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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 male longer, the female shorter and more simple, all lepidote
including the calyces; pedicels of former 1 mm. long; bracts ovate,
acute; calyces 5-dentate, thick disk glands of the male as cylindric
rudimentary ovary and cupulate entire female disk tomentulose;
ovary glabrous; fruit about 5 mm. long, ellipsoid, acute both ends.
F.M. Neg. 24359.
San Martin: Zepelacio near Moyobamba, Klug 3588 (det.
Standley). Loreto: Mouth of Rio Santiago, Tessmann J^.165 (det.
Mansfeld). Balsapuerto, Klug 3113 (det. Standley). Mexico to
Bolivia and southern Brazil.
Hyeronima peruviana Pax & Hoffm. Pflanzenreich IV. 147.
xv : 37. 1922.
Densely lepidote except the leaves above and the older branch-
lets; stipules ovate, acute, contracted at base, 1-2 cm. long; petioles
2-5 cm. long; leaves broadly elliptic or obovate-elliptic, acute at
base, very shortly and acutely acuminate, 10-14 cm. long, 6-9 cm.
wide, rigid coriaceous, punctate with scattered lepidote indument
but scarcely scaberulous above, the 7-9 nerves with the veins
prominent beneath; panicles robust, the male 10-15 cm. long, the
female 4-5, the small broadly ovate bracts acute, the flowers sub-
sessile; male calyx poculiform, 5-dentate, pubescent, the firm dilated
disk pilosulous and marginally undulate-crenate; ovary rudiment
subcylindric, pilose; female calyx cupulate, lightly denticulate, the
hypogynous disk ciliate, the ovary densely lepidote. Noted to
attain 15 meters. Allied to H. akhorneoides of eastern Brazil but
differs sufficiently in the rigid coriaceous leaves, robust inflorescence,
the rachis 3-4 mm. thick, the subsessile male flowers and character
of disk (authors). The Ecuadorean related species H. asperifolia
Pax & Hoffm. has leaf nerves pilose beneath while H. macrocarpa
(M. Arg.) Pax & Hoffm. is said to differ in its obovate leaves, rounded-
obtuse or emarginate and only 5-7 cm. long, 3.5-4.5 cm. wide, disk
56 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
entire, glabrous; there are other closely related species to be expected,
maybe distinguishable by their membranous asperous leaves.
Huanuco: Huamalies above Monzon, open woods, 900 meters,
Weberbauer SU7; 285. Chinchao, Weberbauer 6822Jvnin: West
of Huacapistana, in shrubs, 2,700 meters, Weberbauer 2185, type.
8. DRYPETESVahl
Reference: Monachino, Phytologia 3: 32-35. 1948.
Trees or shrubs with alternate entire or sometimes serrulate
coriaceous pinnate-nerved leaves, minute or rarely foliaceous cadu-
cous stipules and apetalous dioecious flowers fasciculate in the leaf
axils or on the older wood. Calyx in both sexes with 4-5 (6-7) often
coriaceous or fleshy sepals, the male with central disk, and sometimes
with glandular ring, 3-12 free stamens opposite sepals, rudimentary
ovary variously developed, large, scale-like, none or obscure, the
female with annulate or cupulate disk, l-2(-4)-celled ovary, sessile
or subsessile stigmas, geminate ovules. Fruit drupaceous, scarcely
or slightly fleshy, indehiscent, with 1 or 2 ecarunculate seeds.
Similar or confused genera include Discocarpus Klotzsch, Pflanzen-
reich IV. 147. xv: 203, and Chonocentrum Pierre, I.e. 205, the former
with petaliferous male flowers with annulate lobed disk, styles
developed, fruit subglobose, capsular, the latter with apetalous
flowers, male with urceolate disk, large rudimentary ovary, female
unknown; this is C. cyathophorum (M. Arg.) Pierre, originally in-
cluded in Drypetes; as to Discocarpus, D. Spruceanus M. Arg. and
D. brasiliensis Klotzsch perhaps are most likely to be found within
Peru, the former with subequal calyx and petals, pedicellate smooth
capsules, the latter with subsessile female flowers (male unknown),
muricate capsules.
Drypetes amazonica Steyerm. Field Mus. Bot. 17: 420. 1938;
35.
Glabrous except the inflorescence; stipules not observed; petioles
4-10 mm. long; leaves elliptic, entire, typically 12-24 cm. long, 3-7
cm. wide, gradually acuminate; male flowers 7-many on slender
pedicels 3-4 mm. long; sepals 4, subrotund, sparsely strigillose with-
out and marginally ciliolate, 2.6-2.9 mm. long, 2.1-2.3 mm. wide;
stamens 8-10, the obvious central disk obscurely puberulent (fide
Monachino); rudimentary ovary none; female flowers 1-3 with 5
rounded sepals 3-3.5 mm. long, 2.2 mm. wide, pilosulous without;
FLORA OF PERU 57
fruits pyriform, tomentose, to 1.7 cm. long (immature). D. variabilis
Uitt. of the Guianas has 1-3 flowers, 4-8 stamens with glabrous
anthers; the Peruvian material according to Monachino belongs
with these two species but their precise position "is dubious";
their specific validity as indicated by Standley in herb, seems to be
too questionable to give him as authority so for the convenience
of this work they may be disposed of as D. amazonica var. peruviana
Macbr., var. nov., foliis oblongo-ellipticis vel late ellipticis, 10-15
cm. longis, 4-7.5 cm. latis, subabrupte breviterque acuminatis;
floribus masculis numerosis, pedicellis 5-10 mm. longis; sepalis
suboblongis, circa 2.2 mm. longis, 1.5 mm. latis; staminibus paullo
inaequalibus. Probably Krukoff 5624 belongs to the variety but
Monachino observes that it has 8-11 stamens. The Peruvian plants
do not seem to have the anthers at quite the same level and in this
respect suggest the pubescent branched D. Spruceana M. Arg. of
Brazil in which species they are obviously at two levels, and, more-
over, glandular-ringed at base, the rudimentary ovary large. In
the other Brazilian species with gland-ringed stamens, D. Krukovii
Monach. I.e. 34, the anthers are at the same level and the rudi-
mentary ovary consists of three scales. A tree 5-11 meters high.
San Martin: Juanjui, Klug 3869 (type, var. peruviana); also
3880. Rio Acre: Krukoff 5624. Brazil.
9. CROTONL.
Julocroton Mart. Flora 20, Beibl. 2: 119. 1837.
Reference: Mueller, Argoviensi, in DC. Prodr. 15, pt. 2: 513-
700. 1866.
Herbaceous or ligneous, usually lepidote or stellate pubescent,
the leaves mostly alternate and often biglandular at base or tip of
petiole, seldom lobate or pinnate-nerved, the spicate or racemose
flowers commonly monoecious, the male usually disposed above the
female, these sometimes petaliferous, solitary under each small
bract or sometimes with 2-3 male which are ordinarily petaliferous
with 5 perianth segments, 5-many stamens, erect at anthesis.
Female perianth 3-5-lobed, sometimes irregularly. Ovary 3(2-4)-
celled, the styles 1-many times parted; ovules solitary; capsule
separating into 3 bivalved cocci; seeds smooth carunculate.
Julocroton, defined only by the unequal female sepals, sometimes
glandular or appendaged, is an arbitrary segregation, as the character
occurs elsewhere and in Peru even in Croton proper, since C. Boissieri
58 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
and C. tarapotensis have unequal calyx segments; and Croizat,
Rev. Arg. Agron. 10: 144. 1943, cites C. leucophyllus Torr. of Texas
as a species with irregularly lobed perianth segments but accepts
Julocroton as it differs materially according to him from Croton in
the sum of characters that distinguish it. There are many examples
in taxonomy to support his argument but it fails to carry weight
unless the "sum of characters" is consistently developed; here it
does not seem to be.
Since there is no modern revision of this large genus, many of
the species based on Peruvian plants, whether restricted to type or
interpreted according to tradition, may prove to be the same or
variants of others from adjacent areas; it seems possible too that
many of the local forms given names recently and by former students
are the result of introgressive hybridization and apomixis; under
these circumstances the following key at best is only suggestive.
C. Lorentzii M. Arg. Goett. Abh. 19: 96. 1874 was reported by
Bruns from Cachendo, Arequipa (Guenther & Buchtien 249). The
species has been omitted since the determination is highly uncertain.
Annuals, weedy, herbaceous or the stems sometimes hard but then
very slender.
Leaves deeply lobed C. lobatus.
Leaves dentate.
Leaves obtuse both ends or merely acutish; stem indument often
spreading at least in part C. glandulosus.
Leaves ovate-acute or -acuminate; stem indument appressed.
C. trinitatis.
Ligneous plants and clearly shrubs or trees.
Leaves (mature) small, 1-2 (2.5) cm. wide (a small state of C.
Ruizianus might be sought here).
Leaves rhombic-ovate, acuminate, about 2 cm. wide, lepidote.
C. adipatus.
Leaves rotund, barely apiculate (see also C. balsameus).
C. nitidulifolius.
Leaves ovate-lanceolate or narrower, lepidote or stellate.
Leaves acute both ends, closely lepidote; stamens about 20.
C. Boissieri.
Leaves rounded or subacute at base; stamens about 10.
Leaves narrowly ovate, 1.5-2.5 cm. long.
FLORA OF PERU 59
Female pedicels 1 mm. long; styles glabrous.
C. rubiginosus.
Female pedicels to 3 mm. long; styles tomentulose.
C. andinus.
Leaves lanceolate, to 4 cm. long; female pedicels elongate.
C. pedicellatus.
Leaves distinctly larger, at least many broader.
Indument predominantly stellate, sometimes minute, scurfy
or lepidote-stellulate, sometimes nearly lacking, rarely
simple (see p. 63).
Leaves ovate to rotund, rarely oblongish, and the petioles
often shorter than 3 cm.
Leaves not acuminate or in any case mostly or all narrowed
well above the middle, never acutely acuminate from
about the middle or caudate.
Leaves rhombic-ovate, softly stellate-tomentose above.
C. thurifer.
Leaves various, rarely rotund, often soon glabrate or, if
stellate, asperous rather than tomentose above.
Leaves at least mostly somewhat emarginate, glabrate
above.
Leaves orbicular-ovate, glandular at base.
C. balsameus.
Leaves broadly ovate, distinctly or somewhat longer
than wide, eglandular C. heliotropiifolius.
Leaves usually more or less acute, sometimes rounded,
rarely shortly acuminate.
Leaves acute or subrounded to openly cordulate at
base, submembranous, smooth and glabrate
above; stamens 15-25.
Leaves obtuse or apiculate; glands usually 2,
sessile or subsessile or lacking.
C. rivinaefolius, C. alnifolius.
Leaves shortly acute-acuminate ; glands 2-4, stiped.
C. spurcus.
Leaves usually minutely cordulate, often coriaceous,
somewhat scabrous and rugose above.
Stamens at least typically 30-40; leaves often
subrotund. . . .C. Ruizianus.
60 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Stamens at least typically 15-22; leaves often
ovate.
Leaves pinnate-nerved, entire; filaments pubes-
cent only below C. collinus.
Leaves 5-7-plinerved, crenate-denticulate; fila-
ments hirtellous C. pycnanthus.
Leaves acutely acuminate and narrowed from about the
middle, or, if above, caudate-acuminate as in C. Pilgeri.
Leaves 3-5-plinerved from base, eglandular, often about
10 cm. long.
Calyx lobes entire; leaves membranous, green above.
C. scaber.
Calyx lobes (female) lacerate; leaves firm, cinereous
above C. flavispicatus.
Leaves 3-7-plinerved, coriaceous, biglandular, often 4-8
cm. long C. Baillonianus.
Leaves pinnate-nerved to glandular base.
Leaves subabruptly acuminate, bicolored, with 2
sessile glands C. Pilgeri.
Leaves subconcolored with 2-4 stiped glands.
C. spurcus, C. churumayensis.
Leaves oblongish or ovate-rotund but the petioles soon mostly
or all even on flowering branchlets 3-many cm. long.
Leaves ovate-rotund, never oblongish, always cordate or
at least broadest at base except C. fragrantulus (see
p. 62).
Petioles 3(2)-4 cm. long or rarely half as long as the
usually medium-sized leaves, the often dense pubes-
cence ferrugineous, not or scarcely barbate.
Inflorescence paniculiform; lax or scandent shrub;
leaves ovate but acutish at base. . .C. fragrantulus.
Inflorescence spiciform or strobiliform; leaves rounded
to cordate at base.
Female calyx lobes equal, entire; stamens 15-many;
inflorescence spiciform; leaves glandular.
Basal glands 2, sessile or shortly stiped.
Leaves obtuse to acute or not clearly acuminate,
often submembranous, the subpinnate
FLORA OF PERU 61
nerves usually obscurely impressed; sta-
mens in type 30-40 C. Rui&ianus.
Leaves typically narrowed from middle, acumi-
nate, 3-5-nerved at base, nerves prominent
above; stamens ex char. 15-32 (apparently
there is a membranous-leaved state).
C. Baillonianus.
Basal leaf glands 4(2)-several, conspicuously
stiped C. pungens, C. quadrisetosus.
Female calyx lobes deeply dentate, unequal; stamens
about 10; inflorescence strobiliform; leaves
eglandular.
Petioles early 3-4 cm. long; disk entire.
C. triqueter.
Petioles mostly 2-3 cm. long; disk acutely lobed.
C. flavispicatus.
Petioles soon elongate, all unless the uppermost 5-many
cm. long, the leaves usually ample (or the uppermost
petioles shorter but subbarbate) unless in C. palano-
stigma.
Pubescence of branchlet tips and petioles subbarbate;
basal leaf-glands stiped or indument in part
reddish-fulvous; leaves greenish-brown (cf. C.
Tyndaridum}', species doubtfully distinct.
Glands stiped, 2 or more, conspicuous (cf. C.
pungens).
Leaves closely tomentose-stellate beneath, shal-
lowly or openly cordate.
C. callicarpaefolius, C. quadrisetosus.
Leaves rather openly hispidulous-stellate beneath,
deeply and narrowly cordate . C. perspeciosus.
Glands sessile or minute and nearly hidden in
tomentum.
Leaves distinctly longer than wide, acuminate.
Leaves asperous above with rather sparse
lepidote-stellulate indument.
C. Rehderianus.
Leaves coarsely corrugate and softly stellulate
above C. bryophorus.
62 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Leaves ovate-rotund, scarcely longer than wide,
short-acuminate C. Tyndaridum.
Pubescence of branchlet tips and petioles short, ap-
pressed or not at all shaggy-barbate; glands
depressed (cf. C. Tyndaridum); leaves usually
drying dark, the indument beneath cinereous if
present.
Flowers small, to about 3.5 mm. long.
Leaves ample, glabrate in age; seeds finely striate.
C. draconoides, C. Lechleri.
Leaves cinereous beneath or if glabrate rarely
10 cm. long; seeds estriate, somewhat punc-
ticulate-rugulose (or coarsely costate, known).
Leaves rounded or lightly cordate at base, sub-
caudate-acuminate.
Leaves densely stellate-villous beneath, 1-1.5
dm. long; capsules about 5 mm. long.
C. tarapotensis.
Leaves lightly or scurfy-stellate beneath, 6-10
cm. long; capsules 3-4 mm. long.
Indument of leaves beneath scurfy.
C. xanthochylus.
Indument of leaves beneath sparse, stellate.
C. peltophorus, C. erythrochilus.
Leaves deeply cordate C. Macbridei.
Flowers about 5 mm. long, the female sepals some-
what accrescent; leaves greenish and glabrate
both sides or cinereous beneath, ample.
C. huitotorum, C. palanostigma.
Leaves oblongish, acuminate, or rarely, as in C. perlongi-
florus, acute, rarely broadest at base.
Leaves soon glabrous both sides; glands slender-stiped.
C. olivaceus.
Leaves more or less pubescent, at least beneath; glands
obscure or sessile.
Indument of leaves above simple or mostly simple, lax.
C. peruvianus, C. stenosepalus.
Indument of leaves above asperous-stellulate or ol
scure.
FLORA OF PERU 63
Leaves scattered scurfy or appressed-stellate be-
neath C. sampatik.
Leaves villous-stellate, at least on nerves beneath.
Sepals in fruit to 6 mm. long; leaves smooth above.
C. Astianus.
Sepals not accrescent; leaves scabrous above.
Leaves subcuneate-rounded at base, tomentose
beneath C. sapiiflorus.
Leaves cordulate at base, stellate-villous.
C. perlongiflorus.
Indument predominantly lepidote; leaves green at least above;
petioles often less than 2 cm. long.
Leaves eglandular at rounded or subacute base, sparsely
lepidote above C. glabellus.
Leaves biglandular at cuneate or acute base.
Indument soon lacking above, usually silvery beneath;
leaf nerves about 20 pairs C. matourensis.
Indument about equally sparse both sides; nerves about
15 pairs (above basal) C. Tessmannii, C. cuneatus.
Croton adipatus HBK. Nov. Gen. & Sp. 2: 68. 1817; 552.
Branchlets dichotomously branched, acutely angled as the
petioles, flowers, as leaves above, lepidote with small multiradiate
trichomes, these medially lustrous, obtusely umbonate; petioles
stout (stipules obsolete), carinate beneath, about 1 cm. long; leaves
rhombic-ovate, subobtuse at minutely glandular or eglandular base,
acuminate, entire, rigid but fragile, 3-4 cm. long, 2-2.5 cm. wide,
3-plinerved, the nerves prominent beneath, the scales there a little
broader, paler and more lustrous; racemes stout, floriferous nearly
to base, the rachis multiangled; bracts ovate, 1-3-flowered; male
pedicels subequaling the lepidote calyx, this 2 mm. wide before
open ; stamens about 15 ; filaments glabrous. Relationship unknown,
the female flowers not seen. P.M. Neg. 5049.
Cajamarca: Tomependa, Prov. Jae"n, Bracamoros, Bonpland,
type. Near mouth of Rio Chinchipe, Weberbauer 6218 (det. Mans-
feld). "Ullucina."
Croton alnifolius Lam. Encycl. 2: 205. 1786; 604.
Younger branchlets depressed stellate, finally sublepidote, angled;
stipules minute; petioles 1-2 cm. long; leaves biglandular at base,
64 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
broadly obovate or elliptic, commonly rounded-obtuse, sometimes
oblong-elliptic, 3-6 cm. long, 2.5-4.5 cm. wide, subentire, firm-
membranous, the youngest fulvous tomentose both sides, early
scattered stellate puberulent above, soon glabrate; secondary nerves
parallel, the lower 4 approximate; racemes a dm. or two long, dense;
bracts ovate-lanceolate, denticulate; female flowers few or lacking,
often with about 10 rudimentary stamens, fruiting calyx not accres-
cent; male flowers at opening 3 mm. across, 5-parted, stamens 12-20,
filaments pilose at base; ovary stellate- tomentose; capsules 7.5 mm.
long, globose-ellipsoid, glabrescent; seeds 6 mm. long, smooth except
finely costulate. While the species is typically of lomas it appears
to be the same when in rocks and river plains of the interior. Here
might be sought C. rhamnifolius HBK., apparently in Bolivia, with
some hispidity on branches, the petioles often only a few mm. long,
the flowers pedicellate; cf. also C. heliotropiifolius HBK. The
Stork specimen could be C. rivinaefolius, not clearly distinguishable.
F.M. Neg. 5051.
Piura: Serran, river alluvial plain, 100 meters, Stork 11379.
Lima: Cheuchin, Ruiz & Pavdn, type; Dombey; Martinet. Lurin,
low clumps on sandy lomas, 5934 (det. Johnston). Atocongo, clefts
of limestone rocks, Pennell 14755. Santa Eulalia, 1,300 meters,
among large rocks, Goodspeed (det. Croizat). Arequipa: South of
Mollendo, in nitrate dust on hard sand behind beach area, Eyerdam
25178. Prov. Canete, hills of Asia, stony slopes, 500 meters, Vargas
9312 (det. Johnston). Apurimac: Quisapata, 2,420 meters, Vargas
8912.
Croton andinus M. Arg. Linnaea 34: 126. 1865; 639.
Much-branched little shrub similar to C. pedicellatus; leaves
greener above, more ovate and subacute both ends, 1.5-2.5 cm. long,
4-8 mm. wide; racemes about 1.5 cm. long; pedicels equaling the
calyx, the female nearly 3.5 mm. long, the male 1.5 mm. wide;
lanceolate male petals long-ciliate only at base; filaments glabrous;
style divisions tomentulose; capsules 4 mm. long. Otherwise
scarcely differs from the related species and is yet another case
requiring the study of more material to determine the significance,
if any, of the characters noted. Croizat, Darwiniana 5: 458. 1941,
suggested that it has a common range within Peru with C. pedicellatus.
F.M. Neg. 7071.
Peru (probably; cf. note above). Bolivia.
FLORA OF PERU 65
Croton Astianus Croizat, Journ. Arnold Arb. 21: 95. 1940.
Type from a 4 meter shrub, the tips sordidly yellow-tomentulose,
the bark becoming suberosely fissured; stipules subsetaceous, entire,
1-1.5 cm. long; petioles to 3.5 cm. long; leaves elliptic or elliptic-
lanceolate, cuneate-rounded at base, obvious glands none, 6-16 cm.
long, 3-5 cm. wide, coriaceous, rugulose, yellowish-villous-tomentose
beneath, marginally with a dense papillose-setose zone; veins im-
pressed above, about 8 pairs, only the first pair subtriplinerved;
racemes (cymes) mostly male, to 3 dm. long, globose male buds about
3 mm. broad, pedicels 5 mm. long; female flowers solitary, calyx
9 mm. long, 6 mm. wide, gray tomentose, the triangular-lanceolate
acute lobes 6 mm. long, 2 mm. wide, glabrous within; ovary hispid,
small, globular; styles bifid nearly to base, stout, black, canaliculate,
to 5 mm. long. The long-acuminate calyx lobes suggest some
affinities with C. speciosus from Venezuela and with C. caldensis
from Brazil but the vegetative characters are quite different and
the female flower is smaller; developed male unknown (Croizat).
Meritoriously named for Susanna Ast, student of the Annoniaceae
of Indo-China.
Junin: Between Panti and Rocchac, 2,400 meters, valley of the
Pariahuanca, Weberbauer 6536, type.
Croton Baillonianus M. Arg. Linnaea 34: 85. 1865; 536.
C. Baillonianus M. Arg. var. pachyphyllus M. Arg. I.e.
Branchlets rather stout, terete, apically reddish floccose-tomen-
tose, the leaves permanently so beneath, the younger typically
pulverulent-floccose above finally sometimes glabrescent and punctic-
ulate-scabrous, often 4-8 cm. long, 2.5-4.5 cm. wide (petioles as
long or a third shorter), ovate or oblong-ovate, acuminate, (3) 5-7-
plinerved, more or less obscurely biglandular at base, in type sub-
coriaceous; secondary nerves arcuate-ascending, prominent beneath;
stipules small, subulate; racemes dense, often entirely male; bracts
short-ovate, 1-3-flowered; male pedicels 4-6 mm. long, firm, spread-
ing, reddish villous as the subdepressed-globose 3 mm. wide male
flower, this with elliptic-ovate ciliate and basally lanate petals, 13-
32 pilosulous filaments, densely whitish villous receptacle; ovary
rufous tomentose, styles deeply biparted, the laciniae shortly bifid,
bilobed or subentire. Var. submembranaceus M. Arg. has the
broadly ovate leaves reddish tomentose both sides, then above
sparsely, underneath densely, stellate-puberulent, finally cinereous
beneath. As interpreted here this probably includes C. perlongiflorus
66 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
(said to have smaller male flowers) as to Cuzco plants, and in turn
C. Baillonianus is perhaps not specifically distinct from C. ferrugineus
HBK. or C. rivinaefolius HBK. but its leaves are rather uniformly
cordulate-ovate, coriaceous with impressed veins, short- or long-
acuminate. There is also a great resemblance to C. quadrisetosus
except for the lack of the long-stiped leaf -glands. F.M. Negs. 5056;
34019; 5057 (var.).
Huanuco: Near Pozuzo, 4806. Junin: Near Tarma, Ruiz &
Pav6n, type. Cuzco: Ollantaytambo, Cook & Gilbert 283. Santa
Ana, Cook & Gilbert 1483; 1560; 1561; 1562 (leaves long-acuminate).
San Miguel, Cook & Gilbert 916. Machu-Picchu, West 6464 (det.
Wheeler, Julocroton triqueter). Salapunco, Vargas 8112. "Cabra-
cabra" (Cook & Gilbert).
Croton balsameus M. Arg. Linnaea 34: 107. 1865; 589.
Much-branched shrub, the short intricate nodulose branchlets
densely leafy, the younger as all growing parts densely tomentose;
stipules dentiform, subulate; petioles 2-4 times shorter than the
leaves, these 1-6 cm. wide, often a little longer, orbicular-ovate,
subemarginate or obtuse both ends, subentire, submembranous,
minutely puberulent or glabrate above, 2-4-glandular at base, 3-7-
plinerved; racemes equaling the leaves, densely few-flowered, bracts
linear-lanceolate, female flowers 5-8 or often solitary; calyx segments
oblong-ovate, subobtuse, the calyx in fruit 5-6 mm. long; ovary
fulvous tomentose, styles biparted, the divisions a little broader
above, sometimes 1-lobed; calyx at opening nearly 3 mm. wide,
both sexes tomentose; male petals oblong-obovate, cuneate at base,
long-barbate, the margins ciliate villous; stamens about 15, the
filaments slightly scabrous; capsules ellipsoid, 8 mm. long, tomen-
tose; seeds smooth. A specimen from the Madrid collection does
not conform to the description but could be C. Ruizianus, of which
nevertheless it may be a part and probably should go to that or /and
C. collinus.
Huanuco: Near Huanuco, (Ruiz & Pavdn, type).
Croton Boissieri M. Arg. Linnaea 34: 138. 1865; 677.
Densely and mostly dichotomously branched small-leaved shrub,
early conspicuously silvery with a close lepidote indument, this
becoming more or less ferrugineous especially on the leaves beneath
and slightly lanate; petioles often 2-3 mm. long; leaves lanceolate-
ovate or ovate-lanceolate, acute at both ends, 2-2.5 cm. long, 8-15
FLORA OF PERU 67
mm. wide, denticulate, subcoriaceous, finally obscurely punctate by
the central remains of the indument; racemes only 2-5-flowered;
bracts setaceous-subulate; larger lance-ovate segments of the fruit-
ing calyx 5 mm. long, the smaller narrower; male calyx 4 mm. long,
the petals lanate at base, glabrous above; stamens about 20; filaments
barbate at base; ovary lepidote as the shortly bifid styles, the lightly
lepidote capsules 6 mm. long; seeds smooth. Suggests some species
from Argentina. F.M. Neg. 5060.
Peru: Without locality, Ruiz & Pavdn, type.
Croton bryophorus Croizat, Journ. Arnold Arb. 21: 96. 1940.
Shrub or small tree, the branchlets tardily glabrescent, early
almost barbate with the abundant indument; petioles 3-5 (10) cm.
long with some minute glands hidden beneath the tomentum; leaves
ovate-elliptic, incised-cordate at base, longish-acuminate or sub-
cuspidate, 7-9 cm. long, 3.5-5 cm. wide, rigid-coriaceous, yellow-
olivaceous and subscabrous, rather resinous-tubercled-corrugate
above, more or less floccose-tomentose beneath, the 6-8 pairs of
ascending quite pinnate nerves notably orange fasciculate-tomentose;
racemes bisexual, the younger with the incurved setaceous entire
5 mm. long bracts, especially of the male flowers, catkin-like, hispid-
lanulose, to 11 cm. long, 3-4 mm. thick; immature male flowers
hispid, 1.5 mm. wide; female 7 mm. broad, lobes linear, acuminate,
about 4 mm. long, disk glands interrupted; capsules glabrescent,
subellipsoid-trigonous, 1 cm. long, 8 mm. broad; seeds lead-colored,
coarsely rugulose-costate, 5 mm. long, 3 mm. broad. Differs from
C. abutiloides HBK., 533, of Ecuador in the peculiar texture of the
leaves and in the comose catkin-like young cymes; probably related
to the C. Baillonianus complex (Croizat).
