader entire or subentire appendages; styles longer than 1 
mm., bifid, clavate apically; capsules hirtellous, cocci subcarinate; 
seeds oblong, acutely tetragonous, under lens finely papillose as well 
as rugulose. Var. viscidula Boiss. similar to type but glutinose. 
The type was from the Bolivian side of Lake Titicaca. In the 
scrap seen the appendages are slightly crenate. The Ecuadorean 
E. melanocarpa Boiss. in DC. Prodr. 15, pt. 2: 41. 1862, placed near 
this by the author and in general similar, is completely glabrous, 
while E. quitensis Boiss. I.e. p. 43 has very oblique leaves rounded 
at base, oblong, a little hirtellous at the stems. F.M. Neg. 18059. 

Arica: Lechler 1557 (type of the var. in herb, as E. maculata). 
Northern Bolivia; Paraguay. 

Euphorbia micromera Boiss. ex Engelm. Proc. Amer. Acad. 
5: 171. 1861; W194. Chamaesyce micromera (Boiss.) Woot. & Standl. 
Contr. U. S. Nat. Herb. 16: 144. 1913. 

A prostrate small-leafed annual with very slender dichotomously 
branched glabrous or pubescent stems and shortly petioled (petioles 
0.5 mm. long) obliquely rounded ovate or oblong obtuse entire 
leaves only about 3 (2-7) mm. long; stipules linear-lanceolate, cilio- 
late, the lower often united; involucres axillary, solitary, ovoid- 
globose, glabrous with ciliate lobes; glands pink or red, orbicular or 
oblong, unappendaged, or in pubescent plants with minute white 
margins; bracteoles reduced to solitary pink appendage; ovary 
glabrous to pubescent; carpels slightly grooved dorsally; capsules 
about 1.3 mm. long; seeds narrowly oblong, sharply tetragonous, 
1.1-1.3 mm. long, smooth or obsoletely 2-3 undulately rugulose. 
The specimen of Pennell seen has some leaves with a few denticula- 
tions, narrow but obvious gland appendages. Illustrated, Wheeler, 
pi. 658C, opp. p. 169. 

Piura: La Brea, (Haught & Svenson 11586). Shale cliff above 
sea, Paita, Pennell 14815, det. Wheeler; Haught 51. Talara, Beetle 
26202. Parinas Valley, Haught 116. Cajamarca: Near Cajamarca, 
Osgood & Anderson (very young; det. Millspaugh E. quitensis Boiss., 
43, but that with broad appendages). Mexico and southwestern 
United States. 



26 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Euphorbia Milii Des Moulin, Bull. Hist. Nat. Soc. Linn. 
Bordeaux 27-30. pi 1. 1826; B79. E. splendens Bojer ex Hook. 
Bot. Mag. pi. 2902. 1829. 

Erect to dependent or more or less scandent spiny shrub with 
small (1-5 cm. long) membranous subsessile obovate leaves, many 
conspicuous often brown-red spines on the similarly colored thick 
branches and long-peduncled cymes from the upper leaf-axils, the 
bright red bracts showy; involucre campanulate with fimbriate ovate 
lobes and fleshy red glands; seeds tuberculate. This is one of a 
number of plants which, in some part, may be fluorescent; here 
bluish-white and from the roots (see Goodwin and Kavanagh, Bull. 
Torrey Club 75: 1-17. 1948). Doubtfully ever adventive in Peru 
but its bizarre appearance makes its name of interest wherever 
cultivated. Croizat, Journ. Arnold Arb. 21: 506. 1940 and later, 
called attention to the correct scientific name which Boissier had 
indicated applies to the same species, but the native name of Mada- 
gascar seems somehow more suitable! Illustrated, I.e. and many 
others as Bailey, Stand. Cycl. Hort. 1170. 

Loreto: Cultivated at Pebas, Williams 1821. Rio Mazan near 
Iquitos, Williams 8137. Madagascar. "Corona de Cristo," "soongo- 
soongo" (Bojer). 

Euphorbia orbiculata HBK. Nov. Gen. & Sp. 2: 52. 1817; 
B31. 

Glabrous, branched from the base, the diffuse or ascending 
dichotomously much-branched filiform stems about 1 dm. long, 
shrubby below; leaves thick, rounded at base, orbiculate, 4-5 mm. 
wide, entire; stipules minute, denticulate, in type subulately incised, 
often subconnate into a solitary interpetiolar one; flowers axillary, 
solitary, subequaling the leaves; involucres dichotomous and 
terminal, the throat hirtellous, the red, transversely oblong glands 
equaled by the white, entire or obsoletely repand semi-orbicular 
appendages; styles shortly bifid; cocci carinate; capsules mostly 
glabrous; seeds blackish, obtusely ovoid-tetragonous, smooth. 

Peru (probably). Colombia; northern Bolivia. 

Euphorbia Peplus L. Sp. PL 456. 175S; B141. 

Glabrous annual, the greenish-yellowish stems often curved- 
ascending at base, the more or less promptly deciduous leaves slender- 
petioled, the many (in age) erect branches with sessile obovate very 
thin leaves, these usually 1-2 cm. long, acute at base, rounded at 



FLORA OF PERU 27 

summit, the uppermost verticillate; bracts deltoid or broadly ovate; 
involucres subsessile, campanulate, fimbriate; glands semilunate, 
much broader than long and provided with slender or filiform horns; 
capsules deeply sulcate, narrowly alate-carinate, the whitish-oblong- 
ovoid seeds with 1-4 transverse rows of darker gray pits. Young 
plants may be simple with 2-3 flowering branches at apex. Illus- 
trated, Kept. Mo. Bot. Gard. 11. pi. 30, and many others. 

Lima: In ditch, Killip & Smith 21667. Supe, in Salix thicket 
near seashore, Eyerdam 9069 (det. Johnston). Huanuco: Near 
Huanuco, Sawada P 61. Junin: Carpata above Huacapistana, 
Killip & Smith 24412. Cuzco: Yucay, Soukup 746. Naturalized 
from Europe and Asia. 

Euphorbia peruviana Wheeler, Contr. Gray Herb. 127: 59. 
pi. 3a. 1939. Chamaesyce arequipensis Croizat, Journ. Arnold Arb. 
25: 194. 1945. 

Often more or less spreading hispid-pilose or sometimes glabrous 
annual, the 1-4 erect or suberect stems 4-8 cm. high; stipules often 
obsolete or obscure, or 1-2 linear segments 0.6-1 mm. long; leaves 
approximate, sparsely pilose to glabrate, ovate, oval or oblong, 4-8 
(-15) mm. long, about as wide, subentire or irregularly dentate, 
petioles 1 mm. long; cyathia solitary (pedicels to 4 mm. long), 
cupulate-campanulate, 1.7-2 mm. in diameter, pilose; glands trans- 
verse-oblong, 0.8-1 mm. long, the one and a half to twice as wide 
white appendages irregularly dentate; styles 1.1-1.3 mm. long, one- 
third to one-fourth connate, bifid, the stigmas ellipsoid; capsules 
sparsely pilose, acutely 3-lobed, 2.3 mm. long, immature seeds 
quadrangular, about 1.7 mm. long, smooth. Distinguished from all 
other species of subgenus Chamaesyce by the extra long styles with 
thickened tips (Wheeler), but those of E. Meyeniana, known to me 
only from a scrap, seem to be the same. Also the Croizat plant (to 
my regret) appears to be identical except it is slightly more pilose; 
the distinctive (according to Wheeler) styles are equally obvious. 
A duplicate specimen by Metcalf has a firm ascending base but col- 
lector wrote "annual." 

Arequipa: Dry stream bed, Caraveli to Atico, 1,800 meters, 
(Metcalf 30341, type, C. arequipensis). Moquehua: Mount Estu- 
quina, Weberbauer 7441, type. 

Euphorbia Poeppigii (Kl. & Gke.) Boiss. in DC. Prodr. 15, pt. 
2: 56. 1862. Leptopus Poeppigii KL & Gke. Abh. Akad. Wiss. 
Berlin 1859: 46. 1860. 



28 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Erect, attaining several dm., alternately and dichotomously 
branched from often decumbent base, the branches dichotomous, 
glabrous, more or less reddish; leaves petioled, somewhat appressed 
pilose-hirsute, ovate, obtuse, the larger 10-12 mm. long; stipules 
minute, glanduliform; involucres shortly pedicelled in the axils and 
terminal, turbinate, hirsute without and in the throat, the oblong 
lobes fimbriate, the ovate glands with obovate, shortly 3-4-dentate, 
greenish-white appendages more than twice as long; styles short, 
bifid; capsules hirtellous; seeds ovoid. The larger, more openly 
branched var. laxa Boiss. has the leaves less pubescent and is of 
doubtful taxonomic merit. Determinations by Standley for Junin 
and Ayacucho except as noted. F.M. Neg. 32515 (var.). 

Huanuco: Piedra Grande near Muna, 3676 (det. Johnston). 
Pozuzo, 4595 (det. Johnston). Junin: Carpapata, Soukup 31*51. 
Huacapistana, Kittip & Smith 2260. La Merced, 5281 (det. Johns- 
ton); Killip & Smith 23436. Loreto: Yurimaguas, Poeppig 3118, 
type; Spruce 3908 (type, var.). Ayacucho: Aina, 22701; 23098. 
Ccarrapa, Killip & Smith 22427. Cuzco: Colpani, Urubamba 
Valley, Cook & Gilbert 1045; Vargas 755. Quellomayo, Prov. Con- 
vencion, brushy banks, West 8027 (det. Wheeler). Puno: Sandia, 
Weberbauer 509. Bolivia. "Monte huachanca" (Cook & Gil- 
bert). 

Euphorbia raphanorrhiza (Millsp.) Macbr., comb. nov. Tithy- 
malus raphanorrhizus Millsp. Field Mus. Bot. 4 : 95. 1925. 

Glabrous perennial, the few filiform branches prostrate from a 
fusiform root; stipules none; leaves petiolate, the lower scattered- 
alternate, crenulate, obtuse, the upper opposite, crenulate-serrate, 
acute to apiculate, all orbicular; inflorescence solitary in the axils, 
sessile, involucres campanulate, the oblong lobes fibrillate at apex; 
glands transversely ovate, thick, stiped, the stipe prolonged to the 
base of the tube; styles bifurcate one- third, the stigmas globose; 
capsules glabrous, deeply trigonous; seeds triangular ovoid, white, 
2 x 1.5 mm., the facets smooth, the angles sharp; caruncle or funicular 
vestigial, papyraceous papillate. Section Ipecacuanhae (Mill- 
spaugh). 