Ayacucho: Wooded hillside, Carrapa, 2,800 meters, between
Huanta and Rio Apurimac, Killip & Smith 22287, type.
Croton callicarpaefolius Vahl in Geisel. Croton. Monogr. 27.
1807; 532. C. abutiloides HBK. Nov. Gen. & Sp. 2: 86. 1817, probably
at least as to Peru. C. coriaceus HBK. I.e. 87?
Type incomplete; petioles 5-10 cm. long, about equaling the
leaves, these shortly 5-plinerved, basally 2-6 stipitate glandular,
coriaceous-membranous evoluting stellate-puberulent above or some-
times apparently sparsely villous, beneath densely subfloccose tomen-
tose; stipules herbaceous, sublinear, elongate, few-dentate; bracts
linear-lanceolate; racemes of specimen seen all male, open calyx
68 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
4-5 mm. broad; petals spathulate-lanceolate, rigidly lanate-ciliate
(to glabrate?) ; stamens about 40 (30-45), filaments barbate at base.
To this has been referred material from Venezuela as var. pubescens
M. Arg., 532, the leaves villous above, twice as large, ovary rufo-
hirsute, styles biparted, the laciniae incised denticulate; capsules
globose, the seeds biseriately rugulose-costate. Compare note under
C. quadrisetosus, which is probably the same. C. abutiloides HBK.
and C. coriaceous HBK., both of Ecuador, have setaceous or subulate
sepals, the first having male petals subglabrous above, while those
of the second species are cottony-lanate; both these forms seem to
be very similar and Diels, Bibl. Bot. 116: 103. 1937, wrote: it is
possible that C. abutiloides and Vahl's plant are the same species,
but the leaves of the former are not entire; both the Raimondi speci-
mens from Cajamarca were referred to it in herb. Dahlem. F.M.
Neg. 7064 (Fendler).
Piura: Chalaco, Raimondi (det. Dahlem, C. coriaceus). Caja-
marca: Chota, Raimondi. Nancho, Raimondi. Without data,
Jos. de Jussieu, herb. Paris, type. Venezuela? Ecuador? "Tango
grande" (Raimondi), "palo bianco."
Croton churn mayensis Croizat, Darwiniana 6: 466. 1944.
Glabrate shrub or little tree; stipules fleshy, minute; petioles
1.5 cm. long; leaves ovate-lanceolate, basally subcordate or truncate,
apically rather long-acuminate, 3.5-8 cm. long, 2-4.5 cm. wide,
early crustaceous-lepidote scabrous, finally glabrate, the adult brown
and subglabrous beneath, obscurely and irregularly repand-serrulate,
ascending veins about 8 pairs, glands obscure or 2-4, tubulose;
spikes for a long way female, these flowers subsessile, about 3 mm.
broad, 2 mm. long, the 5 broadly triangular foliaceous costate lobes
rather densely indumented, glabrescent within, entire, greenish,
often apically glandular and with 5 glands alternating sepals; ovary
stellate, about twice as long as calyx, styles biparted, 1.5 mm. long;
capsules 3.5-4 mm. long, 3.5 mm. broad, the seeds about 3 mm. long,
ellipsoid, nearly erugulose; male flowers 3-4 mm. broad, 2.5 mm.
long, pedicels 2 mm. long, stamens 15, strongly hispid at base.
Approaches my C. spurcus but seeds never 5 mm. long, female calyx
lobes larger, glands less obvious (Croizat).
Cuzco: Calca, 2,960 meters, Vargas 706. Urubamba, 2,900
meters, Weberbauer 4917. Puno: Churumayo, Soukup 501, type.
Croton collinus HBK. Nov. Gen. & Sp. 2: 78. 1817; 611.
C.ferrugineus HBK. var. collinus (HBK.) M. Arg. in DC. Prodr. 15,
FLORA OF PERU 69
pt. 2: 611. 1866. C. peltoideus HBK., I.e.? C. fermgineus HBK.
var. peltoideus (HBK.) M. Arg. I.e.
Branchlets typically with short appressed indument; petioles
8-10 mm. long or somewhat longer; leaves ovate, acute, rounded
or often slightly peltate at base, 3-5 cm. long, about 3 cm. wide,
subentire or undulate, green (typically) but densely stellulate above,
fuscous tomentose beneath, the pinnate nerves there prominent;
glands sessile, 4 (in type) or apparently often fewer, usually 2 or
even obsolete or concealed; racemes rather short, male bracts 1-5-
flowered, petals and 15-18 (-20) filaments pubescent below; styles
bifid. As remarked by Mueller C. peltoideus HBK. seems scarcely
to differ from the type, with somewhat hirsute-tomentose branchlets,
acuminate leaves peltate, 1.5-3 cm. wide, petioles 12 mm. long,
glands 4-6, stamens 20 or more. The species may well be a high-
altitude form of C. Baillonianus and for that matter passes ap-
parently, as observed by Croizat, intoC. rivinaefolius or C. Ruizianus;
but C. ferrugineus HBK. as to typical form, with somewhat longer
petioles and long-acuminate leaves, seems to be confined to Co-
lombia (Croizat, Caldasia 2: 427. 1944), but of course with as broad
an interpretation as has been given C. rivinaefolius could include C.
collinus as a variety. The young specimen from Huancavelica,
apparently referable here, suggests C. pellitus HBK. of Ecuador,
that however as to type with much smaller leaves. F.M. Neg. 5094.
Cajamarca: Chiclayo to Chota, 1,300-2,200 meters, Weberbauer
4123; 189. Near Cavico at the Rio Chamaya, Ja4n de Bracamoros,
Bonpland, type. Querocotillo, Micuipampa and Montan, Bra-
camoros, (Bonpland, type, C. peltoideus}. Huancavelica: Rio
Huarpa, 2,300 meters, Weberbauer 5669 (det. Croizat, cf. C. fer-
rugineus, var.). Arequipa: Mejia, (Guenther & Buchtien 245?, det.
Bruns, C. ferrugineus}. "Mosquera" (Bonpland).
Croton cuneatus Klotzsch in Hook. Lond. Journ. Bot. 2: 49.
1843; 525.
Striate younger branchlets, petioles (1-2 cm. long) and rachis
densely brown lepidote, this indument soon less dense on the oblong-
or lanceolate-elliptic leaves, becoming sparse beneath, even glabrate
above; leaves shortly cuneate at base or at least acute, cuspidate-
acuminate or acute at tip, the upper below the elongate terminally
clustered or solitary racemes often a dm. or two long, about 5-7 cm.
wide but either smaller or larger, entire or in Peru somewhat serrate,
and with a pair of patelliform glands at base; flowers densely fascicled
70 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
in axils of small oblongish bracts, the uppermost male, the remaining
mixed; calyx lobes oblong-ovate, obtuse, the male 2-3 mm. long,
lanate within, the longer female pubescent (buds globose), both
lepidote without; male petals glabrous without, stamens 14-16,
densely pubescent, anthers ovoid; female petals reduced, glandular,
barbate, ovary lepidote, styles twice cleft; seeds flattened, blackish-
brown. Croizat has indicated an unpublished varietal name for
the Peruvian material. The Tessmann tree was 6 meters tall.
F.M. Neg. 5079.
Loreto: Rio Napa near Mazdn, Mexia 6449; Jose Schunke 43
det. Mansfeld). Iquitos, Tessmann 5127. Rio Itaya, Williams 3300.
Caballo-Cocha, Williams 2272; 2343. Timbuchi, Williams 983 (det.
Standley). Florida, Klug 2003; 2270 (glabrate, leaves more serrate).
To Guianas. "Puma-sacha," "moena" (both Williams), "mamurico-
ey" (Klug; Huitoto).
Croton draconoides M. Arg. Linnaea 34: 90. 1865; 545.
Branches and branchlets covered with depressed stellate trichomes
or these nearly simple; stipules lanceolate-subulate, elongate, entire;
petioles subequaling the broadly cordate-ovate acuminate leaves,
these commonly 10-15 cm. long, 7-11 cm. wide, 5-7-plinerved, patel-
late-glandular at base, membranous, the younger depressed stellulate
becoming puncticulate above, cinereous beneath; secondary nerves
about 12 pairs; racemes very long, the bracts all fasciculately flow-
ered; female calyx segments ovate; stamens 15-18, the filaments
sparsely pubescent below; ovary stellate- tomentulose; styles bi-
parted, slender. Resembles, according to the author, the more
northern C. Draco but lacks the tubercled-stellate trichomes. The
type based on a Sellow specimen from Bahia, Brazil; to this the author
referred two specimens from Peru which are cited here under the
similar C. Lechleri. F.M. Neg. 5087.
Peru(?): Compare note above. Brazil.
Croton erythrochilus M. Arg. Linnaea 34: 93. 1865; 548.
Branchlets dichotomous or subverticillately branched, terete,
the tips as all younger parts shortly stellate-pubescent, finally
grayish-brown with a pulverulent tomentum; stipules subulate,
small; petioles about half as long as the leaves, these 8-10 cm. long,
often 4-5.5 cm. wide at the lower third, rounded or subtruncate at
base, obtusely (or acutely?) caudate-acuminate, entire, at first
densely, finally sparsely, appressed puberulent, biglandular at base,
FLORA OF PERU 71
with 12-17 pairs of nearly equally distant nerves; racemes elongate,
the fruiting 12-30 cm. long, bracts ovate, fruiting pedicels equaling
the calyces, their segments oblong-ovate, subobtuse, small, not at
all accrescent; ovary stellate-tomentulose; styles biparted, divisions
entire; capsules depressed globose, 3.75 mm. long, 5 mm. broad,
subpulveraceous-tomentulose; seeds smooth or shallowly rugulose.
Male flowers unknown in type. In Weberbauer specimen stamens
about 18, villous below, petals narrower than the slightly stellulate
lanceolate sepals. A gummy resinous sap drips in profusion from
incision made in branches (and in some other species) and this may
be one of the Crotons noted from Pozuzo by Ruiz and Pavon the
juice of which "some people gather in calabashes and sell later as
true dragon's blood, and in truth its taste and astringent flavor make
it worth using." F.M. Neg. 5090.
Tumbez: Near Hacienda La Choza, Weberbauer 7688. Huanuco:
Pozuzo (?, see note above), Ruiz & Pavdn, type. Junin: Huacapis-
tana, Killip & Smith 24145 (det. Killip, C. pungens). Apurimac:
Prov. Abancay, Vargas 8967? Cuzco: Cardena, Vargas 6128.
"Sangre de Drago."
Croton flavispicatus Rusby, Mem. N. Y. Bot. Card. 7: 283.
1927. Julocroton peruvianus M. Arg. Linnaea 34: 142. 1865; 704,
not C. peruvianus Briquet. J. peruvianus M. Arg. var. flavispicatus
(Rusby) Croizat, Rev. Arg. Agron. 10: 136. 1943.
Branchlets subtrigonous early angled and densely fulvescent,
with apically stellate trichomes; stipules subulate, 7 mm. long;
petioles 2-4 cm. long; leaves lanceolate-ovate, cordulate or obtuse
at base, long-acuminate, entire to crenulate, (7) 8-12 cm. long,
.(3.5) 3-8 cm. wide, stellate-asperulous above, stellate-tomentose
beneath; spikes dense, 2.5-3 cm. long, the flowers sessile, the bracts
linear-spathulate, acuminate, marginally lacerate above; anterior
female calyx lobes ovate, lacerate-incised, 4.5 mm. long in fruit;
male buds softly tricornute, disk acutely 3-lobed, glands truncate
(male flowers); stamens lanate-fimbriate below; styles short, nearly
free; seeds reticulate-rugose-asperous, 2.5 mm. long, 2 mm. thick.
According to Croizat the Bolivian plant of Rusby differs from
J. peruvianus in the less dense more slender inflorescence. Differs
from C. triqueter in the shorter petioles, bracts and styles and
especially in the acute 3-lobed disk. F.M. Neg. 5209.
San Martin: Tarapoto, Spruce 4290, type; Williams 5401; Ule
6410. Zepelacio, King 3436. To Argentina. "Chivo mashan"
(Williams).
72 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Croton fragrantulus Croizat, Darwiniana 6: 466. 1944. C.
paniculatus M. Arg. Linnaea 34: 87. 1865; 537, not Lamarck, 1768.
Lax, perhaps scandent, the upper branchlets somewhat angled,
all the younger parts fulvous or reddish villous-tomentose with
stellate trichomes, these becoming few on the leaves above; stipules
setaceous, 5-8 mm. long, deciduous; petioles 2.5-4 cm. long, apically
patellate-biglandular; leaves rhomboid-ovate, acute at base, acumi-
nate, 8-12 cm. long or smaller, usually 4.5-5.5 cm. wide, pinnate-
nerved, or shortly 3-plinerved, membranous, minutely serrulate,
secondary nerves 9-13, the lower often approximate; inflorescence
paniculiform, the branches racemiform, terminal, lax or often apex
of racemes dense, either male or female, 5-8 cm. long; bracts seta-
ceous-linear, 3-6 mm. long, flexuous, 1-3-flowered; female pedicels
short, male equaling or exceeding the calyx, this 3 mm. broad before
open, the disk glands fleshy; petals obovate; stamens about 13,
filaments below as receptacle lanate; female calyx with ovate acute
segments; ovary tomentose; styles deeply 4-cleft or -parted, the
divisions entire. Vargas 3694 from Sisal, Cuzco, incomplete, seems
possibly to be referable here or related. F.M. Neg. 5142.
Huanuco: Chacahuasi, Ruiz & Pavdn, type.
Croton glabellus L. Sp. PI. ed. 2. 1425. 1763; 517.
Suggests C. matourensis and C. cuneatus in Peru but lacks the
lower basal glands; the leaves are usually less argenteous than those
of the former but more densely so beneath than those of the latter,
the axillary racemes slender, often branched; leaves variable in size,
rarely acute at base, shortly acuminate; lateral nerves 8-12 pairs;
pedicels often longer, male about 2 mm., female to 6 mm.; calyx
lobes stellate-lepidote as the petals, the female persisting about the
densely lepidote ovary; styles more or less bifid; capsules oblong-
globose, lepidote and slightly tuberculate; seeds smooth. The
racemes are usually much shorter than the leaves, the female flowers
few.
San Martin: Near Tarapoto, Williams 6769 (det. Mansfeld).
Juanjui, King 3774 (det. Standley). Chazuta, King 4010 (det.
Standley). Loreto: Puerto Arturo, Williams 5070 (det. Mansfeld).
Yarina Cocha, Tessmann 3388 (det. Mansfeld). To Mexico and the
West Indies.
Croton glandulosus L. Syst. ed. 10. 1275. 1759; 683.
Shortly stellate-pilose herbaceous annual, usually branched and
several dm. high; stipules setaceous; petioles about a cm. long or
FLORA OF PERU 73
sometimes longer; basal leaf glands 2, patellate; leaves usually
ovate or oblong-ovate, rounded or somewhat truncate at base, obtuse
or acute, crenate, often 3-5 cm. long, 3 cm. or so wide, rather paler
beneath; racemes 1-4 cm. long, the flowers subsessile; male sepals
2 mm. long, petals little longer; stamens 10; female sepals unequal,
3 mm. long, accrescent, spathulate; ovary hirsute; styles nearly
completely biparted; capsules more or less stellate-hirsute; seeds
with small caruncle, minutely pitted, to 4 mm. long. A variant by
many treated as distinct is C. hirtus L'He>. or var. hirtus (L'HeY.)
M. Arg. (subsp. hirtus (L'HeY.) Croizat, Bull. Torrey Club 75:
401. 1948), the stems, particularly, yellow-hispid, glands stiped,
leaves green both sides, seeds said to be only 2-3 mm. long, characters
regarded as specific elsewhere in the genus, probably incorrectly,
but accepted by me because of expediency.
Junin: La Merced, 5292 (det. Croizat; det. Johnston, C. hirtus}.
Rio Pinedo north of La Merced, KiUip & Smith 23584- (det. Mans-
feld). Chanchamayo, Isern 2328 (var.). Huanuco: Pozuzo, Ruiz &
Pavdn (det. Mansfeld). Cuzco: Illapani, Bues (det. Standley).
Warmer Americas.
Croton heliotropiifolius HBK. Nov. Gen. & Sp. 2: 83. 1817;
634.
Branchlets ashy stellate-tomentose, the leaves softly so beneath
but sparsely stellate and green above; petioles to 2.5 cm. long;
leaves ovate, rounded or lightly cordate at base, obtuse or subretuse,
often retuse and mucronate, to 7.5 cm. long, 4 cm. wide, entire,
reticulate-veined, the midnerve prominent beneath, membranous,
eglandular; racemes 7-10 cm. long, the lanceolate-linear bracts and
rachis hirsute, the flowers pedicellate; male calyx double, the teeth
ovate; stamens 16-18; filaments villous at base; female calyx simple,
the teeth linear, hirsute as the ovary; styles bifid; capsules sub-
globose; seeds smooth. Not seen but the branchlets described by
Mueller as in part hirtellous; the original description does not
suggest this but if so the species seems to be distinct from C. rhamni-
folius HBK. with some hispid indument, to which Mueller referred
it as a variety; it seems probable that it is a variant of C. alnifolius
with some of the leaves retuse. F.M. Neg. 5153.
Cajamarca: Chamaya and on the Rio Chinchipe, Prov. Bra-
camoros, (Bonpland, type). Nancho, Raimondi (det. Dahlem, C.
rhamnifolius'). "Tango Chico" (Raimondi).
74 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Croton huitotorum Croizat, Caldasia 3: 9. 1944.
Resembles C. palanostigma; leaves seem to be smaller, the largest
12 cm. long, 10 cm. wide, the indument beneath less dense; female
flowers differ in having the lobes within strongly hispid, style
laciniae 10-12, slender, ovary hispid; male stamens shorter, 2.5-3
mm. long, the filaments not at all dilated at base. It is intermediate
to C. palanostigma and C. Killipiana Croizat of Colombia (Croizat) ;
it seems reasonable to assume that these differences will be shown to
be within the range of a single species-concept, and the material I
have seen from Guiana and Venezuela suggests this; it is noteworthy
that Standley referred the type to C. palanostigma without query.
Loreto: Florida, King 2149, type. "Bogoro-ey."
Croton Lechleri M. Arg. in DC. Prodr. 15, pt. 2: 545. 1866.
C. Draco Schlecht. var. cordatus M. Arg. Linnaea 34: 90. 1865.
Comparable to and maybe not distinct from C. draconoides;
stipules scarcely 2 mm. long, rigid, early dentiform, sublanceolate
but neither subulate nor setaceous; petioles shorter than the leaves,
these 12-20 cm. long, 5-14 cm. wide, the younger more or less vividly
ferrugineous-tomentose both sides becoming glabrescent and stel-
late-puberulent; racemes lax, more than 3 dm. long; stamens about
15, filaments pilose below; capsules 3 mm. long, 4.5 mm. broad,
depressed globose, tridymous. The specific status of this and a
number of forms proposed as distinct remains to be proved. The
Spruce and Poeppig specimens included by Mueller in his C. draco-
noides are cited here following Croizat, Caldasia 3: 17. 1944, but it
seems to me doubtful that C. Lechleri is distinct; the type was referred
by Grisebach to C. alnifolius. Trunk of Tessmann tree -5 dm. in
diameter. F.M. Negs. 5128 (Ruiz & Pavon); 5087 (Spruce).
San Martin: Tarapoto, Spruce 5482. Pongo de Cainarachi,
Klug 2675 (det. Croizat). Huanuco: Chinchao, Ruiz & Pav6n
(det. Mansfeld). Junin: Chanchamayo Valley, Schunke 296; 493.
Loreto: Yurimaguas, Poeppig 1846. Puerto Metendez, Tessmann
3908 (det. Mansfeld, C. draconoides). Above Pongo de Manseriche,
Mexia 6301 (det. Standley, C. palanostigma). Cuzco: Quillabamba,
Vargas 7589. Puno: San Govan, Lechler 2400, type. Without
locality, Ruiz & Pav6n. "Sangre de drago," "palo delgrado"
(Tessmann).
Croton lobatus L. Sp. PI. 1005. 1753; 667. C. riparius HBK.
Nov. Gen. & Sp. 2: 90. 1817, fide Kew Index.
FLORA OF PERU 75
A variable annual often several dm. high, branching, green but
usually sparsely stellate-pubescent and with deeply 3 (-5) -parted
leaves, these on slender sometimes glandular petioles and commonly
3-5 cm. long, their oblanceolate lobes more or less acuminate;
stipules subulate; racemes ordinarily lax; male flowers shortly
pedicelled, glabrous, the sepals elliptic, the petals lanceolate; stamens
10-13; female flowers subsessile, a little glandular; ovary stellate and
pilose, the sometimes 8 mm. large capsules often finally glabrate;
seeds 5 mm. long. A number of variants given recognition by
Mueller; the plant of HBK. was said by the authors to differ in the
leaves being stellate beneath.
Piura: Talara, Haught 41- Cerro Prieto, Haught 36. La Brea,
(Haught & Svenson 11682}. Cajamarca: Chamaya, Bracamoros,
Bonpland. Tomependa, (Bonpland, type, C. riparius}. San Martin:
Tarapoto, Williams 6682. Widely spread as a weed from Mexico.
Croton Macbridei Croizat, Darwiniana 5: 465. 1944.
Shrub or little tree, the innovations early hispidulous; stipules
minute; petioles 2-4 (6) cm. long; leaves exactly ovate-cordate, often
long-acuminate or subcaudate, 6-11 cm. long, 3-5 cm. wide, tomen-
tose both sides, the marginal trichomes subsimple as in C. stenosepalus
but shorter and more delicate, veins 7-10 pairs, the first pair
branched, glands 2, patelliform; spikes slender, to 2.5 dm. long;
male flowers hispid-tomentulose, to 3 mm. large, stamens about 25,
pedicels about 3 mm. long; female subsessile, under fruit scarcely
4 mm. broad, the 5 linear-lanceolate free entire lobes 1.5-2 mm.
long; petals none, disk subentire, slender fruiting column hardly
3 mm. long; capsules 5 mm. long; seeds rounded-rugulose. Affine
C. aequatoris Croizat of Ecuador and C. tarapotensis M. Arg. but at
variance in the soft indument and the exactly cordate leaves; the
Ruiz specimen referred to C. pungens by Mueller could belong here
(Croizat). A duplicate so labeled bears a marked resemblance to
C. peltophorus (a distinct species) var. cuzcoanus Croizat. Certainly
near C. tarapotensis but as to type itself leaves narrowly caudate
and deeply cordate, easily variable characters.
Huanuco: Pampayacu, mouth of Rio Chinchao, 5131, type.
Cuchero, Poeppig 1351? (young, but maybe this; scarcely C. Ruizi-
anus var. cordatus as determined in herb.).
Croton matourensis Aublet, PL Guian. 2: 879. pi. 338. 1775;
550.
76 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Trees beautifully marked (unless in variants) by the contrasting
foliage tones, the oblong-elliptic or elliptic leaves green and glabrous
above, usually conspicuously argenteous with a dense lepidote
indument beneath, this extending to the terminal aggregate racemes,
their flowers usually, except for a few apical ones, female; petioles
2-3 cm. long; leaves acute or rounded at base, shortly acuminate,
usually 1-2 dm. long, 4.5-6 cm. wide, the 2 (1-4) basal glands
depressed or shortly stiped, often also some glands above the base
near and on the entire margins; racemes usually a dm. long or
longer, in Peru apparently half as long, bracts lanceolate or narrower;
male petals lanceolate-spathulate, villous as the 11 stamens; female
sepals lanceolate, medially glabrous within, marginally villous with
simple trichomes, lepidote without as the ovary; styles 3-4-parted
at least above, the divisions entire or 2-3-parted; capsules 3-5 mm.
broad, lepidote; seeds about 2 mm. long. The Peruvian variant
seems to be var. Poeppigianus M. Arg., the styles longer, connate.
King 3355 was given a herbarium name by Croizat and is aberrant
in having a number of male flowers nearly to base of racemes but
the material is scarcely adequate for possible segregation. Williams,
Field Mus. Bot. 15: 272. 1936, noted trees about 30 meters high,
common in the lower Huallaga, the wood used for general construc-
tion. F.M. Neg. 5133.
San Martin: Zepelacio near Moyobamba, King 3355 (det.
Standley). Loreto: Yurimaguas, Williams 4711 (det. Mansfeld).
Fortaleza, Klug 2794 (det. Croizat, var.). Yarina Cocha, Tessmann
3471 (det. Mansfeld). To the Guianas. "Yurac-siprana" (Williams).
Croton nitidulifolius Croizat, Darwiniana 6: 465. 1944.
Innovations fuscous-orange; stipules obsolete; petioles 5 mm.
long; leaves broadly ovate, rounded at base, very shortly acuminate-
apiculate, 2-3 cm. long, quite as wide, rather thick, beneath sub-
orange submetallic pubescent with shortly radiate stellate trichomes,
above densely squamulose, the pinnate veins often impressed, about
6 pairs, dark posterior glands 2; spikes short; male flowers unknown;
female subsessile, 3 mm. long, about 2 mm. wide, the erect broadly
triangular lobes longer than the globose suborange-lepidote ovary;
styles divided. Indument of C. adipatus but differs in character
of female flowers; differs from C. balsameus M. Arg. with similar
flowers in the orange or suborange indument (Croizat); however as
to type female flowers of C. adipatus unknown.
FLORA OF PERU 77
Cuzco: Ollantaytambo, 3,000 meters, Cook & Gilbert 543, type;
536. "Cabra-cabra."
Croton olivaceus M. Arg. Linnaea 34: 93. 1865; 547.
Glabrous at maturity and olive-green including the compressed
branchlets, the younger small ovate stipules and leaves sparsely
pubescent with fulvous depressed-stellate trichomes; petioles slender,
4-7 cm. long; leaves lanceolate-elliptic, cuspidate or acuminate,
9-16 cm. long, 3.5-6 cm. wide, membranous, subentire, biglandular
at acute base with 2 cylindric stiped glands, pinnate-nerved with
about 8-10 pairs arcuate-ascending and incurved-joined; racemes
lax, 1.5-2 dm. long, slender, the rachis glabrous; male pedicels
slender, the glabrous calyces 3 mm. wide; stamens 30-40; filaments
pilose below; petals oblong-obovate, puberulent; female flowers
sessile, scarcely 2.5 mm. long, ovate-lanceolate; ovary fulvous
pubescent; styles biparted; capsules sparsely depressed-stellate;
seeds 7-8 mm. long, smooth except biseriately and lightly undulate-
costate. Color suggests many species of Palicourea, leaf -nervation
species of Tabernaemontana (author). Nearly C. sampatik but may-
be constantly different in glands and number of stamens. F.M.
Neg. 7055.
San Martin: Tarapoto, Spruce 4939, type.
Croton palanostigma Klotzsch in Hook. Lond. Journ. Bot. 2:
48. 1843; 538. C. Benthamianus M. Arg. in Mart. Fl. Bras. 11, pt.
2: 105. 1873, fide Croizat.
Younger branchlets and petioles reddish pulveraceous-flocculose;
stipules 7-8 mm. long, lanceolate, glandular-dentate; leaves broadly
ovate, base cordate or subobtuse, acute, 12-17 (30) cm. long (petioles
about a third as long), 9-14 (28) cm. wide, elobate or unequally
trilobate, patellate-glandular at base, the younger cinereous tomen-
tose beneath, finally more or less softly pubescent with radiately
branched trichomes and the upper surface also punctate-scabrous
with some minute patuliform glands; racemes elongate, bisexual
even below, the lanceolate bracts 1-5-flowered; female calyx turbi-
nate, acute at base, 5-angulate, to 4 mm. long (7 in fruit), 2 mm.
broad, depressed stellate-tomentose (as male), the segments broadly
triangular-ovate; petals obsolete or linear-spathulate; pedicels long,
in fruit as much as 10 mm.; ovary orange-hirsute, style branches
2-parted, each branch 4-6-laciniate (Croizat); stamens often 11,
the hispidulous filaments dilated, 3.5-4 mm. long (Croizat) ; capsules
78 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
small, about 4.5 mm. large, red-hirsute, the nearly smooth seeds
almost 4 mm. long. Williams noted this or some form of it as often
attaining 15 meters, the straight round trunk 3 dm. or more in
diameter. The sapwood and grayish or reddish-brown bark exude
copiously (when injured) a bitter dark red-brown resin used in the
treatment of fractures; see also C. erythrochilus, C. huitotorum, to
which Croizat referred in herb, most of the following specimens.