Lima: Steep, lichen-covered, western slope, Matucana, 85, type. 

Euphorbia Ruiziana (Kl. & Gke.) Boiss. in DC. Prodr. 15, 
pt. 2: 31. 1862. Anisophyllum Ruizianum Kl. & Gke. Abh. Akad. 
Wiss. Berlin 37. 1859. 



FLORA OF PERU 29 

Suffrutescent from subligneous rhizomes, the filiform or slender 
prostrate stems 1-1.5 dm. long, minutely pruinose hirtellous under 
a lens; leaves subsessile, glabrous, thick, 3-5 mm. long, to 4 mm. 
wide, subcordate, obovate, obtuse, minutely apiculate, entire or 
paucidentate apically; stipules ciliate; involucres axillary and 
terminal, solitary or 2-3, the throat as lanceolate lobes densely 
hirsute; glands ovate, puncticulate with short biparted appendages 
or unilateral, liguliform and twice as long as gland; capsules glabrous, 
cocci carinate, style elongate, deeply bifid; seeds oblong- tetragonous, 
in my specimens brownish, faintly striate, about 1.5 mm. long. 
E. Jamesoni Boiss. I.e. of Ecuador has larger leaves twice as long, 
appendages elegantly petaloid (Boissier). F.M. Neg. 18063. 

Lima: Lima and Chancay, Ruiz & Pavdn, type (E. striata in 
herb). Huanuco: Disintegrating shale and gravel slopes, 8213; 
3517; Ruiz & Pavdn (det. Mansfeld). 

Euphorbia serpens HBK. Nov. Gen. & Sp. 2: 52. 1817; W198. 
Chamaesyce serpens (HBK.) Small, Fl. Se. U. S. 709: 1333. 1903. 

Glabrous prostrate annual (sometimes enduring?), the slender 
stems to 5 dm. long and sometimes rooting at the nodes, these to 
3 cm. distant; stipules all united into a white membranous lacerate 
scale about equaling the less than 1 mm. long petiole; leaves ovate- 
orbicular to oblong, oblique at base, entire, 2-7 mm. long and wide; 
peduncles to 2 mm. long; cyathia solitary at nodes; involucres 
turbinate, 1 mm. long and broad, glabrous unless for a few trichomes 
on margins, acute or acuminate at least as long as the transversely 
oblong ochroleucous glands, their white, little wider appendages 
mostly crenate; fifth gland longer; flower parts completely glabrous 
including the 3-lobed ovary and markedly clavate styles, these 
usually more than medially parted; capsules about 1.2 mm. broad; 
seeds smooth, ovoid, the more or less prominent angles rounded, 
about 1 mm. long, 0.5 mm. in any diameter. After Wheeler. The 
more northern ranging E. ammannioides HBK., W128, with distinct 
stipules, appendaged glands, larger seeds, regularly of coastal sands, 
is to be expected in the Lomas. Illustrated, Wheeler, I.e. pi. 661C, 
opp. p. 192. 

La Libertad: Chicama Valley, in a garden, Smyth 84 (det. 
Wheeler). Lima: Prostrate mats on seaside plain, Callao, 5885 
(det. Johnston) ; also Dombey;Gaudichaud. Lurin, Pennell 12208 (det. 
Standley). To Paraguay, Mexico, Venezuela and introduced north- 
ward. 



30 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Euphorbia Spruceana Boiss. in DC. Prodr. 15, pt. 2: 53. 1862. 

Perhaps perennial but the several dm. tall flexuous glabrescent 
stems herbaceous; stipules conical, subulate; leaves petiolate, the 
upper 3-5-verticillate, all sparsely hirtellous, entire, ovate from 
rounded base, 18-22 mm. long, 14-16 mm. wide (Klug specimen, to 
5 cm. long, 3 cm. wide); cymes terminal, sometimes dichotomous, 
exceeding the upper leaves, the bracteolate branches 8-12 mm. long; 
floral leaves lanceolate-subulate; involucres narrowly turbinate, 
lateral mostly aborted, glabrous, the lobes lanceolate, the binate 
glands transversely ovate, concave as the slightly wider appendages; 
capsules 3 mm. wide, depressed, sparsely hirtellous, the cocci sub- 
carinate; seeds ellipsoid, deeply pitted, about 1 mm. thick, 1.3 mm. 
long (Klug). F.M. Neg. 18067. 

San Martin: Tarapoto, Spruce 4378, type. Juanjui, Klug 3917 
(det. Standley, E. eotinifolial). "Yuquillo" (Williams). 

Euphorbia tacnensis Phil. Anal. Mus. Nac. Chile, Bot. 77. 
1891. 

Erect pubescent little annual with few lower leaves, the upper 
opposite, oblong, equally acuminate both ends, shortly petioled, 
nearly glabrous, concolor; stipules none; cymes terminal, dicho- 
tomous, rather many-flowered; involucre glands pectinate and with 
white pectinate appendages; capsules glabrous; seeds coarsely 
tuberculate. After Philippi and obscure to me; maybe perennial? 

Tacna: Near Tacna, (Philippi, type). 

Euphorbia Tessmannii Mansf. Ber. Deutsch. Bot. Ges. 46: 
674. jigs. A &E. 1929. 

Unbranched glabrous 3-meter tree, the trunk of type 4 cm. in 
diameter; petioles about 6 cm. long; leaves alternate, oblanceolate, 
gradually narrowed to base, abruptly caudate-acuminate, 27-36 
cm. long, 7.5-10 mm. wide, drying pergamentaceous, marginally 
minutely revolute, rather densely pinnate-nerved, the lateral nerves 
about rectangular, scarcely conspicuous; inflorescence including the 
peduncle about 1.5 dm. long, laxly branched dichotomously, the 
ovate bracts soon deciduous, ultimate equaling the cyathium; 
involucre 3 mm. long, cyathiform, with 5 rotund-ovate lobes, pedicels 
3 mm. long, glands 5, depressed, shortly stiped, male flowers 5, 
the involucre with 2 broadly obovate exterior lacerate-dentate 
bractlets connate and laterally adnate to the included involucre; 
central female with 5-lobulate disk. Belongs to the Laurifoliae of 



FLORA OF PERU 31 

Section Tithymalus and nearest E. elata Brandegee of Mexico, with 
which it shares the distinctive character of partly adnate bractlets 
(Mansfeld). This description is from the author's, Notizbl. Bot. 
Gart. Berlin 11: 137. 1931. P.M. Neg. 5186. 

Loreto: Flood-free wood, mouth of the Santiago, Tessmann 4071, 
type. 

Euphorbia thymifolia L. Sp. PI. 454. 1753; W252. Chamaesyce 
thymifolia (L.) Millsp. Field Mus. Bot. 2: 412. 1916. 

Prostrate glabrate or pubescent annual (or more enduring?), the 
much-branched stems forming mats; petioles 0.5-1 mm. long; 
stipules about 1 mm. long, linear-subulate, entire or parted, distinct 
or nearly, fimbriate; leaves opposite, elliptic to narrowly oblong or 
ovate-lanceolate, rounded or acute, oblique at base, serrulate, nearly 
glabrous above, to 10 mm. long, ordinarily shorter; cyathia sub- 
sessile, mostly in small cymes of 2-several on short lateral branch- 
lets; involucres broadly obconical, soon distended by partly enclosed 
capsule, strigose above including the lobes, only the distal equaling 
the red glands; appendages twice as long to rudimentary; ovary 
densely pubescent; capsules subappressed puberulent; seeds sharply 
quadrangular, reddish, transversely rugose, 0.8-0.9 mm. long. A 
common tropical weed of both hemispheres, and found at Salinas, 
Ecuador, by Svenson. After Wheeler. Illustrated by him, pi. 655 A 
opp. p. 100. 

Peru: Without locality, Ruiz & Pavdn (det. Mansfeld). Coastal 
Ecuador and widely distributed in tropics. 

Euphorbia trancapatae (Croizat) Macbr., comb. nov. Chamae- 
syce trancapatae Croizat, Journ. Arnold Arb. 27: 290. 1946. 

Repent from stout root, often with many woody rosulate branches, 
the internodes 2.5 cm. long or shorter, all gray lanuginose; petioles 
less than 1 mm. long; stipules interpetiolar, acute, suberect, not 
abundantly lanate; basal leaves rotund, subentire, about 3 mm. 
long, the other (anisophyllous) leaves round-triangular to 10 mm. 
long, 5 mm. wide, serrate, gray or white lanate both sides; involucres 
solitary, about 2 mm. long, 1.5 mm. wide; 5 glands plicate, narrow 
appendage entire, pale rose or whitish; ovary pedicelled, hispid- 
lanate; styles short, bifid, stigmas minute. Certainly notE. boliviana 
Rusby. The young vigorous shoots identify it easily by their heavy 
pubescence; the leaf shape suggests small forms of E. hirta L. 
(Croizat). But compare the apparently too similar E. Meyeniana. 



32 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

However there seems to be a difference how constant to be proved 
by more collections in the shape of the merely undulate appendages. 
I am indebted to my friend Dr. C. Vargas C. for the loan of the 
original material. 

Apurimac: Cuzco boundary, Cuahuasi, Trancapata, 2,800 meters, 
stony slopes, Vargas 1257 (1237; type, Univ. of Cuzco). 

Euphorbia viatilis Ule, Verb. Bot. Ver. Brandenb. 50: 85. 1908. 

Prostrate filiform stems as all parts densely tomentose villous; 
stipules lanceolate-subulate, laciniate-hirsute, 1.5-2 mm. long; 
petioles 1 mm. long; leaves ovate-oblong, slightly inequilateral at 
subcordate base, obtusish or rounded, denticulate, about 6 mm. 
long, half as wide; involucres few, in axillary, nearly sessile cymes, 
turbinate, villous without, lobes lanceolate, fimbriate-hirsute, glands 
4, transversely elliptic, appendaged, the 2 larger 1.5 mm. wide, 
oblong- or ovate-elliptic, repand, 2 much smaller broadly elliptic, 
all yellowish- white; stamens 5-7; ovary and capsules villous, the 
latter 1 mm. high, 1.5 mm. wide; styles short, pilose, bifid, 1.5 mm. 
long; seeds red-brown, transversely and deeply 4-5-sulcate. A 
small low woolly annual with delicate branches, related toE. prostrata 
Ait., W265, staminate flowers 4, involucres axillary, seeds closely 
ridged (E. Chamaesyce L., see Croizat, Bull. Torrey Club 72: 213-318. 
1945) but woolly pubescent disk thick and appendaged, and more 
numerous stamens (Ule). It suggests tooE. dioica HBK., of Mexico 
and Central America with typically more numerous involucres, the 
appendages glabrous; and E. densiflora (Kl. & Gcke.) Kl., known 
from as near as Panama and apparently, ex char., unless more 
delicate, scarcely distinct. 

San Martin: Tarapoto, streets and dry places, ( Ule 6839, type). 

Euphorbia viridis (Kl. & Gke.) Pavon ex Boiss. in DC. Prodr. 
15, pt. 2: 62. 1862. Alectoroctonum viride [Pavon] Kl. & Gke. Abh. 
Akad. Wiss. Berlin 1859: 39. 1860. E. Mandoniana Boiss. I.e. 1264, 
at least as to Peru. 

Glabrous fleshy tortuous-stemmed and more or less ligneous, the 
flexuose dichotomous branches herbaceous; petioles about 1 cm. long, 
ternate in type; leaves ovate, rounded or acute at base, obtuse, 
l-2(-3) cm. long, the opposite floral lanceolate acutish; cymules 
terminal, forming a leafy panicle of 3-5 heads, the shortly pedicelled 
hemispheric involucres glabrous except the hirsutulous throat, the 






FLORA OF PERU 33 

ovate lobes dentate; glands usually 4, oblong, with broader subor- 
bicular denticulate greenish or reddish appendage; styles bifid; 
capsules deeply trisulcate, about 3 mm. broad, cocci rounded; 
seeds ovoid-subtetragonous, apically abruptly attenuate, irregularly 
foveolate and densely tuberculate. The allied E. haematantha M. 
Arg., B63, of Ecuador is lightly pubescent, with filiform procumbent 
stems, purple involucres and the glands with purple deeply 4-5- 
dentate appendages. My collections were referred by Millspaugh 
toE. Mandoniana, said to have seeds not tuberculate; probably this 
is variable in Peru at least. F.M. Neg. 32511 (Mandon). 

Piura: Stems black, leafy only one month, Cerro Prieto, Haught 
153; 154. Lima: Cheuchin, Ruiz & Pavdn, type. East of Sayan, 
1,550 meters, Prov. Chancay, Goodspeed 33028 (det. Croizat, E. 
Mandoniana}. Chosica, rocky hillside, branched only above, 2872; 
fleshy-woody among large rocks, 8 dm. high, 490. Matucana, 
woody at base, spreading in loose granite 277. Huariaca, 3127. 
Huanuco: Yanahuanca, 1245. Mufia, slender half shrub 1 meter 
high, brush-lands, 3943. Piedra Grande, loose soils, grass-shrub 
slopes, 3677; 3704. Bolivia? 

Euphorbia Weberbaueri Mansf. Repert. Sp. Nov. 29: 221. 
1931. 

Cactus-like shrub with succulent alternate lax branches, the 
ligneous obtusely longitudinally carinate-costate erect or suberect 
branchlets 5-8 mm. in diameter; innovations articulate; leaves 
rudimentary, scale-like, 1.5 mm. long, 1 mm. wide, ovate-triangular; 
cymes very short from lateral axillary pulvinus, cyathia mostly 3, 
subsessile, basal foliar leaves 2, opposite, 2-8 mm. long, glabrous, 
the obovate lobes lacerate; glands 4, transverse-elliptic or subsemi- 
lunate, concave, obscurely bilabiate, unappendaged, very shortly 
stiped; capsules glabrous, 4 mm. in diameter; cocci dorsally obtusely 
carinate; seeds small, brown, verrucose or nearly smooth. Placed 
by the author in a species-group which he defines to include the 
interesting Brazilian species E. phosphorea Mart., B176, named in 
reference to the phosphorescent sap; however Croizat, Journ. Arnold 
Arb. 24: 182. 1943, thinks the plant of Martius requires another 
position. 

Cajamarca: Open areas in cactus-shrub formation, 900 meters, 
Weberbauer 3802, type. Side valley to the right of the Rio Huanca- 
bamba, rain-green formation, Weberbauer (without number). 



34 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

2. PEDILANTHUS Necker or Poiteau in 1812 
Reference: Millspaugh, Field Mus. Bot. 2: 353-377. 1913. 

Erect copiously lactiferous shrubby plants more or less fleshy 
above and often nearly or quite leafless, the alternate entire succulent 
leaves usually falling soon after the appearance of the oddly shaped 
monoecious flowers, which actually owe their curious appearance 
to the ordinarily shoe-shaped and colored enveloping involucre. 
Involucres in terminal or axillary cymes, cleft on upper side, bilabiate, 
the tip of lower lip bifid and the fissure more or less closed by 3 lobes, 
the tube at base above with a bifid or 2-3-lobed spur-like appendage. 
Flowers pedicellate, the male numerous, the solitary female with 
finally exserted usually declinate style with 3 often apically distinct 
stigmas. Ovary 3-celled, ovules solitary. Fruit capsular, the 
seeds ecarunculate. Croizat, Journ. Wash. Acad. Sci. 33: 19. 1943, 
expressed the opinion that "the cyathium is homologous with a 
much coarctate inflorescence of Dalechampia, the upper part of the 
inflorescence which bears glands and male flower being replaced by 
a chamber with glands in Pedilanthus; the genus can readily be 
keyed on floral characters." But these, according to Standley & 
Steyermark, Fieldiana: Bot. 24, pt. 6: 141. 1949, are of doubtful 
significance. 

Pedilanthus retusus Benth. in Hook. Journ. Bot. Misc. 6: 321. 
1854; 355. 

Glabrous shrub a meter or so tall; leaves subsessile at the cuneate 
base, obliquely ovate, retuse, lightly carinate beneath, 2.5-5 cm. 
long, 2-3.5 cm. wide; cymes short, terminal and in the upper leaf- 
axils; involucres about 9 mm. long; glands 4, upper truncate lip 3 
times shorter than tube; upper lobes of lower lip glandular; pedicels 
of both male and female flowers hirsute; capsule 6-7 mm. long, 9 mm. 
thick, the cocci rounded, the seeds subcarinate dorsally. Near the 
widely distributed, sometimes cultivated P. tithymaloides (L.) 
Poiteau, with acute or merely obtuse leaves and the pedicel of the 
female flower glabrous. 

Loreto: Iquitos and vicinity, Williams 3567; 8133. Upper 
Amazonian Brazil. "Zapatillo del Nino," "zapatito del Nino." 

3. PHYLLANTHUS L. 

Reference: Mueller, Argoviensi, in DeCandolle Prodr. 15, pt. 2: 
274-436. 1866. 



FLORA OF PERU 35 

Various in habit (one species aquatic), herbaceous or ligneous, 
with entire usually alternate often distichous leaves that on the 
branchlets may appear to be pinnate, the petioles short or wanting. 
Flowers usually monoecious, solitary or in clusters in the axils, 
sometimes in short inflorescences, apetalous, with 4-6 imbricate free 
or nearly free sepals, the male flower with disk glands free or annulate, 
stamens 3-6, filaments sometimes connate, ovary rudiment always 
absent, the female disk often urceolate, ovary 3-5-celled, styles free 
or connate, 3, bifid or lobed; ovules 2. Capsules dry at least at 
maturity separating into bivalved cocci, the ordinarily 3-angled 
seeds ecarunculate. 

Without a modern revision, greatly needed, this compilation is 
no doubt incomplete. Besides the following, the distinctive shrub 
or tree of India (P. acidus (L.) Skeels), known in English cultivation 
as "Indian gooseberry" from the fleshy juicy green or yellowish 
fruits, may be planted in Peru since it is not uncommon in tropical 
gardens; it is a shrub or small tree with leaves 3-5 cm. long and 
wide and unlike other Peruvian species has the flowers in raceme- 
like panicles clustered on the stems. 

Herbs, in Peru annuals or rarely more enduring and the stems firm 
or subligneous only toward the base; leaves mostly 1 cm. long 
or shorter. 

Aquatic, Salvinia-\\ke on slow waters P. fluitans. 

Terrestrial. 
Leaflets nearly round; anther connective often accrescent. 

P. orbiculatus. 

Leaflets nearly oblong; anther connective not enlarged. 
Stems subterete above, neither bi-ridged nor margined, often 

with several short spreading branches. 
Leaf nerves obvious; filaments more or less free; seeds 

minutely verruculose-scabrous P. lathyroides. 

Leaf nerves faint or obscure; filaments connate; seeds 

faintly lineate P. Niruri. 

Stems compressed or bi-ridged or margined above, often with 
several ascending, about equal branches from near base. 

P. compressus. 
Shrubs or half shrubs, the stems ligneous at least below the flowering 

portions; leaves mostly or all 2 (1.5) cm. long or longer. 
Leaves mostly or all 2-4 cm. long or shorter; stems and virgate 
branches about equally slender; styles bifid or parted. 



36 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves acute to acuminate both ends or at least apically. 
Flowers in axillary clusters; anther chinks oblique or hori- 
zontal. 
Leaves oblong-obovate, mostly or all about three times 

longer than wide P. Pavonianus. 

Leaves ovate, mostly or about twice longer than wide 
(species critical, concomitance and constancy of 
characters unproved). 

Styles appressed-recurved; ovary glabrous, smooth; 
glands of male flowers 3; leaves subglabrous, acumin- 

ation marked P. Mexiae, P. acuminatus. 

Styles usually rather more diffuse; glands of male flowers 
6, more or less connate; leaves acute or scarcely 
acuminate. 
Ovary papillose; connective free; leaves pubescent, 

obscurely acuminate P. brasiliensis. 

Ovary glabrous; connective connate below; leaves 

often pubescent in Peru, acute . P. pseudo-conami. 

Flowers (female) in axillary racemes; anther chinks vertical. 