Illustrated, Mart. Fl. Bras. 11, pt. 2. pi. 20. F.M. Neg. 24471
(C. Benthamianus).
San Martin: Rocky stream banks, Tarapoto, Williams 5844
Loreto: Morona-cocha, Mexia 6512 (det. Standley). Mishuyacu
near Iquitos, Klug 966; 1187. Lower Rio Nanay, Williams 644-
Iquitos, Williams 7979; Tessmann 3610; 5111. To the Guianas.
"Shambu kiru" (Klug); "sangre de drago," "rucurana" (Williams),
' 'uksvakiro' ' (Tessmann) .
Croton pedicellatus HBK. Nov. Gen. & Sp. 2: 75. pi 104.
1817; 639.
A shrub a meter or two high with slender terete finally glabrate
branches, the upper scabrous with short dense stellate indument,
this similar on petioles (these 3-9 mm. long, sometimes a little longer)
and leaves above, the pubescence of these more tomentose beneath;
stipules minute, conical; glands apparently lacking or obscure;
leaves narrowly ovate-lanceolate, long-acuminate, obtuse or sub-
acute at base, 2-3.5 cm. long or longer, 3-10 mm. wide or little
wider; racemes subsecundly few-flowered, the female flowers solitary
or few on pedicels 3-16 mm. long, calyx 2 mm. long; lobes not
accrescent; bracts minute, ovate; male calyx scarcely 1.33 mm. wide,
petals densely fimbriate; stamens 8-12, filaments pilose at base;
ovary fulvous- tomentose; styles glabrous, biparted; capsules villous,
5 mm. long; seeds foveolate-punctulate, 3.5 mm. long. Curiously,
no material of this plant has been sent me.
Cajamarca: Rentema, Bracamoros, (Bonpland, type). Colombia;
Brazil.
Croton peltophorus M. Arg. in Mart. Fl. Bras. 11, pt. 2: 154.
1873.
Branchlets glabrate, the tips appressed fulvous pubescent as the
slender 3-4 (10) cm. long petioles; stipules obsolete; leaves tri-
angular-ovate, broadly rounded or lightly cordate at base, long-
acuminate, often about 10 cm. long, 5 cm. wide, obsoletely repand-
FLORA OF PERU 79
dentate, membranous, 8-10 subpinnate-nerved, sessile-biglandular
at base, olive-green, sparsely appressed pubescent beneath, finally
glabrate, the indument depressed stellate; racemes bisexual, small,
the male bracts minute, 3-flowered, ovate, the female sessile, calyx
about 1.5 mm. wide with linear-lanceolate acuminate not accrescent
segments alternate with ovate obtuse petals or joined with an ample
stiped oblique peltate gland, the narrow male petals villous only at
base, the stamens glabrous, about 16, with anthers nearly twice
as long as broad, the calyx fulvous tomentose; styles free, biparted;
columella of dehiscent fruit scarcely more than 2 mm. long (Croizat).
Var. cuzcoanus Croizat, Darwiniana 6: 468. 1944, has the leaves
below the inflorescence often elliptic, the ovary pale yellow. A
duplicate of a specimen labeled C. pungens in the Ruiz & Pavon
herbarium at Madrid referred by Croizat to C. Macbridei looks
remarkably like his variant of C. peltophorus. F.M. Neg. 24515.
Cuzco: Rio Chaupimayo, Soukup 789 (type, var.). Bolivia;
Argentina and Brazil.
Croton perlongiflorus Croizat, Darwiniana 6: 467. 1944.
A 2-meter shrub with hispidulous-velutinous innovations; petioles
soon about 3 cm. long; glands 2 or more, shortly stiped, more or less
concealed; leaves ovate-lanceolate or ovate, subcordate at base,
shortly acuminate, 4-7 cm. long, 2-4 cm. wide, adult glabrate
above, the younger yellowish tomentose beneath, finally tomentose,
veins sometimes well impressed, about 7 pairs, the first branched,
subentire; inflorescences nearly flagelliform, to 3 or 4 dm. long,
often bisexual; male flowers 3.5 mm. wide, 2.5 mm. long, stamens
often 20-25; female flowers subsessile or sessile, scarcely 2 mm.
broad, 2.5 mm. long, lobes triangular-acuminate, alternate glands
manifest; ovary hispidulous; styles biparted, 2 mm. long, tomentose
at base, fruiting column 3 mm. long. Allied to C. thurifer HBK.
and C. spurcus Croizat; in the thicker leaves and impressed veins
simulating C. Baillonianus but easily distinguished by the smaller
male flowers (Croizat). But compare also C. Ruizianus and allies
which area is noted for intermediate forms. C. Mandonis M. Arg.,
532, Bolivian, would key here but stamens many, spikes shorter
than upper leaves, these larger with 2-4-stiped glands. It is probable
that this form, however related, is typically only in Huancavelica.
Huancavelica: Tayacaja, 1,600 meters, Weberbauer 6518, type.
Cuzco: Valle de Urubamba, Ollantaytambo, 2,800 meters, (Herrera
80 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
2789). Cahacaha, Hacienda Tilque, (Herrera 1668). "Jaratu"
(Weberbauer).
Croton perspeciosus Croizat, Journ. Arnold Arb. 21: 98. 1940.
Little tree, the younger parts sordidly subsericeous with a hispid
indument; stipules pinnatifid, to 13 mm. long, 3.5 mm. wide; petioles
7-14 cm. long, with 4-8 stiped glands at apex; leaves ovate, definitely
cordate, long-triangular-acuminate, shortly mucronate, 12-24 (30)
cm. long, 7.5-14 (20) cm. wide, dentate or rarely sublobulate,
margin minutely denticulate, glabrescent above except the 7-8 pairs
of hispidulous veins, the first pair branching; racemes bisexual, none
seen longer than 1 dm.; immature male flowers 2.5 mm. wide, pedicels
4-5 mm. long; female pedicels after flowering 2.5 mm. long, calyx
scarcely 1 cm. broad, lobes triangular, entire, to 4 mm. long, 2.5 mm.
wide, glabrous within; seeds coarsely few-rugose, 6 mm. long, nearly
as broad, caruncle to 2 mm. broad. Suggests C. speciosus M. Arg.,
528, but that with much wider female calyx segments; certainly
distinct from C. quadrisetosus in pubescence and in the character of
flowers (Croizat). Compare C. callicarpifolius and C. abutiloides of
Ecuador, perhaps the same but both typically with denser shorter
tomentum.
San Martin: San Roque, Williams 7378. Ayacucho: Open woods,
Aina, 1,000 meters, Killip & Smith 22841, type. Carrapa, between
Huanta and Rio Apurimac, densely forested valley, Killip & Smith
22402. Cuzco: Santa Ana, 900 meters, Cook & Gilbert 1642. Rio
Marcapata, Vargas 3066. Pasco: Quillasu, 1,700 meters, Soukup
3313. "Sangre de drago."
Croton peruvianus Briquet, Ann. Cons. Jard. Bot. Geneve
4: 228. 1900.
Upper branches densely stellate pubescent with widely spreading
trichomes; stipules minute; petioles 2-2.5 cm. long; leaves ovate-
lanceolate, rounded-subcordate at the biglandular base, caudate-
acuminate, 8-9 cm. long, 3.5-4.5 cm. wide, above green but sparsely
pubescent with simple trichomes, cinereous beneath with dense
stellate pubescence, entire, nervation pinnate, the lateral ascending
nerves about 8 pairs, scarcely prominent either side; racemes a dm.
long or longer, rachis stelligerous; male pedicels 2 mm. long, glabrous
or stellate as the ovate calyx segments, these 15-22 mm. long;
stamens 16, sericeous-hispid below the upper third. Likened in
herb, by Mueller to C. tarapotensis M. Arg. and C. soratensis M. Arg.;
FLORA OF PERU 81
differs from first in indument, form and nervation of leaves, from the
second even more by the absence of the stellate trichomes; our
specimen seems to approach C. lachnostachys Baillon with, however,
different foliage (Briquet). The midnerve has been described by
Croizat as straw-colored with branched tomentum. Apparently not
distinct from C. stenosepalus. F.M. Neg. 24517.
Amazonas(?): Without data, Mathews 3019, type.
Croton Pilgeri Ule, Verh. Bot. Ver. Brandenb. 50: 74. 1908.
Sometimes several meters tall, the younger branchlets, petioles,
these 1.5-2 cm. long, and the 5 principal pinnate leaf -nerves bright
reddish stellate- tomentose; stipules 3 mm. long, caducous; leaves
broadly ovate-elliptic, long cuspidate-mucronate, biglandular at base
beneath where whitish-green with dense stellate tomentum, dark
green but stellate-asperous above, 9-12 cm. long, 5-6 cm. wide,
with some capitate marginal glands; racemes dense, subequaling the
leaves or about 1 dm. long, bracts 1 mm. long, the lower female,
sessile, with 5 equal lanceolate segments 2 mm. long, glabrous within,
stellate without as the male calyx, this with pedicels 2-3 mm. long,
elliptic segments 2 mm. long; stamens 15-18; ovary as capsules
tomentose-stellate; styles 2-4-parted; seeds 3 mm. long, ellipsoid,
blue-gray, netted and ribbed. Maybe a variant of C. tarapotensis.
F.M. Neg. 5146.
San Martin: Tarapoto, Ule 6411, type; Williams 5409; 5536.
"Supayerillo."
Croton pungens Jacq. Icon. Rar. 3: 19. pi. 622. 1786-93; 540.
Becoming a small tree, the elongate terete branchlets above as
well as stipules (5 mm. long), petioles (3-7 cm. long), leaves beneath
and the many-flowered racemes shortly tomentose with stellate
trichomes or sometimes some longer villous pubescence present;
leaves cordate-ovate, cuspidate-acuminate, 9-15 cm. long, 4-8 cm.
wide, basally 4-12-stipitate-glandular, 5-plinerved, denticulate,
sparsely stellate-puberulent above; racemes 15-25 cm. long, often
bisexual below; male calyces early 2.5 mm. wide, the basally cuneate
petals barbate; stamens 40-65, filaments as receptacle lanate;
pedicels slender, fruiting nearly 4 mm. long; ovary sparsely stellate;
styles biparted, elongate; capsules globose, tridymous, 6 mm. long,
7 mm. broad, stellulate; seeds thick, coarsely undulate-subrugose.
Endemic to Venezuela and even if broadly interpreted does not occur
farther south than eastern Colombia, but C. sarcopetalus of southern
82 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Bolivia and Argentina is easily confused with it (Croizat). How-
ever, Mueller referred to it a Ruiz & Pavon specimen without data
which Croizat has suggested may be C. Macbridei.
Peru (doubtful except sens. lat.). To Central America, Vene-
zuela, and Brazil.
Croton pycnanthus Benth. PL Hartw. 247. 1846; 536.
Rigid, more or less copiously fulvous stellate-tomentose shrub ex-
cept the subcoriaceous broadly ovate obtusely acutish leaves stellate
pubescent above, these lightly cordate at the 2-6-glandular base,
5-7-plinerved, often 4-7 cm. long, 3-6 cm. wide, crenate-denticulate;
racemes compactly many-flowered, 10-13 mm. thick, with rigid
linear-lanceolate bracts; pedicels short; female calyx 4 mm. long,
the segments broadly ovate, subobtuse; petals subulate, apically
glandular; ovary stellate-tomentulose; styles biparted, the divisions
bifid; male calyx 2 mm. wide, the cuneate petals lanate below;
stamens 20-22; filaments hirtellous nearly their entire length.
Perhaps should be drawn to include the similar C. Baillonianus var.
submembranaceus with longer petioles and pedicels. F.M. Negs.
5149, 32537.
Peru (probably; cf. note above). Ecuador.
Croton quadrisetosus Lam. Encycl. 2: 208. 1786; 531.
Upper branchlets reddish-succulent, densely pubescent as the
petioles (3-4 cm. long) and elongate racemes, with a fulvous or early
ferrugineous flocculose indument; stipules lanceolate-subulate, rigid,
about 6 mm. long; leaves cordate-ovate, acuminate, 8-10 cm. long,
5-8 cm. wide, coriaceous-membranous, denticulate, punctiform-
scabrous above with small trichomes, tomentose beneath, 5-plinerved
and 4 (2) -glandular at base, their stout floccose or glabrous stipes
sometimes 4-5 mm. long, nearly as broad as the glands; bracts linear-
lanceolate, the pedicels of both sexes little shorter than the calyces,
this in the male flower opening 3-4 mm. broad, the obovate petals
pubescent, stamens about 50-60; female calyx segments lanceolate-
ovate, styles petaloid-dilated, much divided above, the laciniae
entire; capsules globose, over 13 mm. broad. The Bolivian C.
Mandonis M. Arg., 532, has narrower bracts, style laciniae denticu-
late. In all probability, as suggested by Croizat, Journ. Arnold
Arb. 21: 99. 1940, C. quadrisetosus is the earlier name for C. calli-
carpaefolius Vahl as to type. F.M. Neg. 39570.
FLORA OF PERU 83
Cajamarca: Hualgayoc, Monte Seco, 1,800 meters, Soukup 3837.
Junin: Palca, Soukup 3480. Without locality, Dombey, type,
herb. Paris. Cuzco: Idma, Vargas 8549.
Croton Rehderianus Croizat, Journ. Arnold Arb. 21: 100.
1940.
Subherbaceous shoots covered with a harsh beautiful reddish
tomentum; petioles canaliculate with sessile binate glands both
sides, the anterior smaller, 3.5-4 (8) cm. long; leaves ovate-caudate
or sublanceolate-acuminate, cordate or incised cordate at base, 12-
22 cm. long, 5-9 cm. wide, olive to reddish asperously lepidote-
stellate above, grayish-flocculose stellate-tomentose beneath, the
yellowish pubescent veins unequally 10-12-paired, ascending, sub-5-
plinerved, margins lightly glandular serrulate; racemes seen scarcely
longer than 5 cm. but probably often longer, bisexual but male
flowers unknown; female bracts subequaling the solitary subsessile
calyx, about 3.5 mm. large, lobes triangular to 2 mm. long, 0.5 mm.
wide, ovary coarsely tomentose, globose; styles shortly bifid, gla-
brous, about 3 mm. long. Name commemorates the able student
of ligneous plants, particularly those of northern zones. Affinities
of the species apparently are to be looked for in the vicinity of
C. bryophorus (Croizat).
Cuzco: Brush-covered hillside, 2,200 meters, below Machu-
Picchu, West 6464, type. "Mocco-mocco."
Croton rivinaefolius HBK. Nov. Gen. & Sp. 2: 80. 1817; 628.
C. alnifolius HBK., I.e., not Lam. C. Pavonis M. Arg. Linnaea 34:
115. 1865; 609, fide Svenson. C. Jamesoni M. Arg. in DC. Prodr. 15,
pt. 2: 549. 1866, fide Svenson.
More or less cinereous stellate-puberulent bush, variable also in
size and shape of leaves; branchlets rigid; stipules subulate, small;
petioles only a few mm. (-10) long; leaves usually ovate or -obovate,
sometimes suborbicular, obtuse or acute, obsoletely or evanescently
to prominently biglandular at base, typically 12-13 cm. long, 4-6
cm. wide, sometimes larger or often about half as large, somewhat
dentate or subentire, finally glabrate above, pinnate-nerved or also
shortly 3-plinerved; flowers apparently always dioecious (Svenson);
racemes short or long, dense, the rachis more or less angled; bracts
ovate-lanceolate, 1-3-flowered ; male pedicels rigid or reflexing about
as long as globose calyx, this 1.5 mm. broad, petals cuneate at base,
lanate-ciliate; stamens about 15, filaments pilose or glabrous; female
84 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
flowers sessile, calyx segments narrow, not accrescent; ovary tomen-
tose; styles biparted, the branches slender; capsules subglobose,
lightly tomentose or glabrescent; seeds small. Compare C. alni-
folius which in Piura seems to be the same. I have compiled in part
from Svenson, Amer. Journ. Bot. 33 : 458-459. 1946, and the descrip-
tion is too broad if related forms are to be kept as species, for he
remarks, "close to C. Ruizianus M. Arg. of the Lima region [cf.
especially Worth & Morrison 15692] but differs in the less pubescent
capsules, less rugose leaves and smaller seeds"; such comparative
differences are of course meaningless. As he notes, the type of
C. Pavonis may well have been collected by Tafalla and actually
then came from Ecuador near Guayaquil. F.M. Negs. 7084 (C.
Pavonis); 8507 (C. Jamesonii).
Peru (doubtful; cf. Svenson as noted above). Ecuador; Gala-
pagos.
Croton rubiginosus Croizat, Journ. Arnold Arb. 21: 101. 1940.
Dichotomously or verticillately branched little shrub scarcely 5
dm. high, the shoots and leaves rusty lepidote-stellate; petioles
slender, eglandular, 5 mm. long; leaves ovate or ovate-acuminate,
rounded at base, 1.5-2 cm. long, 8-10 mm. wide, concolor, incon-
spicuously serrulate, with 3-5 pairs of obscure ascending veins;
racemes bisexual to 1.5 cm. long; male flowers about 2 mm. large,
stamens 8-10; petals glabrous, twice as long as the lobes; female
pedicels barely 1 mm. long, calyx about 4 mm. long, 3 mm. broad,
lobes marginally with a few glandular trichomes, 2 mm. long, 1.75
mm. wide, more or less elliptic, apex rounded; ovary subtrigonous,
lanulate-tomentose, nearly glabrous when submature, glabrous styles
parted to base. Not allied to any species at present known from
Peru (Croizat). The actual type sheet was at Berlin-Dahlem,
presumably destroyed.
Amazonas: Open grassy land, eastern bank of the Maranon
above Balsas, 2,000 meters, Weberbauer 4273, type.
Croton Ruizianus M. Arg. Linnaea 34: 89. 1865; 541. C.
pungens Jacq. Icon. Rar. 3: 19. pi 622. 1786-93; 540, probably as
to Peru or the earlier name.
Often low and rather stockily branched, the growing tips and
younger leaves densely and shortly tomentose with somewhat reddish-
fulvous stellate trichomes, these becoming sordid gray and more or
less deciduous from the leaves above; stipules obsolete; petioles often
FLORA OF PERU 85
only a cm. long; leaves ovate, rounded at base, acute, or sometimes
merely apiculate, often 5-6 cm. long and nearly as wide (sometimes
much smaller); lower nerves pinnate but more or less approximate;
basal glands 2, sessile; racemes short, few-flowered, at most moder-
ately long, the narrowly ovate-lanceolate or sublinear female calyx
lobes shorter than ovary and not accrescent in fruit; male petals
cuneate, barbate; stamens (20) 30-40, villous below; ovary densely
stellate-tomentose; capsules subglobose, about 6 (7) mm. long,
slightly broader; seeds dark gray or brownish, about 5 mm. long,
lustrous and smooth unless for some undulation. The var. ovatus
M. Arg. is taken as the typical form: the var. cordatus M. Arg. I.e.
is apparently distinct by virtue of the elongate lax racemes, longer
petioles and the openly and deeply cordate almost caudate-acumi-
nate membranous leaves; Croizat, Darwiniana 6: 448, suggested
that this variety is the same as the Bolivian (not Peruvian as to
type as Croizat states) C. Frieseanus M. Arg., 543, which ex char,
has merely acutish lightly cordate leaves, 18 stamens and denticulate
glandular stipules, doubtfully specific characters. C. Ruizianus
seems to be distinguishable from C. pungens and it is probable that
the Ruiz & Pavon specimen so referred by Mueller belongs as
Croizat suggests, to C. Macbridei. As Svenson remarks, Amer.
Journ. Bot. 33 : 459. 1946, C. Ruizianus is very much like the typically
more northern C. rivinaefolius with less pubescent capsules, less
rugose leaves, smaller seeds; these differences it would seem could
easily be included within a reasonable range of species' variation,
but here as elsewhere in Croton the decision ought to be made by
an able monographer. I have been tempted to include all these
forms in C. collinus, which compare. The Cuzco plant apparently
passes into C. Baillonianus and varies in petiole length; the central
Peruvian plants are separable intangibly as C. balsameus. The
typical form spreads a meter high or less; my collections with a
citrous odor, as in C. balsameus, this probably the same. F.M.
Negs. 5154; 34157 (var. cordatus).
Lima: Huaura and Sayan, Ruiz & Pav6n, type. Above Huaytara,
Pisco region, Weberbauer 5423. San Buenaventura, open rocky
slopes, 2,700 meters, Pennell 14524. Matucana, in rocks, 217;
336; 2882 (det. Johnston) ; Stork & Norton 9135; Soukup 2062 (det.
Croizat, C. Frieseanus); Grant 7477 (det. Croizat, C. Frieseanus).
Ancash: Ocros, Weberbauer 2728. Junin: Quequena, Isern 2040.
Arequipa: Sandy plain north of Atiquipa, Worth & Morrison 15648
(det. Johnston). Posco, Rose & Rose 18806; (Guenther & Buchtien
86 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
250, det. Burns). South of Chala, Worth & Morrison 15692 (det.
Johnston). South of Arequipa, Eyerdam & Beetle 22131 (toward
C. collinus?}. Cuzco: Open rocky slope, Ollantaytambo, Pennell
13668; Cook & Gilbert 536. Piri, Vargas 5929. Sisal, Prov. Anta,
Vargas 7282. Moquehua: Carumas, open mixed formation, 2,700
meters, Weberbauer 7353 (det. Johnston, C. Mandonis Croizat, var.
cordatus). Without locality, Ruiz & Pav6n (type, var. cordatus).
Bolivia? "Upalu" (Weberbauer), "cabra-cabra" (Cook & Gilbert).
Croton sampatik M. Arg. Linnaea 34: 94. 1865; 548.
Branchlets subterete with a few leaves crowded at their tips and
as all younger parts shortly ferruginous stellate; stipules denti-
form, subulate; petioles 4-7 cm. long; leaves oblong-ovate, obtuse
at base, acuminate, 8-17 cm. long, 3-7 cm. wide, now obscurely,
now deeply lobate-dentate, fragile but firm-membranous, very green
but with some scattered depressed-stellate trichomes both sides,
5-7 pairs of distant lateral nerves, the lower two approximate,
patellate-glandular at base; racemes subaggregate (often 3) at apex
of branchlets, elongate, 15-20 cm. long, sometimes basal female
flowers lacking, the male linear-lanceolate bracts 3-7-flowered, the
calyx globose, 2.5 mm. wide before open; petals obovate, puberulent;
stamens about 14, filaments glabrous, anthers short-ellipsoid; female
calyx about twice as long as broad; ovary tomentulose; styles bi-
parted, the branches entire. Commonly the male and female
flowers are mixed below; C. Fraseri M. Arg., 603, of Ecuador, known
only from a completely female spike, seems to be the same. The
Weberbauer tree was 25 meters tall. F.M. Neg. 5157.
Huanuco: Chicoplaya, Ruiz & Pavdn, type. Junin: La Merced,
Weberbauer 1889; 282. Brazil. "Sampatik" (Ruiz).
Croton sapiiflorus Croizat, Darwiniana 6: 464. 1944.
Innovations and leaves beneath densely velvety tomentose with
a pale-ochraceous or cinereous indument; stipules promptly caducous,
triangular, 2 mm. long; petioles tomentulose, 1-3 cm. long; leaves
elliptic or subobovate-elliptic, cuneate at base, shortly acuminate,
7-11 cm. long, 3-5 cm. wide, glabrate above, entire, with about
9-12 pairs of conspicuous ascending pinnate nerves, glands none or
faint; spikes slender, uni- or bisexual, to 2.5 dm. long; male flowers
about 2.5 mm. broad, stamens 15-20; female flowers immature but
sessile, lobes more or less lanceolate, entire, 2-3 mm. long, 1.5-2.5
mm. wide, tomentulose, glabrate within; ovary truncate, hispidulous,
FLORA OF PERU 87
the separate 5 glands ellipsoid; petals minute, alternate with sepals;
styles tomentose or hirsute, 4-5 mm. long, parted nearly to base.
Probably not at all remote from C. leptostachys HBK. of Colombia
(author).
Huanuco: Canyon below Rio Domingo, J^.218, type.
Croton scaber Willd. Sp. PI. 4: 550. 1805; 559.
Branchlets slender, terete, shortly and rather sparsely as all
younger parts reddish or fulvous tomentulose; stipules semi-ovate,
semi-auriculate at base, setaceous above, 5-7 mm. long; petioles
1-2 cm. long; leaves ovate, lightly cordate at base, shortly acuminate,
10-12 cm. long, 4.5-6 cm. wide, membranous, subentire, early
sparsely radiately lepidote, eglandular at base, 5-plinerved with 7-9
pairs of lateral nerves; racemes rather short, dense, bracts lanceolate-
setaceous, 1-flowered, the shortly pedicelled female calyx with broadly
ovate obtuse segments accrescent, in fruit 8-9 mm. long; ovary
tomentulose; styles bifid to below the middle, the branches subentire;
male calyx early about 3 mm. wide, 7 mm. wide when open; petals
oblong-obovate, tomentose without, villous within, lanate ciliate;
stamens about 15, filaments pilose entire length; capsules globose,
tomentose; seeds minutely subimpressed puncticulate. Mueller
recognized several variants, the type being from Caracas; it is
doubtful that a Ruiz & Pavon specimen referred here by him was
correctly assigned; possibly mislabeled. C. rhamnifolius HBK.,
634, apparently in Bolivia, may be expected; it has some hispid
pubescence mixed with the tomentosity, eglandular or partly
glandular. F.M. Neg. 5162.
Peru(?): Without locality, (Ruiz & Pavdn). Venezuela.
Croton spurcus Croizat, Darwiniana 6: 467. 1944.
Glabrate shrub; petioles 1-3 cm. long; leaves ovate-elliptic,
broadly rounded-truncate-cordate at base, more or less acuminate
at apex, 2.5-7 cm. long, 2-4 cm. wide, irregularly and distantly
glandular-serrate or subentire, ascending pinnate veins about 7
pairs, glands 2(-4), stipitate, patelliform; male pedicels 1.5-2 mm.
long, buds about 1.5 mm. large; stamens 13; female flowers about
2 mm. wide and quite as long, brownish-orange stellate as pedicels,
the broadly triangular lobes alternate with conspicuous substiped
glands; ovary tomentose; styles biparted, 3-4 mm. long; capsules
glabrescent, stipitate glands obvious, 7-8 mm. long, 5-6 mm. wide,
88 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
the seeds to 5 mm. long. Like my C. churumayensis but larger
capsules and seeds, smaller female perianth lobes, stiped ovary glands
(author).
Lima: Amancaes, 500 meters, Weberbauer 5249, type.
Croton stenosepalus M. Arg. Linnaea 34: 114. 1865; 607.
Fulvous hispid-stellate especially the branchlets, setaceous
stipules (6 mm. long), petioles (early short, finally 5-9 cm. long)
and leaves beneath, these with 2 sessile glands at the subacute base,
elliptic, cuspidate-acuminate, 7-12 cm. long, 4-8 cm. wide, subentire,
rigid-membranous, more or less villous above with slender subsimple
trichomes, pinnate-nerved; racemes dense, long as leaves, bracts
setaceous; female flowers rather many, sessile, the subulate calyx
parts acuminate, 5 mm. long; ovary hirsute; styles deeply parted,
slender; male calyx at anthesis 2 mm., filiform-pedicellate; petals
villous only at base; stamens about 15, the filaments hirsute below;
capsules muricate-hispidulous, subtruncate both ends, 6-7 mm.
long and broad; seeds fully 5 mm. long, lightly costate. Very
distinctive by the long lax trichomes dispersed over the upper leaf
surface. F.M. Neg. 5171.
San Martin: Tarapoto, Spruce 4160, type; Williams 5433 (det.
Mansfeld).
Croton tarapotensis M. Arg. Linnaea 34: 114. 1865; 608.