P. racemigerus. 

Leaves rounded both ends, oval or oblong-elliptic. 
Calyx segments entire except in P. rubellus. 
Leaves orbicular-elliptic, membranous; pedicels, at least of 

female flowers, elongate. 

Leaves about 2 cm. long; pedicels several. .P. graveolens. 
Leaves about 3 cm. long; pedicels (female) solitary. 

P. anisolobus. 
Leaves oblong-elliptic, coriaceous; female flowers 2-3 on 

pedicels 2 mm. long P. rubella. 

Calyx segments lacerate-fimbriate P. lacerilobus. 

Leaves mostly or all about 7 cm. long or longer when grown; 
styles entire, little connate, in P. Poeppigianus or connate in 
P. grandifolius. 
Leaves rotund or broadly elliptic, rounded or little narrowed to 

the obtuse tip P. huallagensis. 

Leaves oblong-elliptic, suboblong or ovate, more or less narrowed 

to the usually acute tip. 

Leaves nearly oblong, rounded, truncate or subcordulate at 
base, often about four times longer than wide; ovary 
3-celled; stamens usually 6-9. 



FLORA OF PERU 37 

Branchlets and leaves beneath, at least prominent nerves, 
puberulent; stigmas stiped; seeds costulate. 

P. salviaefolius. 

Branchlets as leaves typically glabrous or essentially; stig- 
mas sessile; seeds smooth P. grandifolius. 

Leaves various but not suboblong, usually acute at base and 

about three times longer than wide; stamens 2-4. 
Male flowers clustered along short often leafless branchlets; 
calyx segments 4 or 6; ovary (known) 4-5-celled; 
styles bifid. 
Male calyx segments 6; leaves lustrous above, subcoria- 

ceous P. Vincentae. 

Male calyx segments 4; leaves opaque or subopaque, sub- 

chartaceous P. nobilis. 

Male flower clusters all axillary; calyx 5-parted; ovary 3- 
celled; stamens and styles 3, the latter nearly free, 
entire P. Poeppigianus. 

Phyllanthus acuminatus Vahl, Symb. Bot. 2: 95. 1791; 381. 

Much-branched arborescent shrub, sometimes 7 meters high, the 
angulate branchlets lineately puberulent with hyaline trichomes, the 
flowering usually elongate, herbaceous; petioles 1-2 (4) mm. long; 
stipules linear-lanceolate; leaves ovate or oblong-ovate, acute at 
base, more or less acuminate, mucronate, membranous, often 3-4 
cm. long, 1-2 cm. wide, glabrous unless midnerve and ciliate margins; 
flowers monoecious, the female few, the male in axillary clusters 
with linear-ovate scarious sometimes lobate bracts; male flowers on 
capillary pedicels about 2 mm. long, the glands 3 opposite the 3 
outer sepals; stamens 3, the filaments connate, anthers spreading; 
female pedicels 5 mm. long, elongating to 15 in fruit, apically clavate; 
outer sepals broadly ovate; disk glands 3, connate into irregular or 
subentire cup; ovary glabrous, smooth; styles free, bifid, short, 
appressed; capsules 3-4 mm. long, 3-sulcate, net-veined, the reddish- 
brown seeds carinate. According to Boissier the herb, name P. 
foetida of Ruiz and Pavon is this plant. P. Mexiae Croizat, Journ. 
Wash. Acad. Sci. 33: 14. 1943, is similar but female disk glands are 
distinct. Compare Croizat for a discussion of this and related 
species, Caldasia 2: 129-131. 1943. Illustrated, Mart. Fl. Bras. 11, 
pt. 2: pi 7, fig. 2. 

Cajamarca: Prov. Jae"n, Raimondi (det. Mansfeld). San Martin: 
Tarapoto, (Sprite 4928). Huanuco: Pozuzo, Ruiz & Pavdn (det. 



38 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Boissier). Junin: Chanchamayo, Isern 2176. South America; West 
Indies; Mexico. 

Phyllanthus anisolobus M. Arg. in DC. Prodr. 15, pt. 2: 382. 
1866. 

Shrub, apparently entirely glabrous but fruits and male flowers 
unknown; slender lightly angled 10-13-leaved branchlets a dm. or 
two long; stipules triangular-lanceolate; leaves orbicular-elliptic, 
acutish at base, subrotund-mucronulate at tip, 3-3.5 cm. long (as 
known), 22-28 mm. wide, thin-membranous, finely reticulate-veined, 
paler beneath; female flowers solitary, the strict filiform pedicels 
enlarged toward apex, 8-12 mm. long; 3 outer calyx divisions 
elliptic-lanceolate, 2.5 mm. long, the inner 3 much broader, ovate, 
equally long, all paler on margins; hypogynous disk crenate, ovary 
smooth with shortly incurved bifid styles. Thought by the author 
to be related to P. micrandrus M. Arg. of Venezuela with smaller 
leaves and calyces, but compare P. graveolens. F.M. Neg. 8498. 

Peru: Without locality, Pavdn, type, Herb. Boissier. 

Phyllanthus brasiliensis (Aublet) Poir. Encycl. 5: 296. 1804; 
383. Conami brasiliensis Aublet, PI. Guian. 927. pi. 354. 1775. 
P. Conami Swartz, Prodr. 28. 1788. 

Similar to P. acuminatus which probably should be drawn to 
be included as a variant but according to Croizat leaves ellipsoid, 
not manifestly acuminate, prevailingly pubescent, ovary papillose 
to armed with soft processes; styles effuse, long; disk under the ovary 
prevailingly of discrete glands, the male glands connate. 

San Martin: Tarapoto, Spruce 3961 (det. Mueller). Pongo de 
Cainarachi, Ule 6408. Cuzco: Valle de Urubamba, 1,600 meters, 
(Hen era 8621). To northern South America. 

Phyllanthus compressus HBK. Nov. Gen. & Sp. 2: 109. 1817; 
390. 

Erect annual, or more enduring and the stem ligneous below, 
often bushy above to about 3 dm. high, completely glabrous; younger 
branches strongly compressed; stipules cordate at base, produced 
below on one side; leaves subsessile, oblong-elliptic or -obovate, 
acute or obtuse at base, obtuse or rounded at apex, mostly 6-14 mm. 
long, 3-4 (5) mm. wide, opaque, distinctly nerved; flowers monoecious 
on very short pedicels or subsessile; fruiting calyx about 2 mm. 
broad, disk glands urceolately connate, in male flowers free; ovary 



FLORA OF PERU 39 

smooth; styles slender, bilobed, reflexed; seeds brown, lineately 
scabrous. In the similar P. hyssopifolioides HBK. known from 
Brazil the seeds are smooth, merely foveolate, flowers 1-3 in the 
axils, leaves 6-7 mm. long, 3-3.5 mm. wide, the nerves faint. Stand- 
ley has given an herbarium name to Mexia 6154 without any indica- 
tion of distinctive characters; as there are already several names 
for similar plants a critical study of them all seems advisable. 

San Martin: Near Tarapoto, Spruce 4173 (det. Mueller). Loreto: 
River bank at mouth of the Santiago, Mexia 6154? Central America; 
southern Mexico. 

Phyllanthus fluitans Benth. ex M. Arg. Linnaea 32: 36. 
1863; 400. 

Small floating species simulating in habit and form of leaves the 
water fern Salvinia; stems 3-5 cm. long; rootlets many; stipules 
acuminate, 2 mm. long; leaves sessile, cordate-orbicular, slightly 
emarginate, about 12 mm. wide, pellucid-punctate; flowers 2-4 in 
the axils, nearly 1.5 mm. long, shortly pedicellate, calyx segments 
suborbicular, filaments and glands of the male free, the latter in 
female flowers urceolately connate; styles bifid, the branches sub- 
erect; capsules depressed-globose, nearly 3 mm. wide, the seeds 
smooth dorsally. 

Loreto: Floating near shore of Isla Cacao, Rio Amazon, Ramon 
Castilla, F. J. Hermann 11314-5. Brazil. 

Phyllanthus grandifolius L. Sp. PL 981. 1753; 329. 

A tall shrub or small tree with elongate spreading-drooping 
branches, the compressed angulate branchlets more or less puberulent 
as often the leaves beneath, at least within Peru; stipules ovate, 
reflexing, about a third as long as the 4-7 mm. long petioles; leaves 
ovate-lanceolate or oblongish, obtuse or subobtuse at base, acuminate 
or obtusish, 7-15 cm. long, 3-5 cm. wide or also half the size and 
proportionately broader; fascicles 2-12-flowered, commonly with 
1 terminal female flower, the pedicels finally about equaling the 
petioles; disk in both sexes elongate-urceolate; male calyces ac- 
crescent, becoming 3-4 mm. long; stamens usually 6; filaments con- 
nate; style column stout, the sessile stigmas subpetaloid; ovary 
3-celled; capsules depressed, deeply sulcate, glabrous, 8 mm. long, 
12 mm. broad; seeds 5-6 mm. long, smooth. A variant with hirtel- 
lous- or tomentellous-papillose leaves beneath, var. cornifolius 
(HBK.) M. Arg., type from Guayaquil, may rather be P. salviae- 



40 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

folius; the Ruiz and Pavon specimen so determined by Mueller was 
probably, if the same, from that locality. Determinations mostly 
by Mansfeld. There may be some question as to the correct applica- 
tion of the name. 

San Martin: Rumisapa near Tarapoto, Williams 6768. Loreto: 
Lower Rio Huallaga, Williams 4831. Puerto Arturo, Williams 5190. 
Fortaleza, Williams 1>220. Santa Rosa, Williams 4880. To Mexico, 
Brazil, Guianas and the West Indies. "Gallinazo-panga" (Williams). 

Phyllanthus graveolens HBK. Nov. Gen. & Sp. 2: 112. 1817; 
383. 

Shrub with slender lutescent-puberulent branchlets; stipules 
linear-lanceolate, submembranous; leaves orbicular-ovate or broadly 
elliptic, rounded-acutish at base, rounded-obtuse, sometimes min- 
utely mucronulate at tip, 12-20 mm. long, 10-15 mm. wide, thin- 
membranous, glabrous or very sparsely puberulent both sides; 
flowers all monoecious, the few female with the male, or often lacking, 
the pedicels shorter than the leaves, usually 4 together, the former 
glabrous, the latter scabrous; male calyx segments broadly obovate, 
thick connate glands papillose, anthers connate; ovary glabrous and 
smooth; styles rather effuse. P. Millei Standley of Ecuador is 
similar but leaves as to type somewhat smaller and puberulent as 
those of P. pseudo-conami. F.M. Neg. 5027. 