Younger parts ferruginous tomentose; stipules small, lanceolate-
subulate; petioles finally about as long as the leaves, these sometimes
1-1.5 dm., about half as wide, shortly 3-plinerved with 2 appressed
glands at the rounded or somewhat cordate base, broadly or oblong-
ovate, more or less cuspidate-acuminate, denticulate, lower secondary
nerves remote from rest; racemes dense, female flowers many,
their bracts conspicuously uni-patellate-glandular both sides, calyx
segments ovate-lanceolate, the 2 posterior finally distinctly smaller,
not accrescent; ovary tomentose; style branches rigid; male calyx
nearly 2 mm. broad at anthesis, the pedicels suberect; petals sub-
glabrous except at base; stamens 15 or 16; capsules subglobose,
4.5 mm. long, the seeds coarsely costate, 4 mm. long, 2.5 mm. broad
(Croizat). Certainly not in Argentina (Croizat). F.M. Neg. 7086.
San Martin: Tarapoto, Spruce 4138, type. Lamas, Williams
6439. Chazuta, King 3980 (det. Croizat).
Croton Tessmannii Mansfeld, Notizbl. Bot. Gart. Berlin 9:
264. 1925.
FLORA OF PERU 89
Younger branchlets angulate, lepidote; petioles 2-3 cm. long;
leaves oblong-elliptic or elliptic, slightly narrowed to the acute or
subrotund base, acute or shortly acuminate, 7-19 cm. long, 2.5-6.5
cm. wide, lustrous and glabrous above except sparsely lepidote,
especially on the midnerve, more densely lepidote but opaque
beneath, obscurely 3-5-nerved at base, biglandular and minutely
or repandly glandular-denticulate; racemes terminal, to 26 cm. long,
lower bracts bisexual, the upper male, their lepidote pedicels to 6 mm.
long, sepals and petals 3 mm. long, the former ovate, pilose both
sides, the latter glabrous without; stamens 15; filaments glabrous,
receptacle villous; female flowers with lepidote ovate obtuse sepals
3.5 mm. long, petals 3 mm. long, linear; ovary lepidote; styles
dichotomously divided, to 4 mm. long. A little tree, the trunk 10 dm.
in diameter. Allied by the author to C. surinamensis M. Arg., 525,
with pinnate nerved leaves equally lepidote both sides. F.M. Neg.
5186 (Tessmann 3334).
Loreto: Flooded areas, Yarina Cocha, Tessmann 3438, type.
Cumaria, Tessmann 3334 (fide author).
Croton thurifer HBK. Nov. Gen. & Sp. 2: 76. 1817; 606.
Softly stellate-tomentose shrub with the more or less inequilateral
rhombic-ovate or rotund leaves of the related C. alnifolius but the
first pair of nerves usually elongate at least to the middle of the
blade; petioles 1-2.5 cm. long, hispidulous and densely tomentulose
as the lightly angled branches; leaves obliquely rounded to acute
or often cuneate at base, rounded, obtuse or shortly acute at the
often oblique apex, 3-8 cm. long, 2-8 cm. wide, more or less glandular
at base, the glands at least sometimes shortly stiped; racemes dense
or lax when well-developed to a dm. long or longer, the rachis angled,
the bracts minute; female calyx cupulate, the small ovate segments
not at all accrescent; male petals lanate below; stamens about 16,
pilose at base; ovary hispid and stellate as the rather rigid short
styles below the tip. Two meters high (Weberbauer). F.M. Neg.
5187.
Cajamarca: Tomependa, inundated area along Rio Maranon,
Bonpland, type. Valley of the Rio Shumba, 700 meters, Weberbauer
6175 (det. Croizat).
Croton trinitatis Millsp. Field Mus. Bot. 2: 57. 1900. C.
chamaedryfolius (Klotzsch) Griseb. Fl. Brit. W. Ind. 41. 1864, not
Lam. C. Miquelensis Ferguson, Rept. Mo. Bot. Gard. 12: 49. 1901
90 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
(C. Miquelianus Ferguson ex Lanj. in Pulle Fl. Surinam 2: 38. 1932).
Geiseleria chamaedryfolia Klotzsch in Hook. Lond. Journ. Bot. 2: 47.
1843.
Branches and usually stems slender and more or less stellulate-
scabrous with appressed trichomes; petioles commonly a few mm.
long, the ovate leaves 2-3 cm. long, about 1.5 cm. wide or in luxuriant
states petioles 5 cm. long or longer, leaves 7 cm. long, 4 cm. wide,
always with a pair of stiped glands at the rounded or subcordate
base, gradually acuminate and rather to very coarsely crenate-
dentate, paler and somewhat stellulate beneath; racemes axillary
or terminal, few-flowered, bracts basally bilobed, pedicels short;
male calyx 4-5-merous, the ovate segments acicular pubescent
without; stamens 8-10, pilose at base with the receptacle; female
flowers with sepals often unequal, stellulate, petals reduced, disk
annular; ovary hirsute-stellulate as capsules, these 3-4 mm. long;
seeds olive-gray, 3 mm. long, obscurely lineate-puncticulate.
Neither Lanjouw nor Standley and Steyermark, Fieldiana: Bot. 24,
pt. 6: 80. 1949, bothered to indicate the reason for the adoption of
the later name of Millspaugh; common as a weed near Iquitos, only
a few collections cited. Illustrated, Miq. Stirp. Surin. Sel. pi. 30.
^
San Martin: Tarapoto, Williams 5408. Iquitos, Kittip & Smith
26921. Yurimaguas, Williams 3824- Lower Rio Nanay, Williams
264; 474- Alto Rio Itaya, Williams 3231; 3294. Caballo-Cocha,
weed in forest, Williams 2476; 2477 (leaves large, det. Standley).
Pebas, Williams 1958. To Mexico and the West Indies. "Sinchi
pichana" (Williams).
Croton triqueter Lam. Encycl. 2: 214. 1786; 704. Julocroton
triqueter (Lam.) Baillon, Adansonia 4: 368. 1863-4.
Branchlets compressed-trigonous, the trichomes not at all fastigi-
ately radiate, in the angles and at the tips shortly rufous floccose
and horizontally stellate; petioles early 3-4 cm. long, the lower
becoming as long as the triangular-ovate leaf-blades, these often
8-12 cm. long, 3-8 cm. wide, more or less cordate, long-acuminate,
crenate, shortly and densely stellate pubescent above, rather finely
stellate tomentose beneath; spikes nearly strobiliform, 2.5-3.5 cm.
long, 10-12 mm. thick or in fruit at least twice as long as broad;
buds softly tricornute; flowers subsessile, the male calyx segments
finally 4 mm. long, pubescent petals spatulate-linear, filaments
pilose, the fruiting calyx segments 7 mm. long, 3 anterior broadly
FLORA OF PERU 91
ovate, deeply pinnate-lacerate; hypogynous disk entire; capsules
4 mm. long, the rugose-asperous seeds 3-3.5 mm. long.
Peru: According to Croizat. Mexico to Venezuela and Bolivia.
Croton Tyndaridum Croizat, Journ. Arnold Arb. 21: 101. 1940.
Tree, about 7 meters, the shoots early covered with a dense capi-
tate-stelligerous indument; petioles 7-13 cm. long, the dense tomen-
tosity partly concealing the 2 apical glands; leaves exactly ovate-
rotund, shortly acuminate, 10-16 cm. long, 10-14 cm. wide, gray
tomentose beneath, olivaceous and partly glabrescent above, sub-
entire, veins about 8 pairs, the beautifully branched first pair sub-
triplinerved; racemes to 2.5 dm. long, only female observed, these
at anthesis on pedicels about 0.3 mm. long; calyx 5 mm. broad, 3 mm.
long, lobes ovate, acuminate, 3 mm. long, 1.5 mm. wide; ovary 4 mm.
broad, hispidulous-tomentose; styles parted to base, scarcely 2.5
mm. long. The author has named the plant for the mythological
Tyndarides in honor of the botanical brothers Killip and Smith!
He suggests that the species, unlike any other Peruvian form, may
prove to connect C. Urucurana Baill., 545, of Brazil and C. gossypii-
folius Vahl, 538, of Venezuela.
Junin: Along Rio Perene*, near "hacienda 3," in thickets, 600
meters, (Killip & Smith 25229, type).
Croton xanthochylus Croizat, Journ. Arnold Arb. 21: 102.
1940.
Small tree to 4 meters high, the tips sparsely pubescent with a
strongly appressed fasciculate-stellate indument; petioles slender,
2.5-4 cm. long, with 2 subsessile patellate black glands at apex;
leaves ovate-cuspidate, cordate at base, 6-8 cm. long, 4-5 cm. wide,
glabrous above but lightly granulose under a lens, deciduously gray
tomentulose beneath, membranous, entire, with about 8 pairs of
slender anastomosing veins; racemes bisexual, slender, to 4 cm. long;
male pedicels about 2 mm. long; stamens 10-12; female flowers
subsessile; calyx scarcely 2 mm. large, setaceous lobes to 1 mm.
long; ovary globose, about 1.5 mm. large, yellowish tomentulose;
styles scarcely 1.5 mm. long; capsules globose, subglabrous, about
3 mm. large, the brown seeds 2.5 mm. long, 2 mm. broad, coarsely
rugose and puncticulate. Very close to C. erythrochilus, which has
a sublepidote argillaceous indument, female pedicels distinct, about
1.5 mm. long; C. Lechleri is more robust, similar capsules but the
92 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
larger leaves with texture and indument of those of C. draconioides
(Croizat). The type was lemon-scented.
Junin: La Merced, light-barked tree, sandy flat, 5308, type.
Along Rio Perene* near Hacienda "3," Killip & Smith 25172 (det.
Croizat with query).
10. ARGYTHAMNIA P. Br.
Ditaxis Vahl ex Juss. Euphorb. Tent. 27. pi. 7. 1824.
Reference: Pax & Hoffm., Pflanzenreich IV. 147. vi: 78. 1912.
Usually more or less shrubby, sometimes perennial herbs or annual
but always somewhat pubescent with malpighiaceous trichomes or
these only in part simple. Leaves alternate, often subsessile, oblong
or ovate-lanceolate, entire or serrate, 3-nerved from base. Racemes
ordinarily bisexual, often congested, basally female, above male, the
bracts 1-flowered, the short pedicels sometimes reflexed in fruit.
Male calyx at anthesis valvately 4-5-parted, the 4-5 petals entire,
shorter or longer and more or less adnate as the 5 disk glands
to the column, the stamens often 10, 1-verticillate (type, Wheeler)
or 2-3-verticillate on the elongate column (and with or without
staminodia) or this short and stamens only 4 or 6. Rudimentary
ovary none or obsolete. Female sepals 5 (6), as petals these some-
times reduced or none. Disk glands free, rarely filamentous. Ovary
3-celled, styles basally connate or free, often bifid, the branches
sometimes lobed. Capsules dehiscing into 3 2-valved cocci, the
column persisting, the ecarunculate globose seeds often reticulate.
Pax & Hoffmann I.e., 51, maintained Ditaxis while Croizat, Journ.
Arnold Arb. 26: 191. 1945, followed Bentham, in Benth. & Hook,
f. Gen. PI. 3: 303. 1880, in including it in the group of Browne which
as to type has univerticillate stamens. The name has been spelled
variously as Argithamnia by Swartz this adopted by Pax & Hoff-
mann Argyrothamnia by Mueller and Argothamnia by Sprengel;
however, Wheeler, Contr. Gray Herb. 124: 40. 1939, showed that
the name of P. Browne, 1756, is the earliest form.
Leaves oblong-ovate or elliptic, about 1.5 cm. wide; male sepals
4-5 mm. long; glands obsolete A. dioica.
Leaves obovate, about 2 cm. wide; male sepals 8 mm. long; glands
developed, adnate A. Katherinae.
Argythamnia dioica (HBK.) M. Arg. Linnaea 34: 145. 1865;
56. Ditaxis dioica HBK. Nov. Gen. & Sp. 7: 170. pi. 639. 1825.
FLORA OF PERU 93
Much-branched shrub, the lightly angled branchlets sericeous
pubescent as the entire plant except the leaves above including the
sepals and petals both sides (except in male flowers subglabrous)
and ovary, the indument intricate, especially on the leaves beneath;
petioles 1-3 mm. long; stipules triangular, acuminate, coriaceous;
leaves oblong-ovate or elliptic, obtuse or subacute, obtuse at base,
2.5-5 cm. long, 13-17 mm. wide, minutely glandular-serrulate, finally
somewhat glabrescent above, subcoriaceous, the 3 basal nerves
elongate; male racemes few-, female 1-2-flowered, the pedicels much
shorter than calyx, scarcely accrescent in fruit; male sepals lanceo-
late-acuminate, 4-5 mm. long, the longer petals high adnate on
column, spathulate-lanceolate, subobtuse; disk glands not at all
evolute; stamens 10, biverticillate, staminodia papillose-hirsute;
female sepals subcordate-ovate, acuminate, 5-7 mm. long, the lower
margins overlapping, the petals little longer; disk glands confluent,
glabrous; styles pilose, medially connate, free parts biparted, the
lobes dilated and sometimes lobulate; capsules unknown. Described
as dioecious, an observation which Pax and Hoffmann rightly ques-
tion, since related species are all monoecious.
Cajamarca: On the shore of the Amazon at Jae*n de Bracamoros,
Humboldt 3604, type; Weberbauer 6222.
Argythamnia Katherinae (Pax) Macbr., comb. nov. Ditaxis
Katharinae Pax, Pflanzenreich IV. 147. vi: 56. 1912.
A 2-meter shrub, more or less appressed pilose including the petals
and female sepals both sides and ovary, the apically foliate flowering
branchlets and obovate leaves finally more or less glabrescent;
stipules acuminate, 3 mm. long; petioles 1-4 mm. long; leaves rounded
at tip, minutely apiculate, membranous, 3-5.5 cm. long, 15-27 mm.
wide; racemes either with a solitary female flower and 5-8 male or
entirely unisexual; bracts about 4 mm. long; flowers white, with the
leaves developing or precocious, the sepals of the male narrowly
lanceolate, acuminate, glabrous within, 8-9 mm. long, the slightly
longer petals with medial nerve thickened below; disk glands at-
tenuate both ends, completely adnate to column, this as filaments
and 5 staminodia puberulent; stamens 10, biverticillate; female
sepals ovate, cordate at base, 10-13 mm. long, the lower margins
recurved, accrescent after an thesis; female petal with midnerve
less enlarged, 7-12 mm. long; disk glands free, glabrous; styles
shortly bilobed, the thick entire lobes glabrous. Named for Kathe
Hoffmann. D. macrantha Pax & Hoffm., I.e. vii: 426. 1914, of
94 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Ecuador, has very thin glabrescent leaves 5.5-9 cm. long, 2.5-4.5
cm. wide, the female petals only 8-9 mm. long, disk annulate.
Illustrated, Pax, I.e. p. 57.
Cajamarca(?): Los Reyes, route Chepen to Cajamarca, 1,000
meters, stony outcrops, Weberbauer 4809; 4810, types; 188 (under
an unpublished name).
11. CHIROPETALUMJuss.
Reference: Pax & Hoffm., Pflanzenreich IV. 147. vi: 86. 1912.
Similar to Argythamnia but the petals of the male flowers palm-
ately 3-7-parted with narrow lobes. Disk glands free. Stamens 5
or rarely 6. Seeds rugose-asperous. Indument on upper parts
often simple and stellate trichomes intermixed. The genus could
readily be included in Argythamnia as by Baillon, M. Arg., and
Bentham, but apparently there is no species intermediate in the
highly distinctive character of the much-divided petals of the male
flowers.
Indument of the leaves mostly or entirely stellate . . . . C. Pavonianum.
Indument of the leaves malpighiaceous and simple mixed.
Disk glands glabrous; leaves plinerved C. quinquecuspidatum.
Disk glands (male) pilose; leaves basally 3-5-nerved.
C. Ruizianum.
Chiropetalum Pavonianum (M. Arg.) Pax, Pflanzenfam. 3,
Abt. 5: 45. 1890; 89. Argyrothamnia Pavoniana M. Arg. Linnaea
34: 149. 1865.
A small shrub with virgate branchlets becoming glabrate below;
petioles short; leaves ovate, acuminate, shortly contracted at base,
2-3.5 cm. long, 1-2 cm. wide, stellate-pilose, 5-nerved, the longis
ascending secondary nerves not incurved toward the serrate margins;
racemes few, slender, laxly flowered; bracts ovate, acute, not at all
comosely exserted in the younger racemes; male pedicels shortt
than the calyx; female sepals obovate-lanceolate, shortly acuminate,
the male to about 1.5 mm. long, the nearly as long petals deeply
5-parted, the disk glands hirsute pubescent, those of female flower
puberulent apically; ovary villosulous; styles slenderly biparted;
capsules 3 mm. thick, depressed, puberulent, the seeds asperous.-
Perhaps should be drawn to include C. molle [Klotzsch] Klotzscl
FLORA OF PERU 95
ex Pax & Hoffm., 87, of southern Brazil, with twice as large leaves,
a third larger flowers, glabrous glands.
Peru(?): Middle high Andes, Pavdn, type.
Chiropetalum quinquecuspidatum (Juss.) Pax & Hoffm.
Pflanzenreich IV. 147. vi: 92. 1912. Croton quinquecuspidatum Juss.
Euphorb. Tent. 110. pi. 8, fig. 26c. 1824. Chiropetalum peruvianum
Juss. Ann. Sci. Nat. se*r. 1. 25: 22. 1832. Argyrothamnia quinque-
cuspidata (Juss.) M. Arg. Linnaea 34: 150. 1865.
Petioles a few mm. long; leaves ovate or lanceolate-ovate or the
uppermost lanceolate, acuminate, contracted at base, dentate-serrate,
5-plinerved, the long-ascending secondary nerves subsericeous;
racemes exceeding the leaves, the lower half naked, slender, laxly
flowered; pedicels very short; male petals cuspidately 5-7-lobed,
violet-purplish; female sepals triangular-ovate, acuminate; petals
linear, entire; ovary pubescent; styles biparted nearly to base. As
suggested, could be a variant of the Chilean and Argentinian C.
tricuspidatum (Lam.) Juss., 92, with 3-parted male petals and lanceo-
late female sepals.
Peru(?) : Without locality, Dombey, type.
Chiropetalum Ruizianum (M. Arg.) Pax & Hoffm. Pflanzen-
reich IV. 147. vi: 94. 1912. Argyrothamnia Ruiziana M. Arg. Linnaea
34: 151. 1865.
Shrub about 2 meters high with striate-angulate branchlets, the
younger fulvous pubescent; petioles 2-4 mm. long; stipules 2 mm.
long, lanceolate-acuminate; leaves orbicular-ovate or ovate, cuspi-
dately acuminate, contracted or subcordate at base, 4-5 cm. long,
2-4 cm. wide, serrate-denticulate, membranous, softly pilose both
sides with malpighiaceous and simple trichomes, mostly simple
especially beneath, basally 3-5-nerved; racemes slender, many-
flowered, 4-7 cm. long, bracts 1.5 mm. long, pedicels about 2 mm.
long, to 3 mm. in fruit, the 1-4 female flowers with densely pilose
sepals finally 4 mm. long, minute linear petals, broad truncate disk
glands, the acuminate male sepals 2 mm. long, the two-thirds as
long petals 5-7-parted, globose disk glands pilose; stamens 5; ovary
densely pubescent with malpighiaceous trichomes; styles medially
bifid; capsules 5 mm. thick, depressed, pilose, the seeds 2 mm. in
diameter. C. canescens Phil. (C. sponiella (M. Arg.) Pax, 93),
Chilean and Bolivian but accredited to Peru by Pax (wrongly) has
leaves 2-3 cm. long, glabrous disk glands; the more similar C.
96 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
boliviense (M. Arg.) Pax & Hoffm., 94, apparently has only the leaf-
nerves villous pubescent. F.M. Neg. 7121.
Lima: Cheuchen, Ruiz & Pavdn, type. Matucana, among large
rocks, 348 (det. Johnston). Junin: near Tarma, Ruiz & Pavdn.
Hudnuco: Slender shrub along trail, Yanano, 3710 (det. Johnston).
Near Chaglla, 2,800 meters, Weberbauer 6695 (det. Pax). Ayacucho:
Huanta, in bushes above Osno on the Rio Apurimac, Weberbauer
5594 (det. Pax).
12. CAPERONIA St. Hil.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vi: 27. 1912.
Annual or perennial usually setose-hispid and glandular herbs
with alternate acutely serrate ordinarily narrow or oblong-ovate
leaves and axillary peduncled racemes or spikes of generally monoe-
cious petaliferous flowers solitary in the scattered bracts, the more
numerous male above, the few female below, the 5 imbricate calyx
lobes of these often enlarged in fruit. Disk none. Male flowers
with 5 valvate sepals, 5 imbricate often unequal petals. Stamens
usually 10, biseriate; anthers ovoid. Rudimentary ovary on tip of
the cylindric stamen-column entire or minutely tridenticulate. Petals
of female flowers narrower or reduced. Ovary sessile, 3-celled;
styles short, palmately dissected. Capsules tridymous, hispid or
echinate separating into bivalved cocci, the subglobose seeds ecarun-
culate. Commemorates pre-Linnaean pharmacist Caperoni, who
first called these plants to the attention of botanists.
Male petals very unequal; leaves obscurely serrulate. . .C. zaponzeta.
Male petals alike; leaves closely serrate.
Younger stems appressed pubescent, the older glabrate and often
fistulose C. castaneifolia.
Younger stems with more or less spreading often glandular
pubescence, all usually obscurely or not fistulose. C. palustris.
Caperonia castaneifolia (L.) St. Hil. Hist. PI. Remarq. Bre*sil
245. 1824; 31. Croton castaneifolius L. Sp. PI. 1004. 1753.
Much like C. palustris but usually more erect and the older stems
septate and fistulose, glabrous or nearly, a cm. or more in diameter;
stipules broadly ovate; petioles to 4.5 cm. long; leaves to 16 cm.
long, 3-8 mm. wide; male sepals 2 mm. long; capsules 7-8 mm.
thick, 4 mm. long, 3-sulcate. Illustrated, Pax & Hoffm. I.e. p. 32.
FLORA OF PERU 97
Cajamarca: Tomependa, Ja&i de Bracamoros, (Bonpland).
Mexico; West Indies; South America.
Caperonia palustris (L.) St. Hil. Hist. PL Remarq. Bre"sil
245. 1824; 33. Croton palustris L. Sp. PI. 1004. 1753.
Annual often spreading and attaining several dm., the terete
striate unarmed stems (at least above), petioles, these 5-25 mm.
long, and peduncles setulose with glandular spreading rigid trichomes;
stipules lanceolate, acuminate, 3-5 mm. long; leaves variable in
form, the smaller lower elliptic-obovate and obtuse, the upper ovate
to lanceolate, even linear-lanceolate, usually 7-12 cm. long, 1-5 cm.
wide, all membranous, deeply and somewhat irregularly serrate, the
larger with 11-26 straight secondary nerves; racemes spiciform, the
lower 3-5 flowers female with unequal obovate sepals becoming in
fruit about 5 mm. long, setulose, spathulate-lanceolate obtuse or
acute petals about half as long, densely glandular ovary, the short
laciniate styles nearly concealed in the glands; male sepals not accres-
cent, lanceolate, acute, 1-1.5 mm. long, the obtuse petals scarcely
longer; stamens 10; ovary rudiment cylindric, apically denticulate;
capsules 6-7 mm. thick, 5 mm. long, muricate, the ovoid foveolate
seeds 2-3 mm. long. Illustrated, Pax, I.e. fig. 1, p. 4, and fig. 6, p. 32;
O'Donell & Lourteig, Lilloa 8: p. 58.
San Martin: Tarapoto, Spruce 2473. Without locality, Ruiz &
Pavdn (det. Mansfeld). Mexico and West Indies to Argentina;
Africa; Madagascar.
Caperonia zaponzeta Mansfeld, Notizbl. Bot. Gart. Berlin 9:
265. 1925.
Stems lightly puberulent-pilose and aculeate, hollow and about
6 mm. thick in flowering portions; stipules 2.5 mm. long; petioles
pilose, 3 mm. long; leaves elliptic or lanceolate-elliptic, rounded at
base, acute to acuminate, 10-18 cm. long, 3.5-5 (7) cm. wide, gla-
brate or early sparsely appressed pilose both sides, chartaceous, the
remote, dentations about 0.5 mm. long, racemes axillary to 13 cm.
long including the half as long peduncle, basal female flowers 3,
the apical male crowded on the pilose rachis, the former with broadly
ovate abruptly short-acuminate sepals, these like the spathulate
petals 3 mm. long and pilose both sides; ovary densely glandular,
styles obovate-cuneate, lacinulate; male sepals ovate, acute, 3 mm.
long, sparsely pilose, the glabrous petals unequal, the larger ovate,
obtuse, 2 mm. long, smaller spathulate, 0.7 mm. long; stamens 10,
98 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
biverticillate; rudimentary clavate ovary 3-denticulate. The fruits
are densely and rather rigidly echinate, about 5 mm. broad. Near
C. heteropetala but that with retuse male petals, glandular lanceolate
female sepals, more dentate leaves (Mansfeld). The common name
according to the collector means "sea cow teeth." F.M. Neg. 5224.
Loreto: Overhanging shore, shrub-like herb, Yarina Cocha,
Tessmann 3386, type. La Victoria, Williams 3124 (det. Mansfeld);
2823. Puerto Arturo, Yurimaguas, Williams 5197 (det. Standley).
Maucallacta, Rio Paranapura, King 3954 (det. Standley). "Zapon-
zeta."
13. DALEGHAMPIA [Plum.] L.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. xii: 1-56. 1919.
Shrubs, often scandent, the pubescence if any simple and some-
times stinging. Leaves undivided or 3-5-lobed or -foliate, bistipulate
and the petioles usually distinctly bistipulate. Inflorescences with
2 often large and colored bracts, these bistipulate and serving as an
involucre to a lower female 3-flowered 3-bracted dichasium and an
upper 4-bracted male pleiochasium, its bracts often urceolate-con-
nate, and composed of 3-flowered male cymes, 1-flowered female
cyme and sterile flowers. Male calyx 4-6-parted; stamens many,
filaments short. Female calyx with 5-12 imbricate often pinnatifid
indurating sepals. Ovary 3(4)-celled; styles connate, the long
column apically obtuse or dilated. Bivalved cocci on persisting
column; seeds globose, ecarunculate. The genus commemorates
Jacques Dalechamps, early sixteenth century botanist of Lyon.
Leaves undivided.
Female sepals undivided.
Plants scandent; leaves rounded at base D. micrantha.
Plants erect; leaves narrowed to base.
D. brevipedunculata, D. spathulata.
Female sepals pinnatifid.
Style column spreading-capitate, dilated above (typically);
bracts at least 2 cm. long.
Leaves usually glabrate in age; bracts 2-3 cm. long.
D. dioscoreaefolia.
Leaves tomentose beneath; bracts 4-10 cm. long.
D. aristolochiaefolia.
Style column not so dilated apically; bracts 15-18 mm. long.
D. hispida.
FLORA OF PERU 99
Leaves at least in part lobed or trifoliate; female sepals pinnate.
Leaves with 3 leaflets.
Sepal segments (female flowers) early capitate-glandular; leaflets
thin, 3-5 cm. wide D. cissifolia.
Sepal segments not capitate-glandular; leaflets firm, narrow.
D. Weberbaueri.
Leaves more or less deeply lobed, at least in part.
Stipules conspicuously glandular-setose D. stipulacea.
Stipules not glandular-setose.
Involucral bracts 1.5-4 cm. long, 5-nerved.
Bracts 4 cm. long; sepals in fruit about 3 cm. long.
D. Ukana.
Bracts 1.5-3 cm. long; sepals in fruit about 1 cm. long.
D. scandens.
Involucral bracts 3-6.5 cm. long, 7-9-costate . . . D. tiliaefolia.
Dalechampia aristolochiaefolia HBK. Nov. Gen. & Sp. 2:
99. 1817; 47.