Tumbez: Near Tumbez, Weberbauer 7644- Cajamarca: Tome- 
penda, Bracamora, Bonpland, type. 

Phyllanthus huallagensis Standl. ex Croizat, Journ. Wash. 
Acad. Sci. 33: 13. 1943. 

Glabrous tree, older bark much lenticellated and fissured; 
stipules triangular, small; petioles stout, 2-3 mm. long; leaves 
rotund-elliptic, rounded both ends or very shortly and obtusely 
acuminate, on vigorous branches 9-12 cm. long, about 7.5 cm. wide, 
on branchlets about 6 cm. long, 4 cm. wide, chartaceous, the 6-8 
slender lateral nerves prominent beneath, the reticulate venation 
moderately so; inflorescence to 2.5 dm. long, axillary and sub- 
terminal, the slender racemes with many clustered male flowers 
on filiform pedicels (to 10 mm. long) from a pulvinate base, the 
accompanying female flowers few; male flowers 4 mm. wide, lobes 
6, 2 mm. long, alternating 6 glands, stamens 3, connate into a column 
1 mm. long; female flowers 5-6 mm. wide on pedicels 3.5 mm. long, 
the 6 glands curved and almost connate beneath the globose ovary; 



FLORA OF PERU 41 

styles 3, short, reflexed except at tip. Section Elutanthos Croizat, 
I.e. 12, otherwise Mexican and Central American and typified by 
P. glaucescens HBK., distinctive by virtue of pulvinate bases to- 
gether with delicate male flowers. 

San Martin: Juanjui, Klug 4240, type. 

Phyllanthus lacerilobus Croizat, Caldasia 3: 21. 1944. 

Shrub, sparsely and shortly lax-pubescent; stipules 3, lateral 2 
broadly triangular, 1.5-2 mm. long, central more slender; flowering 
branches to 15 cm. long with 10-12 puberulent leaves each side, 
gray-olive, elliptic, rounded or barely narrowed at tip or truncate- 
retuse and mucronate, somewhat oblique to rounded base, (l-)2 cm. 
long, (5-) 10 mm. wide, the slender veins about 4 pairs; petioles 
very short; stipels setaceous; flowers solitary in axils of the upper- 
most leaves, commonly 3-5 and sometimes also in several of the 
axils toward the branchlet bases; male perianth 5 mm. wide, the 4 
(5) lobes 2 mm. long, 1.5 mm. wide, marginally lacerate-fimbriate; 
pedicels to 7 mm. long; stamens 4, coalescent into column; glands 
4 (5), contiguous, margin lobate; female perianth 8 mm. wide, the 
6 ovate laciniate lobes 3 mm. long, 2 mm. wide, disk glands 6, plicate; 
ovary glabrous, styles 6, 0.75 mm. long; pedicels apically compressed, 
6-7 mm. long. Section Eriococcus M. Arg. 420, fide author. Com- 
mon as this appears to be in the vicinity of Iquitos it is strange if 
an earlier name does not exist; cf. P. rubellus. 

Loreto: Caballo-Cocha, Williams 2878, type. Pebas, Williams 
1928. Iquitos, Williams 7896. Mishuyacu near Iquitos, Klug 2567 
(n. sp. Standley, in herb.). "Arbol de Navidad." 

Phyllanthus lathyroides HBK. Nov. Gen. & Sp. 2: 110. 1817; 
403. P. filiformis Pavon ex Baillon, Adansonia 1: 29. 1860, fide 
M. Arg. 

Erect more or less lignescent slender annual (?) sometimes several 
dm. high and more or less branched, the branches slender or filiform, 
terete, or slightly compressed, glabrous; stipules linear-lanceolate; 
petioles 0.5-1 mm. long; leaves distichous, oblong or oblong-elliptic, 
obliquely subcordate at base, commonly rounded-obtuse and apicu- 
late at apex, 6-14 mm. long, 2.5-4 (6) mm. wide, pale green, glauces- 
cent and with 4-5 or 6 pairs of rather conspicuous nerves beneath; 
flowers monoecious, the solitary or geminate male on slender pedicels 
about 2 (-5) mm. long, sepals broadly ovate, subobtuse, disk glands 
free, papillose; stamens 3, connate below; female pedicels 4-5 mm. 






42 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long in fruit, sepals oblong-obovate, venose, disk connate, undulate; 
ovary smooth, the 3 styles bifid; capsules 2.5-3 mm. in diameter, 
the seeds densely scabrous with many longitudinal rows of minute 
points. The type of P. filiformis was collected by Tafalla near 
Guayaquil. The similar and also widely distributed P. Urinaria 
L. is distinguishable by the minutely ciliolate leaves and the minutely 
tuberculate ovary, the seeds transversely rugose. P. carolinensis 
Walt, has somewhat obovate cuneate-based leaves and usually one 
flower of each sex subsessile in their axils; found as near as Galapagos 
where ligneous below. F.M. Neg. 29353 (P. filiformis). 

Cajamarca: Nancho, Cascas and Cutervo, Raimondi. Ayacucho: 
Ccarrapa, wooded hillside, Killip & Smith 22438. Loreto: La 
Victoria on the Amazon, Williams 2729. Yurimaguas, Poeppig 
1655 (det. Mueller). Tropical America to Mexico and the West 
Indies. "Piedra con piedra" (Williams), "orilla del riachuelo" 
(Raimondi). 

Phyllanthus Mexiae Croizat, Journ. Wash. Acad. Sci. 33: 14. 
1943. 

Apparently much like P. acuminatus and possibly supplanting 
that in Peru; male flowers to 2 mm. wide, glands 3 about base of 
the 3 connate stamens, pedicels 3-5 mm. long; female flowers about 
7 mm. wide, pedicels to 10 mm. long, lobes 6, glands distinct, erect, 
incurved, more or less regular, styles short, reflexed. Described as 
entirely glabrous but specimen seen pubescent as P. acuminatus; 
the author points out that true leaves may be present in some 
groups of Phyllanthus; here he regards them as transformed into 
stipules and the leaves, so-called, actually foliose bracts on the 
flowering axis. 

The collector of the type found it used as a fish poison. 

Peru: According to Croizat, Caldasia 2: 130. Ecuador. "Bar- 
basco." 

Phyllanthus Niruri L. Sp. PI. 981. 1753; 406. 

Much like P. lathyroides; stipules subulate from a broad scarious 
base; leaves rounded at both ends, 6-15 mm. long, 2-6 mm. wide, 
the lateral nerves inconspicuous; flowers solitary or one of each sex 
in the same axil; male sepals 5 or 6 as small disk glands, filaments 
connate; female pedicels in fruit only 1-2 mm. long, sepals 6, oblong, 
membranous; capsules 1.5-2 mm. in diameter, 3-sulcate; seeds with 
5-6 faint longitudinal dorsal lines. Like the related species this is 



FLORA OF PERU 43 

a common tropical weed. Determinations by Standley. The similar 
P. diffusus Klotzsch, 409, also widely distributed but always in wet 
places usually has stems basally thickened, smaller leaves, narrower 
stipules, female sepals and the seeds with many transverse lines in 
addition to the dorsal ones. 

San Martin: Pongo de Cainarachi, Klug 2767. Tarapoto, 
Williams 6051. San Roque, Williams 7124- Huanuco: Ruiz & 
Pavdn. Loreto: Yurimaguas, Williams 4466; 5036. Pastures and 
fields, Iquitos, Wittiams 7929; Killip & Smith 26890. Clearing, 
Rio Putumayo, Klug 1735. Widely distributed in the tropics. 
"Chanca piedra" (Williams). 

Phyllanthus nobilis (L. f.) M. Arg. in DC. Prodr. 15, pt. 2: 
414. 1866. Margaritaria nobilis L. f. Suppl. 428. 1781 (excl. male 
pi.). P. antillanus (Juss.) M. Arg. Linnaea 32: 51. 1863. Cicca 
antillana Juss. Euphorb. Tent. pi. 4, fid- 13B. 1824. 

A shrub or small tree sometimes 14 meters high; leaves subsessile, 
elliptic or lanceolate-elliptic, usually 6-13 (18) cm. long, 2.5-5 (7) 
cm. wide, subchartaceous, glabrous or sometimes puberulent beneath; 
flowers dioecious, greenish-white, the male fascicled along a 5-15 
mm. long peduncle, their pedicels 3-5 mm. long, the female solitary 
or 2-4-nate in the axils, the thicker pedicels often 1-1.5 cm. long; 
both sexes with 4 roundish or elliptic sepals in 2 series and with 
annulate fleshy disk; male flowers with 4 free stamens; female with 
4-5-celled ovary, 4-5 thick basally connate styles shortly 2-cleft; 
capsules about 13 mm. long and thick, at first somewhat fleshy; 
seeds usually 4, complanate, trigonous, smooth, olive-green, 3 mm. 
long. Not clearly distinct even as variants are P. nobilis var. 
peruvianus M. Arg., leaves cuspidate-acuminate, fruiting pedicels 
little longer than the mostly 5-celled capsules, and var. Pavonianus 
(Baill.) M. Arg., similar but capsules mostly 4-celled, styles deeply 
biparted. Wheeler, Contr. Gray Herb. 124: 35. 1939, employs the 
name of Jussieu on the ground that that of Linnaeus filius was based 
on a mixture. 

Fruit eaten by Indians, said to be bluish when mature (Mexia). 

San Martin: Tarapoto, Spruce 4472 (var. peruvianus). Juanjui, 
Klug 3812 (det. Standley). Pongo de Cainarachi, Klug 2686; 
2695 (n. sp. in herb., Standley). Without locality, Ruiz & Pavdn 
(var. Pavonianus). Loreto: Florida, Klug 2013?; 2338. Rio 
Santiago above Pongo de Manseriche, Mexia 6140; 6296 (det. 
Standley); Tessmann 3978. Puerto Arturo, Williams 5145. Mouth 



44 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

of Rio Blanco, Tessmann 3046 (det. Mansfeld). Middle Ucayali, 
Tessmann 3270 (det. Mansfeld). Rio Acre: Krukoff 5663 (det. 
Croizat with query). Tropical South America to Mexico and the 
West Indies. "Loronaue," "antachibuca" (both Indian, Mexia), 
"ucariviro" (Tessmann). 