Liana, the younger branchlets puberulent-tomentulose and
sparsely hirsute, the 5-10 mm. long petioles densely so; stipules to
9 mm. long; leaves ovate-lanceolate, shortly acuminate, basally
cordate, 7-11 cm. long, 3-7 cm. wide, palmately nerved, minutely
denticulate, linearly bistipulate, pubescent above, densely tomentose
beneath; inflorescences terminal, often on leafy or leafless branchlets;
bracts suborbicular, 4-7 (10) cm. long, narrowed to obtuse base, in-
cised dentate, sometimes rather trilobate, pilose on the 5-7 nerves,
rose or reddish-violet, in fruit to 10 cm. long; stipules 1-2 cm. long,
lanceolate, often unequal; female bracts reniform-ovate, obtuse, lateral
much smaller, male glabrous, lustrous as the male calyx, the stamens
about 50; female sepals around 10, with 5-6 linear lacinulae, eglandu-
lar, after an thesis 5 mm. long; ovary villous; column 3-4 mm. long,
finally twice as long, often dilated to pileiform stigma; capsules
12 mm. broad, cocci carinate; seeds 5 mm. long, muriculate, varie-
gated. Pax and Hoffmann observed in some specimens style not
pileiform-dilated. Cultivated for adornment (Herrera) and well
worth it for the showy bracts. Illustrated, Pax & Hoffm. I.e. 47.
Cajamarca: San Felipe, Bracamoros, Humboldt, type. Shrub
land, Socota, Stork & Horton 1 0093 (det. Standley) . Below Rambrdn,
2,000 meters, Weberbauer 4801 . Huanuco : Yanano, 3802 (det.
Johnston). Llamapanahui, Ruiz & Pavdn (det. Mansfeld). Valley
100 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
of the Maranon, Weberbauer 4256; 191. Amazonas: Between Balsas
and Celedin, Stuebel 45d. Hacienda Limon, Osgood & Anderson 59.
Apurimac: Chaupimayo, Soukup 812. Pachachaca Valley, Goodspeed
Exped. 10563. Cuzco: Valle de Apurimac, Hacienda Marcahuasi,
2,450 meters (Vargas). San Miguel, Urubamba Valley, Cook &
Gilbert 1022 (det. Standley). "Bella abanquina" (Vargas), "etepen
ccaccha" (Cook & Gilbert).
Dalechampia brevipedunculata Ule, Verb. Bot. Ver. Brand-
enb.50:82.1908.
Like D. spathulata but peduncles only 2-3 cm. long, female sepals
3 mm. long; petioles 3-4 mm. long; leaves 10-12 cm. long, 3.5-4.5
cm. wide; style column 4 mm. long; capsules 10 mm. long, 7 mm.
broad; seeds 4-5 mm. long, aculeolate. Probably will prove to be
a variant of the similar species. F.M. Neg. 5332.
San Martin: Tarapoto, Ule 6504- Loreto: Yarina Cocha, Tess-
mann 5408 (det. Mansfeld). Brazil.
Dalechampia cissifolia Poepp. in Poepp. & Endl. Nov. Gen.
& Sp. 3: 20. 1845; 19. D. triphylla Lam. var. cissifolia (Poepp.)
M. Arg. in DC. Prodr. 15, pt. 2: 1239. 1866.
Climbing, the branchlets and petioles, these 4-9 cm. long, at
least early hispid as also the leaves on both sides, the latter composed
of 3 sessile or petiolulate thin-membranous leaflets, 8-10 cm. long,
3-5 cm. wide, the intermediate oblong-obovate, the lateral obliquely
ovate, all subdenticulate, reticulate-veined; stipules 8, stipels 3-4
mm. long; peduncles 1-1.5 cm. long or in fruit as much as 4 cm.;
involucral bracts orbicular, 3-lobed, denticulate, 2-2.5 cm. long (or
apparently shorter), the stipules lanceolate, 5 mm. long; intermediate
female bracts reniform-ovate, lateral connate into one, all ciliate
and marginally glandular, the 10-12 sepals pinnately parted with
elongate filiform divisions, after anthesis 15 mm. long, the laciniae
3-4 mm. long; ovary puberulent; style column 5 mm. long, slightly
dilated at tip; capsule subglabrous, 8 mm. thick, the pedicel 1 cm.
long; seeds 3 mm. in diameter, black. Not at all closely allied to
D. triphylla of southern Brazil (Pax & Hoffm.) the coarser laciniae
of the female sepals not capitate-glandular as in the Peruvian plant.
However, the constancy of the character is certainly often obscure
and accordingly its taxonomic importance remains to be proved by
the study of more collections. Compare also D. juruana Ule under
D. Weberbaueri; from the material seen it is not clear that there is
FLORA OF PERU 101
specific significance as there is no concomitance between size of
bracts, peduncle length and sepal glandulosity. The San Martin
material with smaller bracts could be D. Weberbaueri. F.M. Neg.
5335.
San Martin: Lomas, Williams 6328. Juanjui, Klug 4208 (det.
Standley). San Roque, Williams 7492. Loreto: Yurimaguas,
Poeppig 2085, type. Iquitos, Williams 8051. Mishuyacu, Klug
523. Balsapuerto, Klug 3102 (det. Standley, but short peduncles
of D. juruana!}. La Victoria, Williams 2888. Letecia, Wittiams 3159.
Dalechampia dioscoreaefolia Poepp. in Poepp. & Endl. Nov.
Gen. &Sp. 3:20. 1845; 46.
Liana with slender sulcate-striate more or less pubescent branch-
lets, petioles (2-8 cm. long) and leaves, all often glabrescent in age;
leaves ovate, shortly contracted into the openly cordate acute base,
5.5-16 cm. long, 4-11 cm. wide, denticulate or subentire, palmately
nerved, membranous; stipels filiform, stipules to 6 mm. long;
peduncles solitary or geminate, 1-6 cm. long; bracts ovate, acuminate,
contracted to petiole, incised-dentate, 2-3 cm. long, 1.5-2 cm. wide,
pink, pilose, 5-7-nerved, deciduous; stipules persisting, often unequal,
ovate, reflexed, to 7 mm. long; flowering bracts glabrous, erose,
the female concave, subsaccate; male calyx pilose, stamens about 15;
female sepals 6-12, with 4-6 laciniae each side of the linear rachis,
long pilose, not capitate, indurating, 7-10 mm. long; intersepalous
glands small; ovary sericeous, style column finally 8 mm. long, dilated
into piliform stigma; capsules 12 mm. broad, 7 mm. long, stipe 2 cm.
long; seeds 5 mm. long, irregularly costulate and rugulose, pale-
mottled. The var. pubescens M. Arg. has the leaves pubescent
beneath, fruiting sepals only 7 mm. long. The Amazonian D. affinis
M. Arg. has linear involucral stipules and laciniae of the female
sepals capitate-glandular. F.M. Neg. 5338.
San Martin: Juanjui, Klug 4186 (det. Standley). Tocache,
Poeppig 1907, type, and 2163; Williams 4228. Tarapoto, Spruce
4476 (var.). Chazuta, Klug 4013. Without locality, Ruiz & Pavdn.
Junin: La Merced, 5384; Soukup 2457. Loreto: Lower Rio
Nanay, Williams 303. Rio Itaya, Williams 32. Iquitos, Williams
8186; Klug 550; 753; 1037.
Dalechampia hispida Poepp. in Poepp. & Endl. Nov. Gen. &
Sp. 3: 19. 1845; 48.
Hispid-hirsute liana, the pubescence extending even to the female
sepals and ovary with the exception of the bracts; petioles 7-7.5 cm.
102 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
long; leaves ovate, openly cordate, acuminate, 14 cm. long, 9 cm.
wide, palmately nerved, membranous, denticulate; stipules 5 mm.
long, stipels filiform-linear; peduncles 1-1.5 cm. long; bracts green,
18 mm. long and little wider, truncate-cordate, shortly 3-lobed,
denticulate, subglabrous; female bracts subequal, broadly reniform-
ovate, obtuse, denticulate, glandular-ciliate, the sepals 10, pinnate,
with 4-5 filiform laciniae, finally 15 mm. long, rachis filiform, indu-
rate, style column slender, slightly enlarged at tip; capsules 5 mm.
long, twice as broad, glabrate, the lustrous mottled seeds smooth.
F.M. Neg. 32441.
San Martin: Tocache, Poeppig 2016, type.
Dalechampia micrantha Poepp. in Poepp. & Endl. Nov. Gen.
& Sp. 3: 19. pi. 222. 1845; 7. Rhopalostylis buettnerioides Klotzsch
ex Baill. Adansonia 5: 317. 1864-5.
High-climbing liana with undivided basally 5-nerved leaves;
petioles 2.5-4.5 cm. long, puberulent as the peduncles and male
calyces, these clavate in bud with about 13 stamens; leaves ovate,
obtusely acuminate, rounded-truncate at base, obscurely denticulate,
9-14 cm. long, 5-13 cm. wide, short-pubescent on nerves; caducous
stipules 3 mm. long, lanceolate, the stipels narrower; peduncles axil-
lary, usually branched at base, to 2 cm. long; involucral bracts 2.5-4
mm. long, broadly lanceolate from base, acuminate, shorter and
narrower than the ovate stipules; female bracts broadly ovate,
acute, subtridentate, the male obovate, subtruncate, denticulate;
flowers lutescent, the 5-6 lanceolate ovate acute entire sepals glandu-
lar at base; ovary pubescent; style column to 9 mm. long, clavate,
glabrous; capsules 13 mm. thick, ligneous, the cocci carinate; seeds
5 mm. across, smooth. Illustrated, Pax & Hoffm. I.e. 8. F.M.
Neg. 7200.
Loreto: Near Iquitos, Vie 625 4; 6255; Williams 3629 (det.
Mansfeld). To the Guianas.
Dalechampia scandens L. Sp. PI. 1054. 1753; 32.
Slender-stemmed vine, the branches often early more or les
pubescent as the leaves beneath, the indument commonly puberulent
or short-pilose, not rarely sparse or sub-lacking especially in age;
petioles 2-many cm. long; leaves 3-5-parted medially or usually
more deeply, sometimes nearly to base, said to range 2.5-14 cm. loi
and at least as wide, lateral lobes oblique, all shortly acuminate or
acute or if obtuse, mucronate; stipules reflexing, 2-10 mm. long;
FLORA OF PERU 103
stipels subulate, 1-3 mm. long; peduncles 1.5-6 cm. long; involucral
bracts pale, 5-nerved, 1.5-3 cm. long and quite as wide, broadly
ovate, cordate at base, 3-lobed often medially, typically glandular-
ciliate and more or less hirsutulous; male calyx glabrous or puberu-
lent, stamens 20-27 (rarely 4-5), female sepals 7-10, rachis ordinarily
lanceolate with 3-7 pairs of laciniae, capitate glandular and setose,
5-9 (12) mm. long; ovary pubescent; style-column often dilated
apically; capsules 7-10 mm. broad; seeds globose, 2.5-4 mm. in
diameter, smooth, whitish or brown mottled. The monographers
discriminated twelve variants which, at least as they have defined
them, do not seem to have taxonomic importance.
Tumbez: Mountains east of Hacienda Chicama in deciduous
bushwood, Weberbauer 7650; 7667. Piura: Parinas Valley, Haught
276. Junin: In brush along trail, La Merced, 5284 (det. Johnston).
Cuzco: Quillabamba, Goodspeed Exped. 104-4-8 (det. Standley).
Generally distributed in warmer regions.
Dalechampia spathulata (Scheidw.) Baill. Etude Gen. Eu-
phorb. 487. 1858; 9. Cremophylla spathulatum Scheidw. Bull. Acad.
Brux. 9, pt. 1: 23. 1842. D. amazonica (Ule) Macbr. Candollea 8:
26. 1940.
Shrub, glabrous unless the young parts, puberulent petioles,
these 2-10 mm. long, leaf-nerves beneath, peduncles, these 2-6 mm.
long, and bracts (sometimes hirsute), ovate or orbicular, acute or
acuminate, 2.5-4 cm. long, 2-4 cm. wide, more or less dentate,
3-nerved, often roseate or lutescent; stipules 3-8 mm. long; leaves
obovate-spathulate, long-decurrent at base, typically subabruptly
acuminate, to 2.5 dm. long, 1 dm. wide; female bracts elliptic,
lateral linear-lanceolate, subulate-acuminate; male broadly ovate;
male sepals puberulent; stamens 15; female sepals 6, linear-lanceo-
late, 1 mm. long, alternating with 1-2 laciniae; ovary pubes-
cent; style column slender, 5-10 mm. long; capsules 5.5 mm. long,
9 mm. broad; seeds globose, softly muriculate. Has been in cultiva-
tion. The Peruvian form is var. amazonica Ule, the bracts hirsute
all over the inner surface. The related species are D. magnoliifolia
M. Arg., 11, the elliptic-lanceolate leaves narrowed both ends, and
D. brevipedunculata Ule, 11, apparently much like D. spathulata but
peduncles only 2-3 mm. long, female sepals 3 mm. long; the former
could be represented by Killip & Smith 28920 but is doubtfully
specifically distinct. Illustrated, Pax & Hoffm. I.e. 10 ex Pflanzen-
familien 3, Abt. 5: 68.
104 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
San Martin: Tarapoto, Ule 6505 (type, var.). Loreto: Yuri-
maguas, Poeppig 2380; Williams 4-189. Santa Rosa, Kittip & Smith
28920. Amazonian Brazil to Central America.
Dalechampia stipulacea M. Arg. Linnaea 34: 221. 1865; 27.
More or less hirsute or villous pubescent liana with conspicuous
stipules 2-3 cm. long, variously glandular-setulose and with dissected
stipules within; petioles 4-9 cm. long; leaves about medially 3-lobed,
deeply and narrowly cordate at base, the ovate or oblong lobes acute
or acuminate, firm-membranous, usually stipitate glandular-ciliate;
peduncles axillary and terminal, 3-4 in fruit 5-6 cm. long; involucral
bracts 3-4 cm. long or larger, greenish-white, rotund, lobate, glandu-
lar-fimbriate and hirsute; female bracts ovate, obtuse, male eglandu-
lose; male flowers 4 mm. wide; stamens 55; female sepals 10-12,
10-18 mm. long, pinnate with many laciniae, these apically capitate-
glandular; ovary tomentose; style column lightly dilated at tip;
capsules 12 mm. thick; seeds smooth, 4 mm. in diameter, white and
brown mottled. The Peruvian plant, very probably a distinct
species (Pax & Hoffmann) is var. major M. Arg. in Mart. Fl. Bras.
11, pt. 2: 656. 1874, the leaves villous both sides with long trichomes,
bracts 5-6 cm. long, 7-nerved; fruiting sepals to 18 mm. long, long
white-setulose; nearly all parts more robust. F.M. Negs. 5355;
29352; 34053.
Junin: Vitoc, Ruiz & Pav6n, type. Brazil; Venezuela; Colombia;
Paraguay; Argentina.
Dalechampia tiliaefolia Lam. Encycl. 2: 257. 1786; 37. D.
peruviana Lam. I.e.
Liana, all the younger parts more or less puberulent-tomentulose,
the leaves becoming glabrescent above; petioles 2.5-8 cm. long;
leaves ordinarily diverse on the same stem or branch, but sometimes
all undivided or 3-lobed, the former usually 7-14 cm. long, 4-12 cm.
wide, cordate, broadly ovate, the latter 7-17 cm. long, nearly as
wide, the middle lobe oblong-elliptic, the lateral very inequilateral,
all denticulate; stipules 3-5 mm. long, subulate, stipels 2-4, 2 mm.
long; inflorescences terminal on few-leaved branchlets, the yellowish-
white bracts 3-6.5 cm. wide and long, merely shortly 3-dentate,
prominently 7-9-nerved, the stipules 2.5 mm. long; female bracts
truncate, often yellowish, 3-lobed, softly pilose-puberulent, the male
ovate; calyx glabrous; stamens about 40; female sepals 12-13 pinnate
with about 13 laciniae, hispid, non-capitate-glandular, after anthesis
FLORA OF PERU 105
attaining 15 mm., plumose-hispid; ovary hispid; style column elon-
gate, campanulate-umbraculiform, 3-lobate; capsule 11 mm. broad,
verrucose; seeds 5 mm. diameter, greenish-gray, brown mottled.
The tropically widely distributed D. scandens L. has regularly 3-5-
lobed leaves, shorter fruiting sepals, smaller seeds. Except as noted
determinations by Standley. Illustrated, Pax & Hoffm. I.e. 38.
Junin: La Merced, sunny riverside brush, 5543 (det. Johnston);
Killip & Smith 23824- Chanchamayo Valley, Schunke 378. Colonia
Perene*, Killip & Smith 24999; 25060. San Martin: Juanjui, clear-
ing, Klug 4331. Loreto: Yurimaguas, Poeppig 1886. Without
locality, "Peru," Jos. Jussieu, type. Rio Acre: Ule 9542. To
Central America, Trinidad and southern Brazil.
Dalechampia Uleana Pax & Hoffm. Pflanzenreich IV. 147.
xii: 31. 1919.
Liana, the branchlets, petioles (7-12 cm. long), 5 leaf- and bract-
nerves more or less ashy puberulent; leaves deeply and openly cordate
at base, 8-16 cm. long, 11-20 cm. wide, chartaceous, reticulate-
veined, more than medially 3-lobed, the middle lobe obovate,
abruptly acuminate, the spreading lateral very inequilateral, all
repand-denticulate; stipules ovate, 7 mm. long, 5 mm. wide, stipels
filiform, 4 mm. long; axillary peduncles 5-7 cm. long; involucral
bracts similar to leaves, greenish-purple, 4 cm. long, 5 cm. wide;
stipules 1.5-2 cm. long, elliptic; female bracts rotund, lateral connate,
sparsely pilose, male suborbicular, free, the calyx pubescent and with
80-90 stamens; female sepals about 10, pinnate with about 27
pubescent non-capitate-glandular laciniae, after anthesis to 3.5 cm.
long loosely enclosing the capsule, this 16 mm. long and broad, verru-
culose, the pedicels 2-4 cm. long. Illustrated, Pax & Hoffm. I.e.
38. F.M. Neg. 24380.
Rio Acre: In woods at Nueva York, Ule 9541, type. Brazil.
Dalechampia Weberbaueri Pax & Hoffm. Pflanzenreich IV.
147. xii: 21. 1919.
Liana with slender striate villous stems, the ultimate filiform;
petioles 1-4 cm. long, pubescent as the branches; leaflets 3, 2-5 cm.
long, 5-13 mm. wide, middle lanceolate, lateral inequilateral, oblong-
lanceolate, base semi-cordate, acute, denticulate, firm-membranous,
pubescent both sides; stipules linear, 4-5 mm. long, stipels subulate,
2 mm. long; peduncle axillary or terminal on leafy lateral branch-
lets, slender, 1.5-3 cm. long; involucral bracts broadly ovate, cordate
106 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
at base, about 14 mm. long, less than medially 3-lobed, denticulate,
green; stipules 2-3 mm. long; intermediate female bracts scarcely
1 mm. long, truncate, the male connate; male calyx pubescent;
stamens 6-7; female sepals about 12, pinnate, 1-laciniate below each
side, apically with 2-3 linear laciniae, pilose, fruiting 5-6 mm.
long; ovary tomentose; style column 3-lobed; capsules 6 mm. thick;
seeds 2.5 mm. in diameter, black and mottled. Perhaps a variety
of the earlier D. juruana Ule, 21, which as to type had stouter
peduncles about 1 cm. long, entire leaflets. Apparently both plants
too near D. triphylla Lam. but the involucral bracts smaller and in
type of D. juruana, one trifid, others entire.
Cajamarca: In rain-green shrub formation, 800 meters, Ja&i,
Weberbauer 6192, type.
14. PERAMutis
Reference: Pax & Hoffm., Pflanzenreich IV. 147. xiii: 1-13.
1919.
Tall shrubs or sometimes trees, the indument often stellate-
lepidote or in Peru sometimes simple, the leaves usually alternate,
always shortly petioled, entire, pinnately nerved, the ordinarily
dioecious sessile flowers subtended by an involucre of 1 or 2 bracts,
sometimes the outer flowers female, the central male but commonly
the involucres unisexual, with or without floral rudiments. Disk
none. Male calyx wanting or valvate, turbinate, stamens 2-5, the
filaments sometimes partly connate; ovary rudiment none. Female
flowers naked; ovary 3-celled, ovules solitary. Capsules composed
of 3 bivalved carpels, the column persisting; seeds carunculate,
black, lustrous, smooth.
Leaves distichous; pubescence simple P. distichophylla.
Leaves opposite or alternate; pubescence lepidote P. benensis.
Pera benensis Rusby, Descr. N. Sp. S. Amer. PL 49. 1920.
P. Mildbraediana Mansf. Notizbl. Bot. Gart. Berlin 9: 265. 1925.
Tree, the branchlets lustrous, the leaves above glabrous, the latter
paler beneath with dense lepidote indument, opposite, oblong-
elliptic or oval, rounded to broadly acute at base, slightly apiculate
at the obtuse apex, 6-16 cm. long, 3-7 cm. wide; petioles 1-2.5 cm.
long, canaliculate above; venation slender; involucres fasciculate at
the often defoliate nodes, lepidote, 4 mm. in diameter, peduncles
FLORA OF PERU 107
5 mm. long; rudimentary female flowers with two unequal mostly
2-3-lobed bracts; male calyx wanting, stamens 12-14, filaments
1 mm. long, anthers 1.5 mm. long; fruit to 6 mm. in diameter,
pergamaceous, glabrous or sparsely lepidote, the open valves about
1 cm. long, half as wide; seeds 5-6 mm. long, ovoid, slightly com-
pressed, black, lustrous with small light brown strophiole. This
species with notably stout angled and nodose branchlets was com-
pared by both Rusby and Mansfeld with P. oppositifolia M. Arg.
of Cuba with lepidote branchlets, looser leaf -pubescence and, accord-
ing to Rusby, much smaller seeds. The Rusby specimen in fruit
has smaller leaves matched however by Krukoff 5710 determined
in herbaria P. glabrata (Schott) Baill. with alternate glabrate leaves;
P. ferruginea (Schott) M. Arg., also Brazilian, is similar in pubes-
cence but the leaves are all alternate; possibly the character is not
valid as some specimens show lower alternate leaves, upper opposite.
In this case the Peruvian specimen may be a variant of P. ferruginea.
P. citriodora Baill., more distinct, has alternate acuminate sparsely
lepidote leaves. F.M. Neg. 5374 (P. Mildbraediana).
Loreto: Flooded high woods Yarina Cocha, Tessmann 3406 (type,
P. Mildbraediana). Rio Acre: Mouth of Rio Macauhdn, Krukoff
5627 (det. Steyermark, P. ferruginea)', 5710 (det. Steyermark, P.
glabrata). Bolivia.
Pera distichophylla (Mart.) Baill. Etude Gen. Euphorb. 434.
1858; 7. Spixia distichophylla Mart. Flora 24, Beibl. 2: 30. 1841.
Branchlet tips early reddish sericeous; petioles 1-5 mm. long,
canaliculate, soon glabrate; leaves distichous, lanceolate-elliptic or
-ovate, 7-15 cm. long, 2-5 cm. wide, more or less acuminate, often
obtusely contracted to petiole or acute, dark above, fuscescent or
glaucescent beneath, glabrous, minutely patellate-glandular beneath
near midrib, the 5-10 secondary nerves slender; male involucres
spicate in axils, the spikes only a cm. long, peduncle 2-4 mm. long,
unibracteolate, sericeous tomentose, about 3-flowered, 4-6 mm. in
diameter, coriaceous; calyx 1.5-2 mm. long, subentire, densely
sericeous with reddish rigid trichomes; stamens 3-5, stamen column
accrescent, glabrous above; female flowers and fruit unknown.
Another Amazonian species with simple pubescence that may occur
is P. heterodoxa M. Arg., 3, the involucre bisexual, bract 1; stellate-
lepidote species likely to be found include P. cinerea [Poepp.] Baill.
8 (P. tomentosa (Benth.) M. Arg.), and P. citriodora Baill. 11, the
first with male calyx developed, leaves tomentose beneath, the latter
108 FIELD MUSEUM OP NATURAL HISTORY BOTANY, VOL. XIII
with alternate sparsely lepidote leaves, male calyx none. The
Williams specimen from a 15 meter tree with thin rather smooth
bark. Illustrated, Pax & Hoffm. I.e. 5. F.M. Neg. 7165.
San Martin: Tarapoto, Williams 6089? Loreto: Yurimaguas,
Poeppig. Amazonian Brazil.
15. HEVEAAublet
Reference: Ducke, Archiv. Inst. Biol. Veg. Rio Jan. 2: 217-246.
1935; with bibliography.
Tall lactescent trees with trifoliate alternate leaves on long
petioles that are glandular at apex, the entire shortly petiolulate
leaflets pinnate-veined and usually membranous. Flowers small,
monoecious, apetalous, in paniculate cymes, the central flower of
which is often female, the others male, these in bud ovoid or globose,
at anthesis 5-dentate or 5-lobed. Disk glands small, 5, free or con-
nate. Stamens 5-10, the filaments connate, anthers sessile, regularly
or irregularly 1-2-verticillate, the column extended. Female flowers
with 3-celled ovary, ovules solitary, stigma stout, more or less
bilobed. Capsule large, splitting into subfleshy (usually) bivalved
follicles with large seeds. The similar Joannesia Veil, is weakly
distinguished by the larger petaliferous flowers; J. hevioides Ducke
is the species.
Source (most important) of natural rubber, the genus has received
much study, but unless genetic investigations prove to be conclusive
the status of its components will not be clearly understood. Ducke,
Arquivos Serv. Florestal 2, No. 1: 38. 1943, listed the four most
important species, commercially, as H. brasiliensis, H. Benthamiana,
H. lutea and H. guianensis. While Pax recognized 17 species, Ducke
12 (at most), J. T. Baldwin Jr., Amer. Journ. Bot. 33: 215. 1946,
wrote: "there are nine or fewer species characterized by much
introgressive hybridization (one could with reason reduce the genus
to a single species)." This student considers H. paludosa Ule and
H. humilior Ducke from Iquitos probably representatives of a
hybrid swarm of H. confusa, (i.e. H. pauciflora sens, lat.) and H.
guianensis (?). The following synopsis is prepared from the notably
acute and sensible work of Ducke, which, based on many years of
field observations and studies in conjunction with consideration of
type materials, furnishes a fine example of the methods that must
be employed before many groups, at least the more complex, can
be interpreted taxonomically with a reasonable degree of usefulness
FLORA OP PERU 109
for all students. His revision, it may be noted, gives the Brazilian
native names with English equivalents, I.e. 222-223, mostly not
repeated here; he has a supplementary paper detailing distribution
and natural hybrids, Arquivos Serv. Florestal 2, No. 1: 25-43. 1943.
Hevea, according to Seibert, Ann. Mo. Bot. Gard. 35: 117-121.
1948, has been known in cultivation for seventy-one years, trees
coming into feasible rubber production only after ten years; the
seeds see remark under H. pauciflora may be a source of economic
products.
Leaves dull and usually paler beneath.
Anthers 5(4-8) in a single sometimes irregular verticil, rarely
incompletely in 2 verticils; leaves erect; pedicels rarely 4 mm.
long; male buds obtuse or barely acuminate. . H. guianensis.
Anthers 7-10 in 2 more or less regular verticils; leaves reclinate
or semi-erect, if erect, pedicels elongate in part.
Pedicels short; leaves not erect.
Disk of male flowers minute or obscure, their buds acuminate;
anthers normally 10 in 2 regular verticils . H. brasilensis.
Disk of male flowers well-developed, their buds obtuse or
obtusely acuminate; anthers 7-10, the 2 verticils more
or less regular.
Male buds obtusely subacuminate; disk glands elongate.
H. humilior.
Male buds obtuse; disk shortly 5-glandular or -lobate.
H. pauciflora.
Pedicels of female flowers 5-10 mm. long; leaves erect.
H. paludosa.
Leaves lustrous both sides, concolored, reclinate to suberect.
H. nitida.
Hevea brasiliensis (Willd.) M. Arg. Linnaea 34: 204. 1865;
235. Siphonia brasiliensis Willd. ex Adr. de Juss. Euphorb. Tent.
40. pi 38b, figs. 1-6. 1824, not S. brasiliensis HBK. Nov. Gen. &
Sp. 7: 171. 1825. H. janeirensis M. Arg. in Mart. Fl. Bras. 11, pt. 2:
706. 1874. H. brasiliensis (HBK.) M. Arg. var. janeirensis (M.