Phyllanthus orbiculatus Rich. Act. Soc. Hist. Nat. Paris 1: 
113. 1792; 401. Orbicularia orbiculata (HBK.) Moldenke, Rev. 
Sudani. Bot. 6: 178. 1940. 

Clearly annual, erect, with usually many always very slender 
branches, these sometimes branched, several to 10 cm. long; stipules 
triangular, acute; petioles 1-2 mm. long; leaves about rotund, 
usually around 5 mm. wide, sometimes twice as large, rather truncate 
at base, often mucronulate, glaucous beneath, membranous, the 
lateral nerves 3-4; pedicels axillary, 1-2 in the axils, nearly filiform 
or the female slightly clavate, 2 mm. long, or the fruiting to 6 mm. 
long; sepals 6, oblong or the narrower female acute; stamens 3, 
nearly free, the 6 crenate disk glands free; ovary 6-sulcate, styles 3, 
bifid; capsules about 2.5 mm. wide, compressed, the trigonous seeds 
longitudinally punctate-scabrous in rows, 1.5 mm. long. 

Piura: Shaded rocks, slope of Loma Prieto (Haught & Svenson 
11631, det. Svenson as affine). Junin: Sandy shady river bank, 
La Merced, 5396 (det. Johnston); Killip & Smith 23768. Chancha- 
mayo Valley, Schunke 617; 618 (det. Standley). Cuzco: In open 
weedy growth near river, Echarate, Goodspeed Exped. 10^72 (det. 
Standley). Brazil; Guianas; Venezuela. 

Phyllanthus Pavonianus Baillon, Adansonia 1: 30. 1860; 380. 
P. oxycladus M. Arg. Linnaea 32: 26. 1863, fide M. Arg. 

Branches subterete but the branchlets acutely and regularly 6 
wing-angled, 2 angles from the petioles, 4 from the stipules, these 
lanceolate, not auriculate, scarious-margined; leaves distichous, ob- 
long-obovate, acute both ends or apically obtusish, 1.5-2 cm. long, 
4-9 mm. wide, ochraceous puncticulate beneath; flowers monoecious, 
both sexes fascicled in axils, the male 2-5 on pedicels 3-4 mm. long, 
the female 1-2 on pedicels 5-8 mm. long; calyces 1-2 mm. long, 
fruiting not at all accrescent; disk urceolate in both sexes, distinctly 
crenate or lobed, that of the male flowers soon unevenly 3-parted; 
stamen column entire, short, stout; ovary smooth, glabrous, the 
biparted styles spreading; capsules nearly 3 mm. in diameter. A 
Ruiz and Pavon collection at Madrid was labeled as from Guayaquil 



FLORA OF PERU 45 

and was made by Tafalla. Apparently comparable or similar is 
P. Haughtii Croizat, Caldasia 3: 22. 1944, with pedicels of male 
flowers only 2 mm. long, the 3 disk glands separate. F.M. Neg. 
24586. 

Peru (perhaps, cf. note above). Ecuador. 

Phyllanthus Poeppigianus M. Arg. in DC. Prodr. 15, pt. 2: 
323. 1866. Glochidion Poeppigianum M. Arg. Linnaea 32: 71. 1863. 

Glabrous, to about 2 meters high, the firm somewhat spreading 
branchlets slender; stipules rigid, linear-lanceolate, deciduous; 
petioles stout, 4-6 mm. long; leaves oblong-ovate or ovate-lanceolate, 
rounded-obtuse or lightly cordate at base, often long-acuminate to 
obtuse tip, 6-11 cm. long, usually 3.5-4.5 cm. wide; flowers dioecious, 
minutely racemulose or fasciculate, calyx 5-parted, lobes of male 
oblong-ovate; disk present in both sexes; male pedicels capillary, 
flexuose, little longer than petioles, the female firmer, shorter; 
style column slender, the 3 linear divisions spreading; stamens 3, 
filaments monadelphous, anthers finally transversely dehiscent. 
Was found by Martius, together with P. Martii M. Arg. in Mart. 
Fl. Bras. 11, pt. 2: 27. pi. 5. 1873, as near Peru as Japura, Brazil, 
and therefore is to be expected; P. Martii is similar but leaves only 
4.5-7 cm. long, 2-3 cm. wide, sepals all equal, flowers all distinctly 
racemulose. Here might key P. guyanensis (Aublet) M. Arg., 376, 
the leaves often acute at base, flowers of both sexes many, on axillary 
pulvinae, styles biparted; the species is to be expected. F.M. Neg. 
5038. 

Peru (probably). Brazil. 

Phyllanthus pseudo-conami M. Arg. in Mart. Fl. Bras. 11, 
pt. 2: 43. 1873. P. pseudo-conami M. Arg. var. pubescens M. Arg., I.e. 

Tall much-branched shrub, the slender branchlets, petioles (about 
1 mm. long), and short female pedicels puberulent in the Peruvian 
type; leaves oblong-ovate or subelliptic, rounded at base, obtusely 
short-acuminate, about 2 cm. long, 8-10 mm. wide, glabrous or 
essentially, olive-green; flowers several in the leaf -axils, the flexuose 
capillary male pedicels 5-7 mm. long; glands of the male flowers 
annulate, anthers free; ovary smooth, glabrous; styles bifid, diffuse; 
capsules 5 mm. broad, 3.5 mm. long, thus markedly depressed, deeply 
sulcate. The variety glaber M. Arg. from the upper Amazon is 
completely glabrous with gradually acuminate leaves 2.5-4 cm. long; 



46 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

capsules unknown and will probably prove to be distinct. Author 
wrote name with hyphen. F.M. Neg. 6595. 

San Martin: Tarapoto, Poeppig, type. Brazil? 

Phyllanthus racemigerus M. Arg. Linnaea 32: 23. 1863; 375. 

A tall glabrous shrub with angulate branchlets about 2 mm. 
thick, at base clothed with 25-30 distichous ascending leaves, and 
racemulose dioecious flowers borne at their early leafless tips; stipules 
ovate-lanceolate; petioles 2-2.5 mm. long; leaves ovate or oblong- 
ovate, both ends acuminate, 2.5-4 cm. long, 1-2 cm. wide, subcori- 
aceous, eglandular, glaucescent beneath; racemes of female flowers 
finally often 2 cm. long, rather densely bracted, suberect; male 
flowers unknown but stamens probably 3; calyx 2 mm. long, coria- 
ceous, the segments broadly ovate; urceolus entire; ovary subprui- 
nose, glabrous. The similar P. adenophyllus M. Arg., 376, of north- 
ern Brazil, has terete branchlets, leaves glandular spotted beneath 
below the apex, 1.5-2.5 cm. long, 8-15 mm. wide, and likewise 
probably will be found in Amazonian Peru; P. racemigerus has been 
collected as near as Solimoes. Illustrated, Mart. Fl. Bras. 11, pt. 2. 
pi. 7.fig.l. 

Peru (probably). Amazonian Brazil. 

Phyllanthus rubellus M. Arg. in DC. Prodr. 15, pt. 2: 379. 
1866. 

Glabrous, the subflexuous terete branches palely lenticellate, the 
branchlets purplish-brown; stipules small, nearly scarious; petioles 
2-3 mm. long; leaves oblong- or lanceolate-elliptic or oblong-ovate, 
obtuse both ends, 3-5 cm. long, 1.5-2.5 cm. wide, green above, 
reddish-glaucous beneath, reticulate- veined; female flowers in axils, 
geminate or ternate, their pedicels about 2 mm. long, the ovate 
lacerate unequal calyx segments deciduous; urceolate disk crenulate; 
styles rigid, semiconnate, bifid, the lobes refracted; capsules globose, 
smooth, glabrous, mature not known. P. symphoricarpoides HBK., 
324, of Colombia and Ecuador, would key here except for the 3- 
parted styles which ally it to P. Poeppigianus; it is distinctive by 
the combination of leaves obtuse at both ends, 2 cm. long, about half 
as wide, and racemulose flowers. F.M. Neg. 8497. 

Peru(?): Without locality, Ruiz & Pavdn, type. 

Phyllanthus salviaefolius HBK. Nov. Gen. & Sp. 2: 116. 
pis. 107, 108. 1817; 330. 



FLORA OF PERU 47 

Allied to P. grandifolius but the shortly connate styles slender, 
the stigmas only 1 mm. dilated at tip, glands of the male flowers 
finally free and seeds costulate; branchlets and often leaves puberu- 
lent tomentose; stipules foliaceous, subequaling the petioles; fascicles 
sessile or shortly peduncled, many-flowered, the male pedicels about 
7 mm. long, capillaceous, the elongating female suberect, rigid, 
filiform, both sexes glabrous as the similar calyces, the former 2, 
the latter 5 mm. long; male nectaries truncate, foveolate-rugose; 
capsules 5 mm. long, 6 mm. broad; seeds finely costulate. Outer 
1-3 stamens sometimes distinct and branches often nearly all male 
or nearly all female. The Peruvian plant is var. glabrescens M. Arg., 
leaves glabrous above, scabrous on nerves beneath. Illustrated, 
HBK. I.e. and Baillon, Etude Gen. Euphorb. pi. 24, figs. 15-19. 

Huanuco: Pink-flowered shrub in rocks by river, Mito, 3278 
(det. Mansfeld). Without locality, Ruiz & Pavdn. To Colombia 
and Venezuela. 

Phyllanthus Vincentae Macbr., nom. nov. Xylosma minuti- 
florum Macbr. Candollea 5: 392. 1934 not P. minutiflorus F. Muell., 
1865-66. 

Shrub with glabrous or glabrate slender more or less flexuose 
branchlets; petioles 1.5 mm. long; leaves ovate-lanceolate or ovate, 
unequally rounded at base, gradually narrowed to the obtusish tip, 
finally to 1 dm. long, 3.5 cm. wide, usually smaller, glabrous, char- 
taceous-membranous, entire, drying darker above; fascicles sessile; 
pedicels glabrous, filiform, to 7 mm. long, somewhat contorted at 
least in age; calyx segments 6, scarcely 0.75 mm. long; stamens 6. 
In this work, Field Mus. Bot. 13. pt. 4, 1: 31. 1941, 1 noted that this 
plant is evidently from fruiting material of the same collection in 
herb. Delessert, Geneva, a species of Phyllanthus, probably P. 
guianensis Aublet or aifine. It appears to be near P. nobilis however 
and if distinct it may well bear the name of the well-known botanical 
bibliographer, Miss Edith Vincent, who with her usual care discerned 
that my original name was preoccupied in Phyllanthus. 

Loreto: Puerto Arturo, Yurimaguas, Williams 4970, type. 

4. SECURINEGA Comm. or Juss.? 

Similar to Phyllanthus but flowers always clustered or solitary 
in the axils, sepals always 5, coriaceous or firm, rudimentary ovary 
more or less obvious, styles usually 3, the divisions recurving. 



48 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Securinega congesta M. Arg. in Mart. Fl. Bras. 11, pt. 2: 76. 
1873; 379. Phyllanthus congestus M. Arg. Linnaea 32: 25. 1863. 

Glabrous tree or tall shrub, the branches terete, the branchlets 
more or less angled; stipules linear-lanceolate, about 1 cm. long; 
petioles 2-3 mm. long; leaves ovate-lanceolate, base acute, shortly 
cuspidate-acuminate, 6-12 cm. long, 1.5-4 cm. wide, firm-mem- 
branous, paler beneath; flowers in axillary glomerules, 1-7-flowered, 
the basal male with 5 short stamens, subcylindric ovary rudiment, 
the female calyx dark brown, subcoriaceous, about 4 mm. long, disk 
urceolate, lobulate; capsules dark-colored, 7 mm. broad, 5 mm. long 
or larger, usually only 1-2 developing, sometimes 3. S. elliptica 
(HBK.) M. Arg. of Ecuador may be known at once by its elliptic 
apically rounded leaves. Illustrated, Mart. Fl. Bras. pi. 11, fig. 1. 
F.M. Neg. 21577. 

Loreto: Caballo-Cocha, Williams 223 %; 2470. La Victoria, 
Williams 2943 (det. Mansfeld). Iquitos, Williams 8044; 8121 (det. 
Mansfeld). Rio Maranon near mouth of Rio Tigre, Killip & 
Smith 27524 (det. Standley). Near Mazan, Mexia 6542 (det. 
Standley). Rio Itaya, Williams 82. Florida, King 2274 (det. 
Standley). To the Guianas. "Puinayma Jipi-ey" (Klug, Huitoto). 

5. RICHERIAVahl 

Reference: Pax & Hoffm., Pflanzenreich IV. 147. xv: 26-30. 
1922. 

Trees with alternate petioled pinnately nerved leaves and dioecious 
apetalous flowers, the male glomerulate in axillary or lateral inter- 
rupted spikes or racemes, the dense female racemes shorter. Sepals 
free or connate, 3-5, imbricate. Male flowers with disk glands free 
between the 3-6 distinct stamens, the anthers oblong; ovary rudi- 
ment small. Female flowers with annulate disk; ovary 3-celled; 
styles short, stout, broad, entire or slightly lobed. Ovules in cells 
geminate. Capsules separating into bivalved cocci. Dedicated to 
Richer de Belleval, 1564-1632, French horticulturist. 

Leaves coriaceous, the male flowers sessile R. grandis. 

Leaves membranous, the male flowers pedunculate. 

R. submembranacea. 

Richeria grandis Vahl, Eclog. Amer. 1: 30. pi. 4- 1796; 27. 
Amanoa divaricata Poeppig in Poepp. & Endl. Nov. Gen. & Sp. 3: 



FLORA OF PERU 49 

22. pi. 226. 1845. Guarania ramiflora Wedd. ex Baillon, Etude 
Gen. Euphorb. 598. 1858. 

Glabrous or essentially except for sepals, disk and young ovary; 
branchlets angled, pruinose, the leaves apically crowded; petioles 
to 3 cm. long, canaliculate; leaves oblongish or variously obovate, 
narrowed to acute or subobtuse base, usually rounded-obtuse, rarely 
somewhat acuminate, ordinarily a dm. or two long and about half 
as wide, entire, coriaceous, in Peru often 2-4-glandular above near 
base (typically broadly obovate and eglandular or obscurely so), 
paler and irregularly reticulate beneath, with 6-8 nerves; inflores- 
cences solitary or several, the male 3-10 cm. long, the 3-7-flowered 
glomerules sessile, the female 3-5 cm. long, racemose, these flowers 
shortly pedicelled; bracts about 1 mm. long, often denticulate; 
male sepals 3-4, unequal, half as long as the filaments, the anthers 
introrse; female sepals 5, ovate, acute, 1.5-2 mm. long; capsules 
subovoid, about 1 cm. long, smooth. Named variants of doubtful 
value include vars. divaricata (Poeppig) M. Arg., leaves obovate- 
lanceolate, acuminate or acute, eglandular, and latifolia Pax & 
Hoffm., leaves broadly obovate, rounded, 2-4-glandular at base. 
F.M. Negs. 5015 (var. divaricata); 32540 (var. latifolia'). 

Huanuco: Cuchero, Poeppig (type, var. latifolia); Poeppig 1441 
(type, var. divaricata). To southern Brazil and the West Indies. 

Richeria submembranacea Steyermark, Field Mus. Bot. 17: 
419. 1938. 

Glabrous unless for the minutely strigillose petioles, these 5-12 
mm. long, and the hirtellous inflorescences; leaves obovate-elliptic, 
acute at base, abruptly caudate, 7.5-11.5 cm. long, 3.5-5.5 cm. 
wide, entire, concolor, membranous or submembranous, with 6-8 
pairs of nerves; male racemes slender, 9-11 cm. long, densely fuscous 
hirtellous, the shortly peduncled flowers 15 mm. broad, with 5 
broadly ovate obtuse sepals slightly hirtellous without; stamens 5, 
the anthers introrse; female flowers unknown. Differs from all 
other species in its submembranous leaves, from R. grandis further 
by its pedunculate male flowers and from R. laurifolia Baillon of 
Upper Amazon by its entire abruptly caudate leaves (Steyermark). 
Since the type was from Sao Paulo de Olivenca, a region which 
has many plants known to occur within Peru, this species also is to 
be expected. 

Peru (compare note above). Amazonian Brazil. 



50 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

6. ANDRAGHNEL. 

Reference: Pax & Hoffm., Pflanzenreich IV. 147. xv: 178. 
1922. 

Low, annual or scarcely suffrutescent (Peru), with small bistipu- 
late leaves, the monoecious petaliferous flowers 1-3 in their axils. 
Male flowers with 5 imbricate sepals, same number of shorter petals, 
free disk glands and stamens (Peru) the erect anthers with parallel 
cells; ovary rudiment small. Female flowers with 5 sepals but small 
or minute petals, disk glands sometimes connate, ovary 3-celled, the 
bifid styles with capitulate stigmas; ovules geminate. Capsules 
separating into cocci with triquetrous ecarunculate seeds. The 
name was an ancient one for Portulaca oleracea. 

Leaves glabrous or pulverulent, 5-7 mm. long A. microphylla. 

Leaves glandular-ciliate, about 1 cm. long A. ciliato-glandulosa. 

Andrachne ciliato-glandulosa (Millsp.) Croizat, Journ. Wash. 
Acad. Sci. 33: 11. 1943. Phyllanthus ciliato-glandulosus Millsp. Proc. 
Calif. Acad. ser. 2. 2: 219. 1889. 

Annual, spreading from filiform root, densely glandular ciliate 
except the oldest leaves, these sparsely so; stems terete, somewhat 
irregularly branched; petioles one-third as long as leaves, the latter 
mostly ovate, obtuse or emarginate, 1.5 cm. long, 1 cm. wide or 
many much smaller, even minute; stipules orbicular, entire; inflores- 
cence axillary, more or less geminate, the flowers pedicellate; male 
calyx with ovate acute lobes and five stiped glands opposite the 
clefts; stamens 5, filaments free; female flowers larger, the lanceolate 
obtuse hyaline-margined calyx lobes with 5 alternating cylindric 
glands at base; styles cleft to middle or free, the stigma slightly 
recurved; ovary globose, ciliate glandular as the depressed capsule; 
seeds pale brown, finely pitted, each pit with a scale. After Mill- 
spaugh. Stems often 1-1.5 dm. long. So closely resembles A. 
microphylla as to be very easily confused with it; both belong to 
section Phyllanthidea with a minute pistillode in the female flower 
and both allied to A. aspera Spreng. from the Punjab to Morocco; 
all undoubtedly pre-Tertiary species (Croizat). The determination 
has been accepted without critical examination. 

Lima: Ambar to Huacho, Stork 11471 (det. Johnston). Lower 
California. 

Andrachne microphylla (Lam.) Baillon, Etude Gen. Euphorb. 
577. 1858; 178. Croton microphyllus Lam. Encycl. 2: 212. 1786. 



FLORA OF PERU 51 

Somewhat glandular, finely and intricately branched; petioles 
about as long as the leaves, these elliptic or ovate, obtuse both ends, 
5-7 mm. long, 2-5 mm. wide, glabrous, or in type sparsely pulveru- 
lent, opaque, lucid-green; flowers solitary, geminate or ternate on 
capillary pedicels about 1 mm. long or sometimes on a very short 
peduncle, the female pedicels firmer, reflexed in fruit and glandular- 
hirtellous as the branchlets and petioles; male sepals hyaline, oblong- 
ovate (the female ovate-lanceolate, opaque, hyaline-margined), about 
as long as the rhombic-obovate petals, these much shorter than 
sepals in female flower, liguliform; stamens free as the stipitate 
urceolate disk glands; disk entire, membranous; styles nearly free; 
capsules glandular-hirtellous, 3.5 mm. broad. A much-branched 
annual (?) with incurved branches forming a subglobular mass, 1-6 
dm. high; detached from the weak root it becomes a "tumbleweed" 
(Haught). 

Huanuco: Near Huanuco, Dombey, type. Piura: Talara, east of 
La Brea, Haught 39 (det. Croizat). Lima: Viscas, along Rio Chillon, 
1,800 meters, Pennell 11^92 (det. Croizat). 