Arg.) Pax, Pflanzenreich IV. 147: 121. 1910.
Gross character of H. guianensis but the glabrous membranous
acuminate leaflets with duller and as to type paler lower surface and
curved downwards instead of erect and, especially, the anthers
normally 10 in 2 regular verticils; flowers whitish-yellow, acuminate
110 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
in bud as the lanceolate lobes after anthesis; pedicels tomentulose,
those of the female flowers longer and maculate above as the calyx-
base; ovary typically puberulent with sessile stigmas; capsules rather
large with oblong spotted seeds 2.5-3 cm. long. Var. Randiana
(Huber) Pax or forma, (Huber) Ducke, I.e. 224 and 236, origin
unknown and scarcely in Peru, has more lanceolate leaflets, glabrous
smaller ovary; var. subconcolor Ducke, Archiv. Jard. Bot.
Rio Jan. 6: 55. 1933, or forma, Ducke, Archiv. Inst. Biol. Veg.
Rio Jan. 2: 224 and 236. 1935, the leaves nearly equally green both
sides, has been collected as near Peru as Sao Paulo de Olivenga
and doubtless occurs; var. acreana Ule, Bot. Jahrb. 50: Beibl. 114:
14. 1914, "may correspond to the luxuriant individuals which were
frequently observed in the fertile soils of Acre territory." (Ducke.)
For the cytology of this species see Arquivos Serv. Florestal 2,
No. 2: 46.
Synonymy after Schultes, Bot. Mus. Leaflets Harvard 14: 79-
86. 1950, who gives a lucid historical summary of the nomenclature
together with copies of the analytical plate and photograph of the
Paris type of Jussieu. Illustrated, Hook. Icon. 26: pi. 2573.
Trees normally 20-30 meters, sometimes 40 meters tall, generally
in the forests of recent alluvial soils, inundable or not; is now nearly
the sole rubber furnishing Hevea and the only one in plantations, its
latex exceeding all others in quality and quantity (Ducke) . Williams,
Field Mus. Bot. 15: 48-49, 274. 1936, gives a good account of rubber
collection accompanied by a photograph of this species in cultiva-
tion. His collections were from dense lowland but flood-free forests,
the cylindric trunks to 3 dm. in diameter, bark pinkish to dark
brown, scaly, twigs grayish pubescent.
Loreto: La Victoria, Williams 2931. Near Iquitos, Williat
8071(1}. Pro, Williams 2003 (det. Seibert). Caballo-Cochz
Williams 2176 (det. Seibert). Rio Itaya, Williams 206 (det. Seibert).
Cerro de Escaler, Ule 6909. Rio Acre: Mouth of Rio Macauhan,
Krukoff 5468 (hybrid with guianensis, var. lutea, det. Seibert).
Amazonian Bolivia and Brazil. "Siringa," "shiringa legitima,"
"jebe debil fino" (Williams), "seringuerira branca," "seringuein
preta" (Brazil).
Hevea guianensis Aublet, PL Guian. 2: 871. 1775; 227. H.
peruviana Aublet, I.e. 4: pi. 335. H. lutea (Spruce) M. Arg. Linm
34: 204. 1865. Siphonia lutea Spruce ex Benth. in Hook. Journ.
Bot. Misc. 6: 370. 1854. H. peruviana Lechl. ex Benth. in Benth.
FLORA OF PERU 111
& Hook. f. Gen. PI. 3: 290. 1880. H. cuneata Huber, Bol. Mus.
Goeldi4: 578. 1906.
Typically with glabrous foliage borne in an erect position, very
dark green, dense, the older branchlets with persistent scales at the
terminal bud, the mature leaflets firm-membranous to thin-cori-
aceous, commonly obovate, obtuse to rounded, apiculate, about
10 cm. long, petiolules (2) 5-7 mm. long, petioles often 1 dm. long;
panicles more or less reddish-tomentulose, early floriferous nearly
to base, becoming about as long as the leaves and flowerless below;
male buds subglobose, typically obtuse (to acuminate in vars.), the
calyx lobed to the middle, about 3 mm. long, disk insignificant,
anthers 5 (4-6) ordinarily in one perfect verticil, sometimes with
one anther lower, or, in var. lutea, verticil irregular; female flowers
with sessile stigma or style developed; ovary glabrous or sericeous;
capsule deeply divided, 2.5-4 cm. long, the subglobose seeds with
brown spots and lines, 17-20 mm. wide, 14-18 mm. thick. The
first described species and highly variable; the most important form
is doubtless var. lutea (Spruce) Ducke & Schultes, Caldasia 3: 249.
1945, the leaves somewhat, the flower buds distinctly acuminate,
and, especially, the anther-verticil irregular, ovary sericeous, stigmas
usually not sessile. An apparent form of this with the leaves some-
times lightly pubescent on the nerves beneath has been designated
var. cuneata (Huber) Ducke, Archiv. Jard. Bot. Rio Jan. 6: 51.
1933 (H. lutea, var. cuneata Huber, Bull. Soc. Bot. Fr. 49: 48. 1902),
or H. lutea var. pilosula Ducke, I.e. 231, these possibly the same
fide Ducke, Arquivos Serv. Florestal 2, No. 1: 21. 1943, or the former
var. occidentalis Ducke, I.e. 229. 1935, subsp., 223, based on cubical
angled seeds with few lines, very large spots; cf. also Ducke, I.e.
229. 1935. H. peruviana Lechl. seems to be merely a form of
var. lutea, the stigmas sessile, this "a very inconstant character."
(Ducke.)
Attains 30-40 meters with cylindrical stem, latex yellow; fre-
quent in the upper rain forests. The rubber of the var. lutea is said
to be better than that of the typical form which hybridizes with
H. pauciflora; see H. paludosa. Illustrated, Pflanzenreich IV.
147: 127; Archiv. Jard. Bot. Rio Jan. 5: pi. 18 (seeds). F.M. Negs.
32472; without number (H. cuneata); 7101 (var. lutea).
Loreto: Cerro de Canchahuaya, (Huber 1377, type, var. cuneata).
Pongo de Manseriche, (La Rue, det. Ducke). Yurimaguas, Killip
& Smith 28706 (det. Seibert). Pinto-cocha, Rio Nanay, Williams
818 (det. Seibert var. lutea). Puno: San Govan, (Lechler, type,
112 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
H. Lechleriana). Rio Acre: Rio Macauhan, Krukoff 5800 (var.
lutea, det. Seibert). Amazonian region to Dutch Guiana. "Shi-
ringa amarilla," "shiringa del cerro," "seringueira itauba" (Brazil).
Hevea hurnilior Ducke, Rev. Bot. Appl. Agric. Trop. 9: 624,
629. pi 18. 1929; 237.
A small tree (as to type) with horizontal or reclinate leaves,
oblong- or elliptic-obovate leaflets usually 8-12 (15) cm. long, 4-5
(7.5) cm. wide, shortly acuminate, firm-membranous to subcoria-
ceous, glabrous, soon lustrous above, dull and paler beneath; panicles
to 15 cm. long, sparsely pubescent; male pedicels 1-2 mm. long,
female twice as long, the buds ovoid, subacuminate or obtuse, sparsely
ashy puberulent, at anthesis sepals obtusely acuminate; anthers
nearly always 10, in 2 verticils, disk glands elongate-lanceolate; ovary
(female flowers) globose, sericeous, stigma subsessile; capsule about
size of H. guianensis but not so deeply sinuate between the carpels
and the spotted lineolate seeds are a little longer in relation to their
thickness. Attains 3-5 meters but not yet known from primary
formations (Ducke in 1935). Illustrated, Ducke, Archiv. Jard. Bot.
Rio Jan. 5: pi 20 (flowers, capsule and seed).
Loreto: Abundant between the low secondary growth near Iquitos
at start of route to Morono-cocha (Ducke 20602, type) ; also Kuhl-
mann.
Hevea nitida Mart, ex M. Arg. in Mart. PI. Bras. 11, pt. 2:
301. 1874. H. viridis Huber, Bull. Soc. Bot. Fr. 49: 48. 1902; 238,
fide Seibert and Schultes.
Unique in the character of the leaflets which are equally green
and lustrous on both surfaces; medium-sized tree, glabrous or the
inflorescence lightly pubescent; leaflets finally subcoriaceous, elliptic-
or oblong-obovate, shortly acuminate, reclinate or horizontal; flowers
nearly like those of H. pauciflora, capsule in form like H. brasiliensis,
not larger than that of H. guianensis; seeds angulate-ellipsoid, about
14 mm. long. Ducke remarks that the leaves of H. brasiliensis var.
subconcolor while nearly green both sides are not lustrous beneath.
The presence of interflush short-shoots as well as the glossy under
surface of the leaflets leaves little doubt that H. viridis should be
referred to H. nitida (Seibert). Illustrated, Ducke, I.e. pis. 2 and 8
(flowers, capsules and seed). F.M. Negs. 6631; without number
(H. viridis).
FLORA OF PERU 113
Loreto: Leche, Pampa del Sacramento, lower Huallaga and
Ucayali and, especially, swampy banks of the Rio Yanayacu, ( Huber
1534, type H. viridis). Rio Nanay, Williams 889 (det. Seibert). To
Amazonian Colombia and Brazil. "Puca shiringa," "shiringa entre-
fina colorada" (Huber), "urcu shiringa mapa" or "jebe debil mu-
erto" (Williams).
Hevea paludosa Ule, Bot. Jahrb. 35: 666. 1905; 237.
Glabrous except the lightly pubescent inflorescence; petioles
shorter than leaflets (in type), petiolules 3-4 mm. long; leaflets
obovate-lanceolate, erect, firm-membranous, 7-10 cm. long, 2.5-3
cm. wide, or larger, acuminate, lustrous above, pale and dull beneath;
inflorescence about 1.5 dm. long, subglabrous, the slender pedicels
2 mm. long for male flower, 5-10 mm. for female, the buds of the
former subacuminate but the apex itself obtuse, the lobes in anthesis
obtusely acuminate; anthers 7-10 in two more or less regular verticils
but frequently incomplete; disk with small obtuse or acute glands;
ovary globose, sericeous, stigmas sessile; capsule 3-3.5 cm. long,
3-4 cm. broad. This plant has been determined by Seibert as a
hybrid between H. pauciflora and H. guianensis var. lutea. Accord-
ing to Ducke it is common at the type locality. Swampy regions
outside of flood-area near Iquitos, the type 15-30 meters high.
Illustrated, Ule, I.e. 668 (flowers and fruit). F.M. Negs. 5369;
24365.
Loreto: Near Iquitos, Ule 6260, type. Mishuyacu near Iquitos,
King 128. Tierra Doble, Rio Nanay, Williams 918 (det. Seibert
as noted above). "Jebe debil fino" (Williams).
Hevea pauciflora (Spruce) M. Arg. Linnaea 34: 203. 1865;
238. Siphonia pauciflora Spruce ex Benth. in Hook. Journ. Bot.
Misc. 6: 370. 1854. H. membranacea M. Arg. in Mart. Fl. Bras. 11,
pt. 2: 299. 1874.
Glabrous (typically) except for the slightly tomentulose panicles
or puberulent flowers, these in bud, at least the male, globose-ovoid
and obtuse; leaflets horizontal, suberect or subreclinate, oblong-
obovate, obtusely acuminate, often about 8-16 cm. long, 4-8 cm.
wide, somewhat lustrous and green above, dull and paler beneath;
calyx lobes acuminate, puberulent both sides; disk glands well-
developed, bifid or emarginate; male flowers with 10 anthers in 2
verticils; ovary (female flowers) sericeous or glabrous (f. leiogyne
Ducke), stigmas sessile; capsules large to rather small but the carpels
always well-separated. Illustrated, Hook. Icon. pis. 2574, 2575.
114 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
A rather tall tree rarely used for rubber in the typical state, the
leaves membranous even in age, firm in var. coriacea Ducke, I.e. 239.
Conservatively this could be drawn to include H. humilior and
H. paludosa. Rather allied, apparently, is H. Benthamiana M. Arg.,
232, that however not yet known from within Peru, the male buds
acuminate and usually with only 6-9 anthers, the leaflets pubescent
beneath. If Williams 2503 actually belongs here it records the
species as attaining 50 meters or more, trunk over 3 dm. in diameter,
bark dark brown, closely fissured, latex turning pink. This and
Klug 812 have been determined by Seibert as a hybrid with H.
brasiliensis.
The seeds of this species according to Seibert in his important
article "Uses of Hevea for Food," Ann. Mo. Bot. Gard. 35: 117-121.
1948, are a potential source of economic products useful to man.
Loreto: Lowland forest, Caballo-Cocha, Williams 2503. Mishu-
yacu near Iquitos, Klug 812. Upper Amazonian Brazil. "Siringa"
(Williams).
16. RICINUSL.
Reference: Pax & Hoffm., Pflanzenreich IV. 147. ix-xi: 119. 1919.
Flowering as an annual but persisting to become a shrub-tree,
the ample peltate leaves palmately lobed, the lobes serrate, and the
rather large monoecious flowers in terminal racemes or narrow
panicles, the lower male, the upper female; pedicels bibracteolate,
the male medially articulate. Stamens a great many, borne on
repeatedly branched filaments. Styles 3, united at base, plumose,
red. Capsules usually echinate, 3-lobed with 3 conspicuously carun-
culate black or often beautifully mottled seeds.
The single species, perhaps native to Africa, has long been widely
distributed and is one of the first plants known for economic use,
greatly varied see an encyclopedia but it is most important as
the source of castor oil.
In the interior of Peru, as at Mito, the oil of the seeds is used
medicinally and collected on water for making candles and "by ex-
pression, for lighting, especially for the lamps in the churches."
(Ruiz and Pavon.)
Ricinus communis L. Sp. PI. 1007. 1753; 119.
Herbaceous state often pruinose and purplish, friable; leaves
several dm. or even a meter broad, 5-11-lobed, glabrous, glaucous;
FLORA OF PERU 115
male calyx 6-12, female 4-8 mm. long; capsules large, the ellipsoid
flattened seeds 9-22 mm. long, 6-15 mm. broad, variously marbled
in color. Many forms have been given taxonomic recognition none
probably stable variations but some said to be rated more useful
for certain economic qualities.
Huanuco: Mito, established along trails, 1578. Loreto: Mishu-
yacu, clearing, King 1013; 94-1- Puerto Arturo, Williams 5141-
Along Rio Itaya, Williams 234- La Victoria, Williams 2886. Cuzco :
Naturalized in great abundance in the Prov. of Convencion (Herrera).
Arica: Juan Isern. "Higuerilla," "iguerilla," "castor bean,"
"castor oil bean," "higuerilla mexicana and de la tierra."
17. PLUKENETIAL.
Reference: Pax & Hoffm., Pflanzenreich IV. 147. ix-xi: 12-16.
1919.
Climbing shrubs with alternate bistipulate ovate usually basally
3-nerved leaves and monoecious apetalous flowers borne in spici-
form racemes, the 1-2 female at the base, the remaining male, the
calyx of these valvately 4-parted with 12-30 free stamens on a conical
receptacle. Female sepals 4. Ovary 4-celled, 4-alate, the styles
connate into a cylindric or obovoid column; stigmas short, entire
or bilobed. Capsules dehiscing, the cocci carinate, winged or biver-
rucose, the lentiform or globose seeds ecarunculate. Eleutherostigma
Lehmanianum Pax & Hoffm. I.e. 11, monotypic, Colombia and ap-
parently Ecuador, has filiform disk glands half as long as the 40-50
stamens, styles free above the short column, inflorescence paniculate-
racemose. Commemorates Leonard Plukenet (1642-1706), English
physician and botanist.
Plukenetia volubilis L. Sp. PI. 1192. 1753; 14. P. peruviana
M. Arg. Linnaea 34: 157. 1865.
Younger parts puberulent or pubescent, finally glabrate; petioles
2-6 cm. long; leaves ovate, basally broadly cordate, abruptly cus-
pidate, 7-13 cm. long, 5-10.5 cm. wide, reddish biglandular above
at insertion of petiole, not beneath, crenulate or dentate (sometimes
very minutely), membranous, puberulent on the nerves; racemes
several to 9 cm. long with 1 female flower on pedicel 5 mm. long at
base, lower male bracts 6-12-flowered, upper about 5-flowered, the
slender pedicels to 3 mm. long; stamens 20-24; female sepals acute,
1.5 mm. long, glabrous as ovary with slender style column 18-25
116 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
mm. long; capsules 3.5 cm. wide, 18 mm. high, the 4 cocci dorsally
carinate-winged, the lentiform corrugately veined seeds 15 mm.
broad. Illustrated, Pax, I.e. (flowers in part and capsule). F.M.
Neg. 7111 (P. peruviana).
San Martin: San Roque, Williams 7731 (det. Mansfeld). Juanjui,
King 3845; 3901 (det. Standley). Loreto: Mishuyacu, King 634;
Killip & Smith 29927 (det. Mansfeld). Iquitos, Williams 8123 (det.
Mansfeld). Florida, King 2056 (det. Standley). Near Mazan,
Mexia 6470 (det. Standley). Yurimaguas, Poeppig 2210. Without
locality, Ruiz & Pav6n (type, P. peruviana). Bolivia; West Indies.
"Sacha Yuchiqui," "sacha yuchi" (both Williams), "amui-o" (Klug,
Huitoto).
18. ELAEOPHORA Ducke
Scandent shrubs with long narrow axillary panicles (male) of
dioecious apetalous flowers, the female green in short axillary
racemes, and entire or subentire petioled leaves. Female calyx with
4 imbricate sepals; receptacle elongating; style column 4(3)-costate
with 4 (3) spreading branches at tip. Fruit indehiscent, 4(3)-lobate,
fleshy, the very large seeds ecarunculate. Male calyx 4-parted,
sepals finally reflexed, the subconic receptacle with many glands
mixed between the filaments, in this respect differing from the allied
Plukenetia and, moreover, with the large seeds of Omphalea (Ducke).
Elaeophora polyadenia (M. Arg.) Ducke, Archiv. Jard. Bot.
Rio Jan. 5: 146. 1930. Plukenetia polyadenia M. Arg. in Mart. Fl.
Bras. 11, pt. 2: 334. 1874. P. macrostyla Ule, Verh. Bot. Ver.
Brandenb. 50: 80. 1908, fide Croizat.
Glabrous; petioles slender, 2.5-3 cm. long; leaves elliptic or ovate,
rounded-obtuse or subtruncate at base, abruptly caudate-acuminate,
8-9 cm. long, 4.5-5 cm. wide, entire (or nearly), lustrous, uniglandular
(or rarely biglandular) above the base, 5-nerved, firm-membranous;
inflorescence axillary, laxly and narrowly paniculiform, naked for
2-3 cm. at base; female flowers with 2.5 mm. long style-column,
branches 1 mm. long, ovary 5 mm. long with 4 elliptic wings, 7 mm.
wide; cymules 2-3 mm. long, stiped; male buds 2.5 mm. long, acute;
receptacle subconic, densely glandular, lobulate; filaments much
narrowed apically, about 25, anthers didymous, cells free below,
connective produced. E. abutaefolia Ducke, I.e. 4: 112. pi. 9. 1925,
of Brazil, the type species, seems to be similar but has style columns
3-4 mm. long, branches 2-2.5 mm. long, fruit to 10 cm. long, 11 cm.
FLORA OF PERU 117
wide, glabrous, lustrous, keeled; seeds 5.5x4x3.5 cm., opaque.
See also Ducke, I.e. 5: 146. 1930. Ule separated his species on the
basis of 2 leaf-glands and acutish buds.
Loreto: Yurimaguas, Poeppig 2385, type. Brazil.
19. APODANDRA Pax & Hoffmann
Reference: Pax & Hoffm., Pflanzenreich IV. 147. ix-xi: 20. 1919.
Habit of Plukenetia but leaves always pinnately nerved and,
especially, the (12) 15-35 anthers sessile on a globose receptacle,
free or more or less confluent. Inflorescences short, unisexual or
finally glomerate-spiciform. Nearly/ 1 ragariopsis St. Hil. of southern
Brazil with indehiscent fruit and could readily be included with it
in Plukenetia as was the former by Bentham.
Leaves oblong or nearly, the larger about 3 times longer than wide;
leaf-nerves 6-7; style column obovoid A. Buchtienii.
Leaves elliptic-ovate, the larger about twice as long as wide; nerves
usually 4-5.
Style column obovoid; stamens 12-15 A. brachybotrya.
Style column cylindric; stamens 15-16 A. loretensis.
Apodandra brachybotrya (M. Arg.) Macbr., comb. nov.
Plukenetia brachybotrya M. Arg. Linnaea 34: 158. 1865; 16.
Younger parts as the tips of the terete branches early appressed
pubescent becoming glabrate as entire plant; petioles 10-15 mm.
long; leaves elliptic, usually twice as long as broad, 8-10 cm. long,
abruptly cuneately narrowed to petiole, abruptly acuminate, the
acumen often short or rounded-obtuse and very acute; minutely
denticulate, membranous, brownish biglandular at base; nerves
slender; racemes commonly shorter than the petioles, male pedicels
longer than calyx, female elongate and even exceeding the male
part of the raceme; stamens 12-15; column style obovoid. Not
seen by the monographers. F.M. Neg. 8594.
Huanuco: Chicoplaya, [Ruiz &} Pavdn, type.
Apodandra Buchtienii Pax, Pflanzenreich IV. 147. ix-xi: 21.
1919. Plukenetia Buchtienii Pax, Repert. Nov. Sp. 7: 110. 1909.
Simulating A. loretensis but leaves lanceolate or oblong, narrowed
to base, 6-15 cm. long, 2.5-5 cm. wide, biglandular at base, glandular-
118 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
denticulate, chartaceous, secondary nerves 6-7; flowers monoecious,
the racemes either unisexual, male 1-3 cm. long, the female 1-
flowered, about 1.5 cm. long, or bisexual with 1 female flower at
base; pedicels filiform, the male 5-8 mm. long, female to 1.5 cm.
long; male sepals 3, orbicular-ovate, acute or obtuse, 2 mm. long;
stamens 27-33; female sepals acuminate, scarcely 1 mm. long; ovary
wings divergent; style column stoutly obovoid; stigmas indistinct;
capsules cruciately 14 mm. wide, half as long, the cocci unitubercled,
the laterally compressed seeds 4.5 mm. long, rugulose and slightly
granulate, lutescent- and brown-spotted.
Rio Acre: Seringal San Francisco, Ule 9539, type. Bolivia.
Apodandra loretensis (Ule) Pax & Hoffm., Pflanzenreich IV.
147. ix-xi: 21. 1919. Plukenetia loretensis Ule, Verh. Bot. Ver.
Brandenb. 50: 81. 1908.
Slender branchlets tardily glabrescent; stipules minute; petioles
pubescent, 8-12 mm. long; leaves elliptic or ovate, contracted at
base, shortly and obtusely acuminate, mucronulate, 5-6 cm. long,
about 3 cm. wide, crenulate, membranous, 2-several glandular above
the base, early pilose on the 4-5 arcuate nerves; racemes 1-2 cm.
long, all parts pubescent, bracts to 1 mm. long, ovate, acute, the
male branches only 2-3-flowered, the pedicels 2-3 mm. long, sepals
4, 1-2 mm. long, elliptic, acute; stamens 15 or 16; female sepals
acute, nearly 1 mm. long, thickened at base within margin; ovary
pubescent, wings refracted, style column 4 mm. long, pubescent,
cylindric, the stigma little broader; capsules unknown. F.M. Neg.
24572.
Loreto: Near Iquitos, Ule 6837, type.
20. MEGALOSTYLIS Sp. Moore
Reference: Pax & Hoffm., Pflanzenreich IV. 147. xiv. Add. VI:
'33. 1919.
Shrub probably scandent with alternate ample petioled basally
5-nerved leaves and monoecious apetalous flowers borne in short
axillary bracteate cymes. Male calyx closed in bud, valvately
3-parted at anthesis; stamens 6, the filaments connate into a cylindric
column, the bicelled anthers didymous; disk and rudimentary ovary
none. Female sepals 6, imbricate; disk none; ovary smooth, 3-celled;
styles large, clavately connate, entire at tip; ovules solitary. Fruit
unknown. The related Angostylis longifolia Benth., 29, of the upper
Amazon has the 2-3 male flowers at the nodes, the female solitary
FLORA OF PERU 119
in the axils, stamens about 20 and ovary muricate, the urceolate
style column with bifid spreading lobes; perhaps rightly in the same
genus are Astrococcus Benth. and Haematostemon (M. Arg.) Pax &
Hoffm., both with racemose flowers, 4 stamens with glandular disk,
the glands extrastaminal in the former, intrastaminal in the latter;
the leaves of A. cornutus Benth. are puberulent on nerves, membra-
nous, of A. coriaceus Baillon, glabrous, coriaceous, this H . coriaceus
(Baill.) Pax & Hoffm., both to be expected.
Megalostylis Poeppigii Sp. Moore, Journ. Bot. 54: 250. pi.
544, fig- 2. 1916; 34.
Branchlets as slender petioles, these 2.5-4.5 cm. long, and
peduncles sericeous puberulent; leaves broadly ovate, truncate or
obscurely cordate, obtuse or sometimes shortly cuspidate, 11 cm.
long, 9 cm. wide, often smaller, membranous, sparsely and shortly
pubescent both sides, nerves nearly horizontal; cymes 2-2.5 cm.
broad, bracts ovate, obtusish, 3-4 mm. long; sepals puberulent, the
male oblong-spathulate, obtuse, 2 mm. long, the female little longer,
ovate-lanceolate, longer than staminal column; style column 8 mm.
long, little curved; ovary globose, sericeous.
Peru(?): Without data, (Poeppig, type).
21. TRAGIA [Plumier] L.
Reference: Pax & Hoffm., Pflanzenreich IV. 147. ix-xi: 32-101.
1919.
Herbs often more or less suffrutescent and climbing and usually
harsh to touch, with stinging trichomes, the alternate petioled leaves
ordinarily more or less cordate at base. Flowers monoecious,
apetalous, in axillary or terminal racemes, only a few at base female,
all usually solitary in the axils of the persisting bracts. Male calyx
valvately 3 (-5) -parted, disk glands often obsolete; stamens usually 3
(exceptionally fewer, or 4-20 or 30-50), free; ovary rudiment minute
or none. Female calyx 3-6-lobed, the imbricate lobes sometimes
pinnate. Ovary 3-celled, the more or less connate styles with entire
stigmas. Capsules breaking into 3 bivalved cocci, the globose seeds
ecarunculate.
Inflorescence biparted; fruiting pedicels to 8 mm. long, usually
shorter.
Stamens 6 with urceolate disk glands; leaves often half as wide as
long; female pedicels to 8 mm. long T. fallax.
120 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Stamens 1-3 with no disk; leaves rarely half as wide as long;
fruiting pedicels about 4 mm. long T. subhastata.
Inflorescence not biparted; fruiting pedicels greatly elongate; stamens
1-3, no disk T. volubilis.
Tragia fallax M. Arg. Linnaea 34: 179. 1865; 36.
Climbing, the hirsute terete stems tardily glabrescent; stipules
about 5 mm. long, acuminate; petioles 2-6 cm. long; leaves ovate or
elliptic from a subhastate-cordate base with narrow sinus, acuminate,
7-13 cm. long, 4-7 cm. wide, irregularly dentate, palmately nerved,
the nerves early puberulent on both sides; inflorescences opposite
leaves, pubescent, biparted, peduncles 2-5 cm. long, male branch
2-6 cm. long, many-flowered, the female 3-7 cm. long, 2-8-flowered;
bracts linear-lanceolate, 1 mm. long, the male 1-3-flowered, pedicels
2 mm. long, articulate near base, buds pyriform, opening 2 mm. long
or little longer, sepals 3, obovate, hirsute, stamens 6(5-7), disk
glands present, filaments apically clavate, rudimentary ovary none;
fruiting pedicels to 8 mm. long, sepals 6, lanceolate, acute, hirsute;
styles connate to middle, little papillose; capsules 8 mm. thick, 5 mm.
long, hirsute; seeds 3 mm. thick. T. japurensis M. Arg., 36, has
3 mm. long bracts, the male branches only 2 cm. long, 9 stamens
with filiform filaments, papillose styles; T. Lessertiana (Baill.). M.