7. HYERONIMA Fr. Allem. 

Reference: Pax & Hoffm., Pflanzenreich IV. 147. xv: 31-40. 
1922. 

Shrubs or trees marked (in Peru) by the lepidote indument espe- 
cially on the leaves beneath, these pinnately nerved, the nerves 
arcuately ascending and joined before the entire margins. Flowers 
dioecious, apetalous, often paniculate, shortly pedicelled, or sub- 
sessile, the bracts minute, the male inflorescences the larger. Male 
calyx somewhat campanulate with 5 more or less connate disk glands, 
3-6 free stamens, the pendulous anthers exserted, the connective 
enlarged. Ovary rudiment small. Female disk entire or lobulate; 
ovary 2(-nearly 3)-celled, the short styles bifid, reflexed; ovules 
geminate. Fruit drupaceous with 1 ecarunculate seed. Genus 
named for Jeronimo Serpa, Brazilian horticulturist, and written by 
the author Hyeronima or Hieronima; Walpers' change to Hieronyma 
has been generally followed but the author's first choice should be 
retained as by Lanjouw. The type species, H. alchorneoides Fr. 
Allem., the "uricana" or "urucurana" of Brazil, is the source of a 
fine wood said to be much sought for furniture. 

The following key is after Pax and Hoffmann but from the little 
material seen it seems possible that the degree of pubescence as a 



IIRRARY 



52 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

character for the distinguishing of species may prove to be unreliable; 
the number of species may therefore actually be fewer than these 
authors indicated. 

Leaves densely lepidote beneath, the scales more or less overlapping. 
Leaves oblong, ovate or obovate, rarely elliptic, about half as 

wide as long, scarcely if at all scabrous. 

Leaf nerves pilose beneath; fruit 3-5 mm. long. H. Moritziana. 
Leaf nerves not pilose beneath; fruit (known) about 1 cm. long. 

Leaves rigid, obtuse H. macrocarpa. 

Leaves membranous, acute ' H. andina. 

Leaves broadly elliptic, more or less scabrous, often nearly as 

wide as long. 
Leaves flexible; rachis branches even below slender or about 

2 mm. thick H. akhorneoides. 

Leaves rigid; rachis branches to 4 mm. thick. . . . H. peruviana. 
Leaves closely to sparsely lepidote beneath but the scales not over- 
lapping. 

Scales evenly distributed on leaves beneath, approximate; nerves 
not at all pilose. 

Leaves rigid, obtuse H. macrocarpa. 

Leaves membranous, acute H. andina. 

Scales scattered on leaves beneath; nerves more or less pilose. 

Leaves oblong or oblong-obovate H. oblonga. 

Leaves rotund-elliptic H. laxiflora. 

Hyeronima alchorneoides Fr. Allem. Diss. c. icon, in Trab. 
Veil. Rio Jan. 1848; 36. 

Stout branchlets, petioles (2-6 cm. long), leaves beneath and 
calyces densely lepidote; stipules deciduous, only the upper present, 
but conspicuous, petiolate, nearly a cm. long; leaves broadly elliptic, 
sometimes suborbicular, often 1-2.5 dm. long, 0.5-1.5 dm. wide or 
even larger, abruptly acute or very shortly apiculate, rounded at 
base or slightly contracted to petiole, membranous or subcoriaceous, 
scattered punctulate-lepidote and slightly scabrous above, glabrate 
or somewhat pilose on the nerves, the 7-10 secondary as the veins 
prominent beneath; panicles erect-spreading, the male to 1.5 dm. 
long, the female much shorter; bracts small, broadly ovate, acute 
or obtuse; pedicels to 2 mm. long; calyx of both sexes poculiform, 



FLORA OF PERU 53 

3-5-dentate, pubescent within as without, male disk entire, shorter 
than calyx, pilosulous, the thicker female pilose-margined; rudi- 
mentary ovary obovate- truncate, pilose; ovary lepidote; fruit ovoid- 
globose, 4-5 mm. long, sparsely lepidote. Becoming a fine timber 
tree, the heartwood red, the sapwood reddish brown when first cut. 

Huanuco: Shapajilla, 630 meters, Woytkowski 30 (det. Standley). 
Loreto: Mouth of the Santiago, Tessmann 4114 (det. Mansfeld). 
Gamitanacocha, Rio Mazan, Jos& Schunke 330 (det. Standley). 
Rio Acre: Mouth of the Rio Macauhan, Krukoff 5640 (det. Steyer- 
mark). To Para and southern Brazil. "Uricana," "urucurana," 
"palo de sangre." 

Hyeronima andina Pax & Hoffm. Pflanzenreich IV. 147. xv: 
37. 1922. 

In general resembles H. macrocarpa especially in pubescence, but 
petioles 2-4 cm. long, leaves elliptic, acute or subacute at base, 
cuspidate, 13-20 cm. long, 6-9.5 cm. wide, membranous; male 
panicles 8-14 cm. long, laxly few-branched, bracts 1 mm. long, 
acute, pedicels 2-3 mm. long, male calyx 3-5-dentate, disk glands 
little shorter than calyx, pubescent as rudimentary ovary; female 
panicles 4-5 cm. long, pedicels 1 mm. long, calyx 5-dentate, longer 
than entire pubescent disk; ovary sparsely lepidote or subglabrous; 
fruit probably small (authors). F.M. Neg. 5010. 

Peru (probably). Colombia to Bolivia. 

Hyeronima laxiflora (Tul.) M. Arg. Linnaea 34: 66. 1865; 39. 
Stilaginella laxiflora Tul. Ann. Sci. Nat. ser. 3. 15: 244. 1851. 

Branchlets not at all stout, densely lepidote as the petioles, these 
rather slender, 4-7 (12) cm. long, and spreading panicles, the male 
12-18 cm. long, the female much shorter; stipules oblong-ovate, the 
petioles nearly as long as the cochleate 3-10 mm. long blade; leaves 
rotund-elliptic, cuspidate-acuminate at the rounded apex, obtuse 
or shortly acute at base, 1-2 dm. long, usually 7-15 cm. wide, mem- 
branous or subcoriaceous, laxly reticulate-veined, early scattered and 
sparsely lepidote both sides, more or less brownish pilose beneath 
on the nerves, these ordinarily about 7, rarely 10; lower bracts 
simulating stipules, acute, floriferous, small; pedicels rigid, 1.5 mm. 
long or shorter; male calyx minutely 3-5-dentate, subequaled by the 
entire rufous-pilose disk, rudimentary ovary elongate, pilose; female 
calyx minutely denticulate, ovary sparsely lepidote; fruit little com- 
pressed, 4.5 mm. long. Crooked trunks with distinct root insertions 



54 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

or very heavy spurs to a high level (Lanjouw). To 25 meters tall, 
the wood very hard. F.M. Negs. 19494, 24358. 

Peru: Without locality, (Martins 2647, in part, fide Pax & 
Hoffmann). To the Guianas; Colombia? 

Hyeronima macrocarpa M. Arg. amend. Pax & Hoffm. Pflan- 
zenreich IV. 147. xv: 34. 1922. H. macrocarpa M. Arg. var. Spruceana 
M. Arg. Linnaea 34: 66. 1865. 

Pubescence entirely lepidote, the younger parts densely, the leaves 
permanently so beneath; petioles 1.5-2 cm. long, acutely canaliculate 
above; leaves obovate, rounded-obtuse or emarginate, 5-7 cm. long, 
3.5-4.5 cm. wide, rigid, prominently reticulate-veined, glabrous 
above; female racemes panicled, bracts at least 1 mm. long, broadly 
ovate; pedicels stout, angled, in fruit to 4 mm. long; female calyx 
obconic, minutely if at all 5-denticulate, disk entire; immature 
fruits at least 1 cm. long, 5 mm. broad above the middle, narrowly 
compressed pyriform, acuminate, glabrous. Perhaps should include 
H. andina and then probably within Peru. F.M. Neg. 7037 (var. 
Spruceana). 

Peru (compare note above). Ecuador. 

Hyeronima Moritziana (M. Arg.) Pax & Hoffm. Pflanzenreich 
IV. 147. xv : 33. 1922. H. macrocarpa M. Arg. var. Moritziana M. 
Arg. Linnaea 34: 66. 1865. 

Angled branchlets, petioles, these 1.5-2 cm. long and leaves 
beneath densely lepidote, the former becoming glabrescent; leaves 
oblong-elliptic or obovate, obtuse, emarginate or acute, obtuse or 
acute at base, 4-11 cm. long, 2-4.5 cm. wide, chartaceous, the 6 or 7 
nerves pilose beneath, the venation there reticulate; stipules lanceo- 
late, cochleate, 5-7 mm. long; panicles to 8 cm. long in fruit, lepidote 
as the 0.5-1 mm. long bracts; male calyces shortly dentate, lepidote 
disk divided into 5 cuneiform lobes, the subglabrous rudimentary 
ovary included; female calyces 4-5-dentate, not at all exceeded by 
disk; fruits glabrous or subglabrous, ovoid or pyriform, 3-5 mm. 
long. The typical form, var. Fendleri (Briq.) Pax & Hoffm., has 
slender petioles, leaves 4-7 cm. long; var. yungasensis Pax & Hoffm., 
petioles stouter, leaves to 10 cm. long, nerves pilose but finally 
glabrescent; as fruit is unknown this may not pertain to H. Morit- 
ziana (Pax & Hoffm.). F.M. Neg. 5014 (var. Fendleri). 

Peru (probably, especially if the Bolivian variety belongs here). 
Venezuela; Bolivia? 



FLORA OF PERU 55 

Hyeronima oblonga (Tul.) M. Arg. Linnaea 34: 66. 1865; 37. 
Stilaginella oblonga Tul. Ann. Sci. Nat. se>. 3. 15: 248. 1851. 

Branchlets terete, densely lepidote as the petioles, these slender, 
1-3 cm. long; stipules cochleate, about 6 mm. long; leaves early 
sparsely and scattered lepidote, more or less pilose beneath on the 
5-7 prominent nerves (veins reticulate) or finally glabrescent, mem- 
branous, lustrous or rarely opaque, varying in shape but often some- 
what obovate and lanceolate to oblong, ordinarily 7-15 cm. long, 
about half as wide; panicles 3-10 cm. long, often branched to base, 
the