Arg., 37, has 10-20 stamens, leaves acute at base. Determinations
by Mansfeld except as noted. F.M. Negs. 7155; 34165.
San Martin: Tarapoto, Ule 6636; Williams 5558; 6795. Juanjui,
Klug 4%07 (det. Standley). Huanuco: Chicoplaya, Ruiz & Pavdn,
type. Ayacucho: Kimpitiriki, Rio Apurimac Valley, Killip & Smith
22948.
Tragia subhastata Poepp. in Poepp. & Endl. Nov. Gen. & Sp.
3:20. pi. 223. 1845; 46.
High-climbing liana, the younger stems as the petioles, these
2-5 cm. long, and the young leaves both sides on the nerves puberu-
lent, stipules 3-4 mm. long, acute; leaves oblong-ovate or ovate,
acuminate, deeply and acutely cordate at base, 7-13 cm. long, 3-6.5
cm. wide, irregularly dentate, palmately nerved; inflorescence bi-
parted, the female branch 3-6-flowered, the male shorter, fewer
flowered; rachis puberulent; bracts lanceolate-ovate; male pedicels
articulate above the base and there dilated into a persisting annulus,
the female 2-3 mm. long; stamens 3; female sepals ovate, persisting,
FLORA OF PERU 121
unequal; ovary densely fulvous-hirsute; styles shortly connate.
F.M. Neg. 32546.
Huanuco: Tocache, Poeppig 1935, type.
Tragia volubilis L. Sp. PI. 980. 1753; 47.
Vine with elongate slender stems, the younger parts somewhat
hirsute, the often stinging trichomes more or less persisting especially
on the nerves of the more or less dentate leaves, these variable in
shape and size but usually oblong-lanceolate from a truncate or
more or less openly cordate base, acuminate, several cm. to a dm.
long or longer, about 1 cm. to 5 cm. wide or wider; petioles 1-3 (5)
cm. long; stipules lanceolate, about 3 mm. long; racemes lateral,
slender, 1-several cm. long, the lanceolate bracts persistent, the
female flower usually solitary at base on pedicel 4-5 mm. long,
much elongating in fruit; male pedicels many, 2-3 mm. long, articu-
late above the middle; sepals 3; stamens 1-3; filaments short, stout;
female sepals 6, persisting; styles more or less connate; capsules
5-7 mm. thick, deeply 3-lobed, setose. A form with triangular
ovate leaves, coarsely serrate-dentate, elongate fruiting peduncle
and 2-3 stamens has been designated var. serra (Poepp.) M. Arg.,
a lanceolate subentire leaved form, var. lanceolata M. Arg. Illus-
trated, Pax & HofFm., I.e.; var. serra Poepp. in Poepp. & Endl. 3:
pi. 223. F.M. Neg. 32547 (var. serra}.
San Martin: Tarapoto, Spruce 4850 (type, var. lanceolata).
Morales, Williams 5688. Loreto: Yurimaguas, Poeppig (type, var.
serra). Widely distributed.
22. ACALYPHAL.
Reference: Pax & Hoffm., Pflanzenreich IV. 147. xvi: 1-177. 1924.
Shrubs, less often trees, sometimes perennial herbs or annual
and then especially simulating Urtica species. Leaves petioled, alter-
nate, bistipulate, more or less dentate, usually ovate, basally 3-5-
nerved or pinnately nerved, frequently scabrous or hyaline puncticu-
late. Flowers usually monoecious, apetalous, in bisexual or uni-
sexual axillary or terminal inflorescences, if the former the female
flowers few at the base. Male flowers glomerulate in axils of small
bracts racemosely or spicately disposed as also the female but these
usually solitary or 2-4, the bracts often dentate. Disk none. Male
calyx valvately 4-parted; stamens generally 8, filaments free, anther
cells divaricate or pendulous, rudimentary ovary none. Female
122 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
sepals 3-5, imbricate; ovary 3-celled, the 3 free styles commonly
lacinulate, ovules solitary. Capsules separating into 3 bivalved
cocci, the seeds ecarunculate. The Peruvian species transferred by
Kuntze in his Rev. Gen. 2: 615. 1891 to Ricinocarpus [Burm.]
Kuntze have not been listed, as his action has become only historically
significant.
At least as concerns Peruvian species their definition by the
monographers is often obscure and it seems probable that there are
more names than natural specific units.
Female flowers at least minutely pedicellate; bracts minute.
Leaves palmately nerved, ovate with rounded-subcordate base.
A. villosa.
Leaves pinnately nerved, lanceolate or oblong-elliptic.
Inflorescence paniculate, sometimes narrowly; petioles mostly
or all elongate; leaves more or less acuminate.
Styles 30-40-lacinulate A. platyphylla.
Styles 10-15-lacinulate A. subandina.
Inflorescence spiciform; petioles rarely 1 cm. long; leaves nar-
rowly caudate-acuminate A. Macbridei.
Female flowers sessile; bracts more or less conspicuous.
Herbaceous annuals unless A. contermina with androgynous spikes,
these in part terminal except in A. arvensis.
Bracts minutely multidenticulate; stems puberulent.
A. contermina.
Bracts laciniate or deeply 3-9-lobed; indument usually some-
what pilose or hirsute.
Bracts not laciniate or setaceous.
Spikes androgynous; leaves ovate or oblong-ovate.
A. Forbesii.
Spikes male or female; leaves often roundish, basally 3-
nerved A. infesta.
Bracts filiform-laciniate or setaceous- tipped.
Leaves acuminate or caudate; spikes in part terminal.
Bracts laciniate to base; stems eglandular.A. subcastrata.
Bracts lobed, long-setaceous; stems glandular.
A. alopecuroides.
Leaves acutish or acute; spikes all axillary. . . .A. arvensis.
Ligneous plants at least below; spikes not androgynous, or rarely.
FLORA OF PERU 123
Inflorescence in part terminal.
Petioles at least some of them, and soon, much longer than
1 cm.
Female bracts entire or merely glandular below, long-
acuminate.
Leaves soon glabrescent beneath, 3-nerved from rounded
base; bracts large A. peruviana.
Leaves pubescent beneath, 5-nerved from cordate base;
bracts small A. subbullata.
Female bracts dentate, the middle tooth sometimes ex-
tended (unknown in A. bullata).
Leaves conspicuously bullate and reticulate-veined.
A. bullata.
Leaves obscurely if at all bullate, the lax reticulation
rarely prominent; species especially in flower similar.
Leaves glabrous or nearly beneath or scabrous with
minute trichomes, sometimes barbate in axils.
Leaf -dentations more or less spreading and distant;
bracts 5-6 mm. long in flower, teeth 5-10, the
middle extended A. stachyura.
Leaf-dentations (or leaves crenate) approximate.
Bracts 3-10-dentate, the teeth little unequal.
A. stenoloba.
Bracts often 9-15-dentate, the middle tooth more
or less extended.
A. mapirensis, A. macrodonta.
Leaves pubescent beneath, if merely puberulent,
densely.
Leaf -pubescence minute or rather sparse; bracts
unequal, 5-6 mm. long in fruit, 9-15-dentate.
A. mapirensis, A. macrodonta.
Leaf -pubescence usually dense; bracts often sub-
equal; some specimens seem to merge with
above.
Leaves more or less softly puberulent beneath;
bracts 3-4 mm. long, typically 5-9-dentate,
the middle tooth little longer. . . .A. benensis.
Leaves villlous-pilose beneath; bracts 4-6 mm.
long, often 12-15-dentate A. stricta.
Petioles all short, 4-10 mm. long.
124 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Female bract dentations unequal; leaves glabrate.
A. Ruiziana.
Female bract dentations subequal; leaves short villous.
A. padifolia.
Inflorescence all axillary.
Spikes bisexual.
Leaves pinnately nerved, oblong-ovate-lanceolate, often
7-15 cm. long, never subcordate.
A. samydaefolia, A. diversifolia.
Leaves palmately nerved, 3-nerved from base or subtri-
plinerved, small or more or less cordate.
Leaves not at all cordate, the petioles about 5 mm. long.
Leaves glabrous or nearly A. aronioides.
Leaves tomentose beneath A. Argomuelleri.
Leaves at least subcordate on petioles finally about as
long; bracts 9-17-dentate.
Leaves obtuse or acute, obscurely cordate; indument
little if at all glandular (obscure species).
Bracts 17-multidentate; branchlets eglandular.
A. reflexa.
Bracts 13-15-dentate; branchlets stiped-glandular.
A. brachyclada.
Leaves obviously acuminate, usually cordate.
Leaves openly or rounded cordate; glandulosity
lacking or not copious.
Indument dense; leaves dentate A. plicata.
Indument sparse and fine; leaves crenate.
A. tenuipes.
Leaves deeply cordate with imbricate lobes; gland-
ular indument pungent-resinous A.fulva.
Spikes unisexual (perhaps abnormal A. reflexa).
Leaves pinnately nerved (A. cuneata), A. obovata.
Leaves palmately nerved.
Indument in no part stellate.
Leaves green; petioles often elongate.
Erect shrub or tree; styles short. .A. macrostachya.
Scandent shrub; styles to 1 cm. long. .A. scandens.
Leaves red-spotted; petioles short A. Wilkesiana.
Indument stellulate A. dictyoneura.
FLORA OF PERU 125
Acalypha alopecuroides Jacq. Icon. Rar. 3: 19. pi. 620. 1786-
93; 22.
Resembles A. arvensis; stem-pubescence at least pilose-glandular;
leaves cuspidate- or caudate-acuminate, often more coarsely serrate;
spikes terminal and axillary, the former often with a few male flowers
at apex; female bracts hispid, 3-5-lobed with long setaceous tips;
styles bifid or entire. A weed like the superficially similar species
but apparently much less common.
San Martin: In clearing at Juanjui, Klug 4402 (det. Standley,
A. arvensis). Loreto: In pasture at Puerto Arturo, Yurimaguas,
Williams 5032. To Mexico and the West Indies.
Acalypha Argomuelleri Briq. Ann. Cons. Jard. Bot. Geneve
4: 229. 1900; 113. A. buddleifolia Pax & Hoffm. Pflanzenreich IV.
147. xvi: 113. 1924.
A shrub with divaricate-ascending branches; stipules caducous,
setaceous, 2 mm. long; petioles to about 5 mm. long; leaves elongate-
elliptic, acutish, obtuse or rounded, 3-4.5 cm. long, 10-15 mm. wide,
crenulate, coriaceous, rugulose-bullate, dark green and lustrous
above, white tomentose and prominently pinnate-nerved beneath,
subtriplinerved at base; spikes axillary, 3-8 cm. long, 1-2 basal
bracts female, the latter 3-flowered, 3-6 mm. long, about 20-30-
denticulate, reticulate, coriaceous, pilose or tomentose beneath and
strongly nerved; sepals acuminate; ovary villous; styles pectinately
lacinulate, about 4 mm. long; capsules smooth. Placed in herb,
by Mueller, Argoviensis, near A. accedens M. Arg. and A. Mandoni
M. Arg. (Briquet). Pax and Hoffmann separated their species on
the basis of the larger (to 1 cm. broad) female bracts; they described
the sepals as oblong, glandular as the ovary but these differences
appear to be varietal, if that, and partly due, probably, to age of
material. F.M. Negs. 24427; 5282 (A. buddleifolia}.
Amazonas: Chachapoyas, Mathews, type. Ancash: Huari below
Chavin de Huantar in the valley of the River Puccha, Weberbauer
3737 (type, A. buddleifolia).
Acalypha aronioides Pax & Hoffm. Pflanzenreich IV. 147.
xvi: 113. 1924. A. divaricata M. Arg. Linnaea 34: 34. 1865, not
Raf. 1836.
Densely branched shrub, the sometimes virgate or flexuose
branchlets as all younger parts puberulent-hirtellous and granular-
126 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
glandulose; stipules rigid, acuminate, 2-3 mm. long; petioles to
10 mm. long; leaves ovate-lanceolate, acute both ends or tips sub-
obtuse, 2-3.5 (5) cm. long, 1-1.5 (2) cm. wide, crenate-serrulate,
firm-membranous, glabrous above from the first, sparsely pilose on
the nerves beneath or nearly glabrous, 3-nerved from base; flowers
monoecious; spikes axillary, androgynous, to 1 cm. long, the male
part dense, shortly peduncled, one basal female, these bracts 1-
flowered, in flower 1 cm. broad, orbicular-reniform, more or less
7-15-dentate-crenate, subglabrous, granular-glandular, sepals ovate,
ciliate; ovary hirsute above, slightly glandular; styles about 4 mm.
long, connate below, somewhat 10-lacinulate above, 1-3 mm. high.
The type of A. divaricata is openly branched, the petioles more
slender, the pubescence minute and nearly wanting but otherwise
seems to be the same. F.M. Negs. 5278; 7123 (A. divaricata}.
Ayacucho: Shrubwood, 3,200 meters, Huanta, Weberbauer 7514
(det. Johnston, sp. nov.). In hedgerows, Ocros, Stork & Horton 10800
(det. Standley). Apurimac: Humid rocky places, Curahuasi, Vargas
9614- Quisapata, Abancay, Vargas 8964. Prov. Andahuaylas,
Vargas 8788. Cuzco: Valle de Urubamba, 2,800 meters, Herrera
2103; 1145; 3348; 3355. Torontoy, 2,400 meters, Cook & Gilbert
(det. Ewan). Valley de Lares, Soukup 45. Puno: Between Sandia
and Cuyocuyo, 2,100 meters, Weberbauer 836, type. Without
locality, Ruiz & Pavdn (type, A. divaricata). "Pespita" (Cook &
Gilbert).
Acalypha arvensis Poepp. in Poepp. & Endl. Nov. Gen. & Sp.
3: 21. 1845; 99.
Creeping-ascending-erect more or less pilose-hispid annual, the
trichomes spreading, those above somewhat yellowish; petioles to
3 cm. long or longer; leaves rhombic-ovate or -lanceolate, obtuse
at base, acute, usually 4-6 cm. long, 3-3.5 cm. wide or larger, thin-
membranous, basally 5-nerved; spikes androgynous, the upper nearly
entirely female, axillary, peduncled, 1.5-2.5 cm. long, 10-13 mm.
wide, the slender male part interrupted-spiciform; fruiting bracts
about 5 mm. wide, 4-7(9)-parted to middle with filiform-acuminate
divisions long-hirsute and glandular as the narrowly ovate sepals;
styles lacinulate; capsules to 2 mm. thick, apically pilose, the broadly
ovoid seeds about 1 mm. long. Williams noted the plant as a weed
in pastures and chacaras. F.M. Neg. 32497.
Loreto: Yurimaguas, Poeppig 2115, type; Williams 3937; 4267;
4465; 5040. To Mexico; Bolivia; Martinique.
FLORA OF PERU 127
Acalypha benensis Britton, Bull. Torrey Club 28: 304. 1901;
12. A. tomentosula Ule, Verb. Bot. Ver. Brandenb. 50: 79. 1908,
ide Pax & Hoffm.
A 1-5-meter shrub, the flexuose branchlets and petioles, these
..5-4 cm. long, early velvety tomentulose; leaves oblong or ovate,
ounded at or sometimes a little narrowed to the somewhat cordate
>ase, 5-16 cm. long, 2.5-10 cm. wide, serrate, membranous, finally
ubglabrous above, softly pubescent beneath, basally 3-5-nerved,
>therwise pinnately; flowers monoecious, the spikes dense, the
,essile male 4-7 cm. long, the terminal subsessile female about a
Im. long, these with truncate bracts 4-5 mm. long, 6 mm. wide,
lirtellous, 1-flowered, 5-9-lacinulate to one-third, the middle of the
larrowly triangular acute lobes a little larger; sepals ovate, acute,
;iliate; ovary pubescent; styles 7 mm. long, pinnately about 10-
acinulate, the lower divisions the smaller; seeds smooth.
San Martin: Tarapoto, Ule 6637 (type, A. tomentosula); Williams
W27 (det. Standley, A. mapirensis); 5658; 5773; 6262; 6688; 6746.
>an Roque, WiUiams 7003; 7475. Zepelacio near Moyobamba,
Klug 3347 (det. Croizat, A. Ruiziana; Standley, A. benensis).
Pongo de Cainarachi, Klug 2611. Hudnuco: Chinchao to Rio
ftuallaga, Seibert 2231 (det. A. C. Smith). Tingo Maria, Soukup
1119. Pozuzo, 4621 (det. Johnston). Junin: Puerto Yessup, Killip
fe Smith 26298 (det. Standley). Loreto: Balsapuerto, Klug 3100
[det. Standley); Killip & Smith 28491 (det. Croizat, A. stachyura).
^bove Pongo de Manseriche, Mexia 6168 (det. Standley). Rio
Mazan, Jose Schunke 22. Recreo, Yurimaguas, Williams 3938.
3uzco: Rosalina, 650 meters, Bues. San Pedro, Prov. Paucartambo,
1,350 meters, Vargas 6759. Bolivia. ' 'Yana-barilla" (Mexia).
Acalypha brachyclada M. Arg. in DC. Prodr. 15, pt. 2: 862.
L866; 125.
A shrub with slender terete glabrate branches and short (about
5 cm. long) divaricate apically densely leafy tomentulose branchlets
svith many axillary spikes; stipules lanceolate-ovate, scariose;
petioles 2-4 times shorter than leaf-blades, these 2-3 cm. long, 1-2.5
cm. wide, ovate, cordate at base, acuminate, obtusely serrate, mem-
branous, basally 5-7-nerved, shortly pubescent and glandular as
the branchlets and petioles; spikes 2-2.5 cm. long, entirely male
or often with 1 female bract at base, this small, 1-flowered, reni-
form-ovate, 13-15-dentate, the ovate teeth equal; sepals ovate,
subacute; ovary glabrous; styles small, rigid, about 5-lacinulate.
128 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
A. soratensis Pax & Hoffm., 126, has somewhat larger leaves softly
villous beneath, eglandular. F.M. Neg. 8505.
Peru(?): Without locality, [Ruiz &] Pavdn, type.
Acalypha bullata M. Arg. Linnaea 34: 17. 1865; 66.
Branches stout, rusty tomentose toward tips as petioles, these
2-4 cm. long, and leaves beneath, these ovate, rounded-cordate at
base, acuminate or obtuse, 6.5-10 cm. long, 3.5-5 cm. wide, serrulate,
the dentations revolutely recurved beneath, subcoriaceous, bullate,
sparsely pilose and lustrous above, 5-7-nerved at base; areola convex
above, the impressed veins often cruciately quadrate or biquadrately
impressed-sulcate; male axillary spikes to 8 cm. long, compressed
to 5 mm. wide, dense, erect, sessile; female flowers unknown and
therefore also the species relationship. Supposedly related species
but with monoecious flowers include the Ecuadorean A. tunguraguae
Pax & Hoffm., 66, male spikes slender, lax, the female bracts 5-
dentate and A. subbullata Pax & Hoffm., origin unknown, the
dense male spikes about 4 mm. broad, the female bracts entire
except for glandular-denticulate base. F.M. Negs. 5283; 34036.
Peru: Without locality, Ruiz & Pavdn, type, herb. Boiss.
Acalypha contermina M. Arg. Linnaea 34: 46. 1865; 25.
Branches subherbaceous, puberulent as the petioles (3-6 cm.
long) and leaves beneath, these sparsely pubescent or glabrate above,
rhombic-ovate or -lanceolate, cuneate to the narrowly cordate
base, acuminate, 6-12 cm. long, 2.5-5 cm. wide, serrate, membranous;
spikes sessile, slender, 1 dm. long or longer, multibracteate, the
terminal and subterminal female at tip with rudimentary male flowers
of elongate rigid sterile setae, the other spikes shorter, compressed,
10-13 mm. broad, the male 8-11 cm. long, 2 mm. broad, female
bracts 2-3-flowered, finally 8 mm. long, about 9 mm. wide, sub-
orbicular, barely acute, with 13-15 minute ovate densely stipitate
glandular teeth; female sepals broadly ovate, acutish; ovary hirsute
and papillose-subechinate, the styles above 4-7-lacinulate; seeds
1.5 mm. long. Possibly from Mexico as most of the closely related
species. F.M. Neg. 7141.
Peru(?): Without locality, ex herb. Pavon, type.
Acalypha dictyoneura M. Arg. Linnaea 34: 12. 1865; 160.
Branchlets, petioles, these 5-7 times shorter than leaves, and
spikes stellulate puberulent; stipules subulate, 2.5-4 mm. long,
FLORA OF PERU 129
deciduous; leaves ovate, acuminate, cordate or scarcely so, typically
about 8 cm. long, 4.5-5 cm. wide, sparsely pilose or glabrate above,
densely stellate tomentulose and prominently reticulate beneath,
firm-membranous, basally 3-nerved; flowers monoecious, the axillary
peduncled dense male spikes often 8-12 cm. long, 3.5 mm. thick,
the female 3-5 times shorter (or reduced to one bract) with flowering
bracts 1.5 mm. long, broadly ovate, obtuse or acute, 13-17-denticu-
late, the terminal tooth larger, stipitate, glandular; ovary hirsute;
styles 3.5 mm. long, rigid, pectinately 12-15-lacinulate. "Forma b."
reducta M. Arg. I.e. 13 seems well enough marked to become more
appropriately var. reducta (M. Arg.) Macbr., comb, nov., indument
more persisting; leaves about 2.5 cm. long, 2 cm. wide, obtuse or
scarcely cordate, tip more obtuse; spikes often reduced to one
bract. F.M. Neg. 5289.
Amazonas: Chachapoyas, Matthews, type. Without locality,
Ruiz & Pavdn (type, var. reducta). Ecuador.
Acalypha diversifolia Jacq. Hort. Schoenbr. 2: 63. pi. 244-
1797; 107. A. microgyne Poepp. in Poepp. & Endl. Nov. Gen. &
Sp. 3: 21. 1845.
Slender shrub or tree sometimes attaining 20 meters, the younger
branchlets early more or less villous or appressed pubescent as often
the leaves beneath or these glabrate; petioles 1-2 (5) cm. long, stipules
5-6 mm. long, linear-setaceous, rigid; leaves ovate to lanceolate, base
obtuse or sometimes narrowed to acute or subcordate base, acuminate
or cuspidate, often 7-15 cm. long, 2-5 cm. wide or larger, dentate,
firm-membranous, 6-9 pinnately nerved; flowers monoecious, the
axillary spikes male or androgynous with 1-2 (5) basal female or
rarely all female, 5-11 cm. long, 2-4 mm. wide, sessile or nearly,
male part dense, later laxly flowered; female bracts ovate, obscurely
dentate, 1-3-flowered, small, the fruit 4-6 mm. wide; sepals broadly
ovate, acute, hirsute; ovary muricate and hispid; styles about 3 mm.
long, pinnately 10-20-lacinulate; capsules nearly 3 mm. thick; seeds
1.5 mm. long, minutely puncticulate. Plants with villous leaves
have been named var. leptostachya (HBK.) M. Arg. in DC. Prodr.
15, pt. 2: 854. 1866, and those with glabrate leaves or the nerves
pilose, var. carpinifolia (Poepp.) M. Arg. I.e. 854. Some of the
following material, incomplete, may be misdetermined, especially
from Junin, which seems mostly var. leptostachya and simulates
A. Ruiziana, but Diels, Bibl. Bot. 116: 104. 1937, noted leaf-form
and pubescence combined otherwise than by Mueller in Ecuador
specimens, which he placed here.
130 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
San Martin: Tarapoto, Mathews 1557; Spruce 4033; 4142;
Williams 6025; 5490 (det. Mansfeld, A. samydaefolia). San Roque,
Williams 7059. Zepelacio, King 3369; 37 51 (det. Standley). Juanjui,
Klug3791 (det. Standley). Huanuco: Pozuzo, 4564 (det. Johnston).
Cuchero, Poeppig (type, A. microgyna). Monzon, Weberbauer 3428.
Junin: La Merced, 5311 (det. Johnston); Killip & Smith 23742;
23533; 24080. San Ramon, Schunke 115; 165; 276; Killip & Smith
24549; 24740. Colonia Perene*, Killip & Smith 25052; 2521 7. Pichis
Trail, Killip & Smith 25887. Loreto: Lower Huallaga, Williams
4391; 4756. Yurimaguas, Williams 4470; Killip & Smith 27663;
28063; 28216. Puerto Arturo, Killip & Smith 27712; 27897. La
Victoria, Williams 2826. Mexico to Bolivia, Brazil and Venezuela;
Guiana. "Yana-varilla" (Williams).
Acalypha Forbesii Sp. Moore, Journ. Bot. 52: 336. 1914; 26.
Stems early densely white pubescent as the petioles, these slender,
about 12 mm. long, and the leaves beneath especially on the nerves;
leaves ovate, rounded at base, obtuse, often shortly 3-lobed, about
3 cm. long, 2 cm. wide, crenate, membranous, sparsely strigillose
above; spikes terminal and axillary, all androgynous, subsessile, the
terminal about 4 cm. long, 8-9 mm. broad, the male part scarcely
3 mm. long, deflexing, the densely bracted female portion oblong;
axillary spikes about 12 mm. long; female bracts becoming 5-6 mm.
broad, 3-5-lobed medially or a little deeper, the lanceolate-oblong
obtuse lobes entire or unidentate, membranous, pubescent without;
ovary shortly hispidulous, styles few-lacinulate, not colored; capsules
trilobed, about 2 mm. in diameter, the seeds 1 mm. long. The related
and widely distributed (at least as a weed) A. Poiretii Spreng., for
which the proper name may be A. alnifolia Poir., has larger leaves,
slender spikes and simple styles; the similar but like A. Forbesii
imperfectly known A. paupercula Pax & Hoffm., 29, of Bolivia is
ligneous at base, has female part of spikes ovoid, the bracts 5-9-
dentate, or spikes all male, ovary long-pilose. A. boliviensis M. Arg.
has the male spikes axillary at base of the terminal female, styles
simple or 3-4-parted.
Lima(?): Valley between Pacasmayo and Railhead, (Forbes,
type).
Acalypha fulva Johnst. Contr. Gray Herb. 75: 29. 1925.
Pungently resinous shrub about 5 dm. tall, the bracts, petioles,
these 5-15 cm. long, and leaf -nerves beneath villous-hirsute with
FLORA OF PERU 131
more or less glandular simple setae, these sometimes 2-3 mm. long;
stipules reflexed, persisting, lanceolate-subulate, subscarious, gla-
brate, 6-8 mm. long; leaves ovate-acuminate, deeply cordate at base
with imbricate lobes, 12-36 cm. long, 9-18 cm. wide, green, villous,
sparsely stipitate glandular and inconspicuously punctate above,
serrate; male spikes axillary, 6-15 cm. long, glandular pubescent,
about 1 cm. above base with 1 or 2 cordate-ovate sessile 1-flowered
female bracts 20-30-dentate, 10-12 mm. long, 11-16 mm. wide;
female sepals oblong-ovate, glandular-ciliate; ovary densely stipitate
glandular; styles 7-9 mm. long, deeply 10-15-lacinulate. Belongs
to group Olygogynae-Cuspidatae, 114, but according to Johnston
readily separated from other members of this section by the glandular
pubescence and deeply cordate leaves with closed sinus, and thus
allied to A. plicata and A. reflexa within Peru.
Huanuco: Muiia, 2,000 meters, 4011, type.
Acalypha infesta Poepp. in Poepp. & Endl. Nov. Gen. & Sp.
3: 22. 1845; 48.
Annual or perhaps sometimes perennial, the branching stems to
5 cm. high, puberulent or above hirsute with retrorse trichomes as
the 0.5-3 cm. long petioles; leaves broadly ovate, rounded, obtuse
or slightly cordate at base, obtusish, 2-5 cm. long, 1.5-4 cm. wide,
crenate-serrate, thin-membranous, early sparsely pilose, basally
3-nerved; flowers monoecious, the male spikes axillary, shorter than
the petioles, the female terminal and 1-several in the leaf-axils, the
former in fruit to 4 cm. long, 1.5 cm. broad, hirsute, the bracts then
6-8 mm. long, 3-5-lobed at least medially, the oblong lobes obtuse
or acute, the terminal sometimes smaller, shortly hirsute, ciliate,
often glandular, 2-flowered; sepals ovate, acute, ciliate; ovary apically
hirsute, muricate or nearly smooth; styles 2-4 mm. long, slenderly
2-5-lacinulate; seeds about 1 mm. long, minutely puncticulate.
Name written infestans by Mueller, Linnaea 34: 23. 1865. The var.
rotundifolia (Vahl ex Baill.) M. Arg. is the form with lobes of female
bracts rounded-obtuse, eglandular, var. stenoloba M. Arg. (typical
form), the bract lobes sometimes acute, always glandular-ciliate. A
weedy annual sometimes rooting at the often decumbent-ascending
base. The Ecuadorean A. stellipila Pax & Hoffm., 49, said to be
related, seems to be recognizable at once by the stellate pubescence
and shrubby habit. F.M. Negs. 32501; 5295 (var.).
Lima: Matucana, sandy gravel along river, 31 9 (det. Johnston).
Huanuco: Near Cuchero, Poeppig 1701 type (and type var. stenoloba}.
132 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Junin: La Merced, ditch bank, 5286 (det. Johnston). Apurimac:
Oropeza Valley, Vargas 9785 (det. Standley). Casinchihua, Prov.
Abancay, Goodspeed Exped. 10583 (det. Standley). Without locality,
Ruiz & Pavdn; Dombey. Ecuador.
Acalypha Macbridei Johnst. Contr. Gray Herb. 75: 28. 1925.
Branchlets terete, the younger as the petioles, these 5-8 mm.
long, densely canescent strigose; stipules lanceolate, deciduous, about
2 mm. long; leaves broadly oblanceolate or oblong-lanceolate, obtuse
at base, attenuate-acuminate, 15-27 cm. long, 2.5-5.5 cm. wide,
membranous, obsoletely crenate, asperulous above, sparsely ap-
pressed villous and paler beneath with the 15-25 primary veins
prominent; male racemes unknown; female axillary, filiform, 8-16
cm. long, spiciform (but the flowers scattered), shortly peduncled,
strigose; bracts minute, ovate-lanceolate, 1-3-flowered, the flowers
scarcely 1 mm. long; pedicels slender; sepals 5, ovate, acute, glan-
dular-dentate; ovary densely papillose; styles about 2.5 mm. long,
15-20-lacinulate. Apparently nearest A. subandina Ule with longer
petioles and paniculate inflorescence (Johnston) and the almost
filiform caudation is sometimes nearly 3 cm. long. Type from a lax
shrub about 9 dm. high; arborescent, 5 meters high (Mexia). A.
salicifolia M. Arg., 19, of Ecuador has linear-lanceolate leaves,
12 cm. long, 12-18 mm. wide, minutely puberulent beneath only on
nerves, female pedicels 1.5 mm. long; otherwise seemingly similar
but male flowers unknown.
Junin: La Merced, Killip & Smith 23525. Pichis Trail, Killip
& Smith 25428. Huanuco: Pampayacu, 5093, type. Loreto: Above
Pongo de Manseriche in tangle on steep banks, Mexia 6213 (det.
Standley).
Acalypha macrodonta M. Arg. Linnaea 34: 51.1865; 85.
Tall shrub, the branches and leaves beneath sparsely pubescent;
petioles early short, soon 1-4 cm. long; leaves ovate, lightly cordate,
acuminate, 6-12 cm. long, about half as wide, or larger, coriaceous-
membranous, serrate; flowers dioecious, the male unknown; terminal
sessile female spikes dense, shorter than the leaves, the ovate-
lanceolate 1 -flowered bracts 5-7-dentate both sides below the tip,
this tooth or terminal part 3-4 times longer than the rest, sparsely
stipitate glandular; sepals ovate, long-acuminate; ovary hirsute;
styles elongate, shortly 10-12-lacinulate their entire length with
2-3-parted laciniae, sparsely pilose. A. stellipila Pax & Hoffm.,
FLORA OF PERU 133
49, of Ecuador with the pubescence of A. dictyoneura, that however
apparently with axillary inflorescence, would be sought here. When
the type of A. macrodonta can be re-examined it may prove to be
the correct name for A. mapirensis; compare also A. Lechleri noted
under the latter. F.M. Neg. 8501.
Ayacucho: Rio Apurimac Valley near Kimpitiriki, Killip &
Smith 22947 (det. in herb. A. macrophylla, i.e. A. stachyura). Peru(?) :
Pavdn, without other date, type, herb. Delessert (Boissier).
Acalypha macrostachya Jacq. Hort. Schoenbr. 2: 63. pi. 245.
1797; 144. A. sidaefolia HBK. Nov. Gen. & Sp. 2: 95. 1817. A.
macrophylla HBK. ex Poepp. in Poepp. & Endl. Nov. Gen. & Sp.
3: 22. 1845. A. tarapotensis M. Arg. in DC. Prodr. 15, pt. 2: 808.
1866. A. tristis Poepp. in Poepp. & Endl. I.e.
A tall shrub or slender tree rarely attaining 12 meters, the stout
branchlets and leaves hirsute-villous in varying degree to nearly
glabrous; stipules basally cordate, obtuse, linear-lanceolate, glandular,
1-1.5 cm. long, persistent; petioles 5-20 cm. long or longer; leaves
ovate, often broadly, obtuse or cordate at base, acuminate, usually
a dm. or two long, 6-17 cm. wide, serrate, membranous, palmately
nerved; flowers apparently monoecious in long (sometimes 4 dm.)
axillary sessile or shortly peduncled spikes, the male dense, to 5 mm.
wide, the rather laxly flowered female sometimes with a few male at
tip or sometimes the upper half male; female bracts broadly ovate
or truncate-obtuse, 13-27-dentate, one-fifth to one-third incised,
1-flowered, the teeth acuminate, frequently gland-tipped, the fruiting
5-7 mm. long; sepals broadly ovate, acute, glandular; ovary hispid;
styles 4-5 mm. long, subpinnately 10-20-lacinulate, dorsally pilose;
capsules nearly 4 mm. thick, pilose, the puncticulate seeds 2 mm.
long. Herbarium materials have been sorted into three variants
according to pubescence: hirsutissima (Willd.) M. Arg., macrophylla
(HBK.) M. Arg. (this with leaves glabrate except nerves), and
tristis (Poepp.) M. Arg., entirely glabrate except for apical puberu-
lence. Since, apparently, these are not geographically distinct, the
sorting appears to be entirely artificial. The last two were treated
as formas by Mueller Argoviensis in DeCandolle's Prodromus, I.e.
810-811. The similar more northern A. heterodonta M. Arg., 146,
has ovate leaves, styles about 7 mm. long and is probably a sub-
species. F.M. Neg. 7134 (S. tarapotensis).
Huanuco: Huamalies, Weberbauer 3614; 286 (cf. A. tarapotensis).
Cuchero, Ruiz & Pavdn; Poeppig (det. Poeppig, A. sidaefolia).
134 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Pampayacu, mouth of Rio Chinchao, 5025 (det. Johnston). Tingo
Maria, Soukup 2119 (det. Croizat). San Martin: Tarapoto, Spruce
4294 (type, A. tarapotensis); Williams 6313; 6765. San Roque,
Williams 7704. Zepelacio near Moyobamba, Klug 3365 (det.
Standley). Junin: Chanchamayo Valley, Schunke 426; 486; Isern
2132. Hacienda Schunke, Schunke A109. La Merced, Killip &
Smith 23405; 23441. Near Perene" Bridge, Killip & Smith 25398
(var. tristis, det. Mansfeld). Loreto: La Victoria, Williams 2822.
Yurimaguas, Williams 3912; 4838 (var. tristis, det. Mansfeld);
Poeppig 2192; 3088 (type, A. tristis}. Puerto Arturo, Killip &
Smith 27756; Williams 5124- Pucallpa, Soukup 3050. Iquitos and
Mishuyacu, Williams 8055; Klug 1464- Cuzco: Valle del Urubamba,
Machu-picchu, 2,200 meters, Herrera 3228; 3237; 3242 (all det.
Standley). Potrero, Rio Chuyapi, Vargas 8249. Cadena, Vargas
6125. Rio Acre: Seringal Auristella, Ule 6536.
Guianas to Mexico, Bolivia and Brazil. "Yana-vara" or "varilla"
(Williams), "pespita" (Herrera).
Acalypha mapirensis Pax, Repert. Nov. Sp. 7: 110. 1909; 65.
Branchlets shortly pubescent apically becoming glabrate as the
petioles, these 3-13 cm. long; leaves broadly ovate or oblong, rounded-
cordate or obtuse at base, acuminate, 12-25 cm. long, 5-15 cm. wide,
serrate-dentate, membranous, early sparsely pubescent especially
beneath on the nerves, the basal 3-5; flowers monoecious, the dense
spikes sessile or shortly peduncled, the axillary male 3-6 cm. long,
2-3 mm. thick, the terminal female in flower 8-15 cm. long, tomen-
tose, their bracts about 6-7 mm. broad, strongly nerved, 1-flowered,
slightly if at all glandular, one-fourth or one-third 9-15-dentate,
teeth linear, the medial often 2-3 times longer than the others;
sepals ovate, acuminate; ovary verrucose, pilose; styles 5-10 mm.
long, shortly and finely 6-15-lacinulate; capsules pilose; seeds
subglobose, 1 mm. thick, nearly smooth. Var. pubescens Pax &
Hoffm. and var. scabra Pax & Hoffm. designate two types of leaf-
pubescence, the former the typical. Compare A. macrodonta, prob-
ably the earlier name. A. Lechleri Britton, 65 (type, Lechler 2408
without data), Bolivian or possibly from Peru may belong here but
the flowers are described as dioecious with styles only 2 mm. long.
F.M. Neg. 5301.
Cuzco: Prov. Quispicanchi, 800 meters, Marin 1551. Puno:
Chunchusmayo, Sandia, Weberbauer 1174 (var. scabra). Rio Acre:
Seringal San Francisco, Ule 9535; 9535b (last, var. scabra). Mouth
of Rio Macauhan, Krukoff 5440 (det. Steyermark). Bolivia.
FLORA OF PERU 135
Acalypha obovata Benth. Bot. Voy. Sulphur 163. pi. 53. 1844;
163. A. cuneata Poepp. in Poepp. & Endl. Nov. Gen. & Sp. 3: 22.
1845; 163.
Tall glabrous shrub or the branchlets and leaves beneath on the
nerves more or less hirtellous; stipules lanceolate-subulate with
thick midnerve, 7-9 mm. long; petioles 1-4 cm. long; leaves oblong-
obovate, long-cuneate to the slightly cordate base, more or less
abruptly caudate-acuminate, often narrowly, cuspidate and mucronu-
late, 1-2.5 dm. long, or longer, 3-9 cm. wide, firm -membranous,
crenate or callous-denticulate except toward base; flowers monoecious
in sessile axillary dense elongate spikes, the upper female, the lower
male, these 12-15 cm. long; fruiting bracts broadly reniform, nearly
5 mm. wide, apically subrecurved, acute, slightly 11-15-denticulate
(apparently also fewer), 1-flowered; sepals broadly ovate, acuminate;
ovary hispid and very muricate; styles rigid with about 20 laciniae
entire length, dorsally pilose and asperous; capsules 6 mm. broad;
seeds smooth. A. juruana Ule seems near but the female bracts
are 7-9-denticulate, petioles 4-10 cm. long and to it was referred
a collection by Rospigliosi accredited to "Lima" which must have
been obtained on the eastern side of the Andes. The common form
seems to be var. cuneata (Poepp.) Macbr., Candollea 8: 26. 1940,
according to Mueller the glabrous state, but I have not seen Ben-
tham's type. Common in the region of Iquitos, only a few collec-
tions cited. Krukoff 5249 has been referred here, but female bracts
not seen. F.M. Neg. 5288.
Huanuco: Rio Cayumba, Mexia 8269 (det. Steyermark). Loreto:
Yurimaguas, Poeppig, type (numbered 2230; 2317; 2807); Killip
6 Smith 28195; Williams 3866. Puerto Arturo, Williams 5134.
Fortaleza, Williams 4271. Lower Huallaga, Williams 5134. Santa
Rosa, Killip & Smith 28888; 28859. Iquitos, King 1400. San
Antonio, Rio Itaya, Killip & Smith 29438. La Victoria, Williams
2767. Rio Acre: Mouth of Rio Macauhan, Krukoff 5249. To
Venezuela and Colombia. "Sapote yacu" (Williams).
Acalypha padifolia HBK. Nov. Gen. & Sp. 2: 97. 1817; 62.
Shrub-tree to about 4 meters high, the terete branchlets, petioles
(4-10 mm. long), leaves beneath and male spikes more or less villous-
tomentose; leaves lanceolate-ovate, obtuse and slightly cordate at
base, acute, 4-8 cm. long, 1.5-3.5 cm. wide, firm-membranous,
serrations recurved, sparsely pubescent and scabrous above, 3-nerved;
flowers monoecious, the known axillary male spikes 8 cm. long,
136 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
slender, laxly flowered, the fruiting terminal 5-6 cm. long, compressed
12 mm. broad, dense, the suborbicular bracts acute or acuminate in
fruit, about 12 mm. wide, 1-flowered, strongly nerved, stipitate-
glandular beneath and marginally, 17-19-dentate, the triangular
acute teeth recurved; sepals ovate, acute, ciliate; ovary hirsute;
styles 7-8 mm. long, pilose, pectinately 10-16-lacinulate. A.
Schimpfii Diels, Bibl. Bot. 116: 103. 1937, of Ecuador has bracts
about 12-dentate but otherwise ex char, seems to be similar, as also
A. eugeniifolia Rusby, 62, of northern Bolivia, but the bracts 7-8-
parted. There seem to be connecting Ecuadorean forms or species:
A. andina M. Arg. and A. ecuadorica Pax & Hoffm. both with
membranous leaves, the former more pubescent, rounded-cordate
at base, the latter obtuse or acute, the female bracts stiped-glandular.
There is further A. tunguraguae Pax & Hoffm., scarcely distinguish-
able unless the leaves somewhat bullate above, more pubescent, the
petioles 1-3 cm. long, and A. controversa (Ktze.) K. Schum., 68, type
from Bolivia, which is apparently a large-bracted species com-
parable to A. peruviana but pubescent. My collection is very young,
the bracts not clearly dentate below but undeveloped; as the leaves
are densely villous-tomentose beneath it could be A. andina or maybe
A. tunguraguae if these are specifically distinct. As remarked by
Diels, Bibl. Bot. 116: 103. 1937, the species definition in this group
is not yet determined; maybe A. padifolia is a basic unit with the
central acumination of the female bracts not produced, as more or
less in the following material. My specimen was from an open
shrub, 1 meter tall; Stork and Horton from trees or shrubs 5 meters
tall with cane-like stems forming a tangle with each other and other
shrubs.
Cajamarca: Above Socota, 2,800 meters, Stork & Horton 10148
(det. Standley, A. controversa). Junin: Huacapistana, 5824 (det.
Johnston, A. controversa). Loreto: Rio Maranon Valley, Dennis
29132 (det. Croizat, with query). Ecuador.
Acalypha peruviana M. Arg. Linnaea 34: 17. 1865; 68.
Branches ligneous, the younger parts densely appressed pubescent
with pale fulvous trichomes, the branchlets and leaves soon glabrous;
petioles 2-6 cm. long; leaves broadly ovate, obtuse at base, acumi-
nate, 9-17 cm. long, 4-12 cm. wide, serrulate except toward sub-
5-nerved base, chartaceous; secondary nerves with veins prominent
both sides; sessile or subsessile axillary male spikes 6-18 cm. long,
dense, about 3 mm. broad, the terminal peduncled fruiting ones
FLORA OF PERU 137
nearly equaling the leaves and the bracts unusually large, said to
attain 2 cm., 8 mm. wide, ovate, acuminate, entire but (except at
tip) densely glandular pilose toward and on margins; sepals broadly
ovate, acuminate, ciliate; ovary shortly pilose; styles 4-5 mm.
long, shortly many-lacinulate; fruit unknown. There are several
similar Bolivian species or variants, some with tomentose leaves
as A. controversa (Ktze.) K. Schum. and more or less dentate, glan-
dular or eglandular but large bracts; cf. A. padifolia. F.M. Neg.
5308.
Huanuco: Compact tree-shrub with spreading branchlets, Muna,
4029; Ruiz & Pavdn, type.
Acalypha platyphylla M. Arg. Linnaea 34: 6. 1865; 20.
Shrub with angled early appressed puberulent branchlets, this
indument also on the 9-13 nerves of the younger leaves; stipules
subulate; petioles 2-several cm. long; leaves elliptic-lanceolate,
obtuse at base, 17-20 cm. long, 7-9 cm. wide, obtusely crenate-
dentate, rigid-membranous, paler beneath; female panicles axillary,
ample, erect, laxly flowered, finally equaling the leaves, the small
oblong-ovate acuminate bracts 1-3-flowered, the flowering pedicels
equaling the calyx, in fruit 7 mm. long, the 5 ovate-lanceolate
sepals acuminate; ovary minutely muricate-subhirtellous; styles
2-2.5 mm. long, divergently 30-40-lacinulate from the narrowly
lanceolate base, the short laciniae subgeminate or ternate. A
Colombian specimen thought to belong here but entirely male has
slender subsessile densely flowered spikes 5-15 cm. long (Pax and
Hoffmann); see also A. subandina. F.M. Negs. 7135; 32505.
Peru (probably). Ecuador; Colombia?
Acalypha plicata M. Arg. in DC. Prodr. 15, pt. 2: 855. 1866;
121. A. flabellifera Rusby, Mem. Torrey Club 6: 119. 1896? A.
cordifolia Griseb. Goett. Abh. 24: 59. 1879, not Hook. f. Trans.
Linn. Soc. 20: 186. 1847.
Densely leafy flexuose branchlets obscurely glandular pubescent
with short trichomes, glabrescent in age; stipules linear, subulate-
acuminate, membranous; petioles 1.5-2.5 cm. long, densely pilose
as the leaf -margins and female bracts with gland-tipped trichomes;
leaves oblong-triangular-ovate, rounded-cordate at base, long-acumi-
nate, 6-8.5 (15) cm. long, 3-4.5 (10) cm. wide or becoming larger,
5-plinerved, firm-membranous, denticulate, paler beneath, the
younger villous-sericeous, the trichomes above in part rigid, beneath
138 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
sometimes entirely subpulveraceous asperous with enlarged tri-
chomes; spikes elongate, finally as long as or longer than the leaves
(male part 2.75 mm. wide), a naked area between the female and
male bracts, the former basal, 1-flowered, broadly reniform, obtuse,
typically entire with prominently plicate nerves, fruiting to 14 mm.
wide, 8 mm. long, cyathiform-obconic, subtruncate, glandular;
female calyx segments broadly ovate; ovary pilose and slightly
muricate above; styles shortly 5-7-lacinulate; seeds 2 mm. long.
Habit of A. cuspidata Jacq., 120, which reaches southern Ecuador
from the West Indies and perhaps is only a more glandular state.
Lourteig & O'Donell, Lilloa 8: 312. 1942, regard the plant of Rusby
(that is Grisebach, not Hooker, f.) as distinct, the female bracts
minutely many-denticulate; Pax and Hoffmann questioned the
identity. The problem of course (curiously ignored) concerns the
validity of bract-dentation and glandulosity as indicators of species
or variants, as varieties or subspecies. Incidentally, in some moment
of aberration, surely no less, I created the superfluous name A.
Hookeri, Field Mus. Bot. 11 : 26. 1931, for the earlier described plant
of Hooker, f.
Cuzco: Quillabamba to Echarate, Vargas 7530. Near Cuzco,
Vargas 1713 (det. Croizat). Potrero, Rio Chuyapi, Vargas 8248.
To Paraguay and Argentina.
Acalypha reflexa M. Arg. Linnaea 34: 33. 1865; 123.
Branchlets many, fulvous villosulous as the petioles, these finally
nearly as long as the orbicular to elliptic or ovate blades of the
membranous leaves, these subcordate at base, rounded to the obtuse
or acutish apex, 3.5-7 cm. long, 3-5 cm. wide, the marginal dentations
oblique and subconcave dorsally below the tips; stipules 6-7 mm.
long, subulate-acuminate, margins hyaline, stipitate-glandular; male
spikes 4-6 cm. long, dense or below interruptedly glomerulate;
female flowers 1-2, the broadly ovate bracts cordate at base, rounded-
obtuse, 6 mm. long, about 25-dentate (teeth refracted), villous with-
out, pilose within, radiately more or less plicate, 1-flowered; sepals
ovate, acuminate; ovary fulvous sericeous; styles rigid, short,
pectinately about 16 lacinulate. Female inflorescence probably not
normal (M. Arg.). F.M. Neg. 7144.
Peru(?): Without locality, Ruiz & Pav6n, type, herb. Boiss.
Acalypha Ruiziana M. Arg. Linnaea 34: 16. 1865; 67.
Ligneous branches lenticellate, fulvous-villosulous apically;
petioles to 5 mm. long; leaves lanceolate or oblong-ovate, base
FLORA OF PERU 139
narrowly obtuse or acute, acuminate, 2.5-4.5 (14) cm. long, 12-20
mm. (or 3-5.5 cm.) wide, coriaceous, opaque, pubescent only on
pinnate nerves, these impressed above, prominent beneath (3-nerved
at base), glabrescent, dentations early revolute beneath; axillary
male spikes 5-12 cm. long, 4 mm. wide, compressed, sessile finally
laxly flowered; sessile or subsessile terminal female spikes 2-4 cm.
long, the congested flowering bracts 2-3 mm. long, in type 5-9-
dentate both margins, long-acuminate, the acumen twice as long as
triangular basal part, entire or few-glandular, sparsely hirsute,
1-flowered; sepals oblong, exceeding pubescent ovary, pubescent
and few-glandular; styles rather rigid, pectinately 12-15-lacinulate.
The incompletely known A. carthagenensis Jacq., 67, of Colombia
may prove to be the earlier name but scarcely on account of the
separation in range. The larger leaf -measurements apply to the
specimen of Soukup. F.M. Neg. 5316.
Huanuco: Muna, Ruiz & Pavdn, type. Junin: Vitoc, Soukup
2448. Chanchamayo, Isern 2388.
Acalypha samydaefolia Poepp. in Poepp. & Endl. Nov. Gen.
& Sp. 3: 21. pi. 224. 1845; 107.
Shrub sometimes 10 meters high, the slender branchlets and
petioles, these 3-15 mm. long, early appressed pubescent; stipules
linear-setaceous, rigid, 5-7 mm. long; leaves lanceolate or ovate,
obtuse at 3-nerved base, narrowly cuspidate-acuminate and mucro-
nate, 4-14 cm. long, 2-6 cm. wide, lightly serrate, firm-membranous,
glabrous above, glabrous or sparsely pilose on the 5-7 nerves beneath;
flowers monoecious, the slender sessile axillary spikes 2-3.5 (rarely
to 10) cm. long, entirely male or 1-4 female bracts at base or these
solitary in the leaf-axils, about 4 mm. broad, orbicular-reniform,
rotund or shortly acute, shortly and obtusely about 12-dentate,
coriaceous, 1-2-flowered; female sepals ovate, acute, ciliate; ovary
densely muricate and pilose; styles about 2 mm. long, subpalmately
to 10-lacinulate at base; capsules 3 mm. broad, seeds said to be
smooth. Not clearly distinct from A. diver sifolia unless the seeds
are smooth. It is noteworthy that the authors described the female
bracts as denticulate while according to Pax and Hoffmann they
are entire; I doubt that the species is distinct from A. diver sifolia.
F.M. Neg. 5318.
Loreto: Yurimaguas, Poeppig 2122, type; Spruce 3910 (det.
Pax). Rio Acre: Seringal San Francisco, Ule 9532 (det. Pax).
Southern Brazil to Venezuela and Colombia.
140 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Acalypha scandens Benth. in Hook. Journ. Bot. Misc. 6: 329.
1854; 147.
High climbing, the tips of the lightly angled branches often
densely fulvous-hirsute with more or less spreading trichomes or
these sometimes appressed or soon deciduous; stipules narrowly
lanceolate, acuminate, typically glandular, 6-8 mm. long; petioles
2-10 cm. long; leaves elliptic-ovate but as interpreted here variable,
rounded-cordate to acute at base, apiculate to well acuminate, 8-16
cm. long, 5-9 cm. wide or smaller, especially narrower, coarsely
serrate to subentire, membranous, basally 5-nerved, rather harshly
scabrous-hirsute above, ordinarily glabrate or scabrous-punctate
beneath; flowers monoecious, the shortly peduncled early densely
flowered male spikes to 2 dm. long, the female to 3.5 (10) dm. long,
lax, slender, sometimes with a few male flowers at tip and always
conspicuous after anthesis by the coral-red styles about 1 cm. long
with many entire or parted lacinulae; fruiting bracts to 3.5 mm.
long, ovate, hirsute, 11-19-dentate to the middle (or shortly in Peru)
with linear gland-tipped teeth; ovary strigose and muricate; capsules
nearly 3 mm. in diameter; seeds smooth, 2 mm. long. Some of the
material cited has glabrous shortly dentate bracts and may not
belong here. F.M. Neg. 5317.
Huanuco: Tingo Maria, Stork & Horton 9475 (det. Standley).
Loreto: Fundo Indiana near Rio Maranon, Mexia 6386 (det. Stand-
ley). Rio Mazan, Jose Schunke 22 (det. Croizat with query). Yuri-
maguas, Williams 4521. Mishuyacu, Klug 1070 (det. Mansfeld).
Rio Itaya above Iquitos, Killip & Smith 29541- Leticia, Wittiams
3170. Lower Rio Nanay, Wittiams 591 (leaves oblong-lanceolate,
about 10x3 cm., det. Mansfeld). Caballo-Cocha, Williams 2311;
2179. Balsapuerto, Killip & Smith 28491. To the Guianas.
Acalypha stachyura Pax, Repert. Nov. Sp. 7: 110. 1909; 65.
A. macrophylla Ule, Verh. Bot. Ver. Brandenb. 50: 79. 1908, not
HBK. or Poeppig.
Glabrous (or the leaves early puberulent) except the puberulent
sulcate branchlet tips and the inflorescences; petioles in age 4-16 cm.
long; leaves oblong, obtuse or rounded at base, caudate-acuminate,
1.5-3 dm. long, 5-11.5 (15) cm. wide, serrate dentate, membranous,
pinnately nerved with 8-10 secondary nerves, the lowest basal or
nearly; flowers monoecious, the slender dense axillary male spikes
6-15 cm. long, the terminal female in flower about 10, in fruit to
25 cm. long, pedunculate; sometimes the female spikes are male
FLORA OF PERU 141
toward apex for about 3 cm.; fruiting bracts finally subglabrous,
to 1.5 cm. long, eglandular, 1-flowered, nearly medially 5-10 dentate,
the middle of the triangular teeth the larger; sepals acute, pulveru-
lent; ovary tomentulose; styles 5-10 mm. long, pinnately 10-15
lacinulate; seeds 2.5 mm. long, smooth.
The species (a slender tree, the bark abundantly lenticellate,
Williams, Field. Mus. Bot. 15: 266. 1936) seems fairly well marked
by the minute distinct leaf-dentation, but this is more crenate and
more approximate in the type of A. stachyura so the Peruvian plant
may not be the same as indicated by the monographers; however,
if another name is required for the plant of Peru there will probably
prove to be an available synonym, for instance A. stenoloba (which
compare) or A. Lechleri Britton, 65, the type without data said to
be distinguishable by its dioecious flowers, styles only 2 mm. long,
the first character probably unimportant and the second perhaps an
error, typographical or in observation. Apparently better defined is
A. grandispicata Britton, 65, also Bolivian with female spikes to 2.5
cm. wide, and 14-16 secondary nerves. A. stellipila Pax & Hoffm.,
49, Ecuadorean, has the pubescence of A. dictyoneura with apparently
axillary spikes. F.M. Neg. 5299.
San Martin: Tarapoto, Ule 6656 (type, A. macrophylla). San
Roque, Williams 7037 (det. A. benensis). Huanuco: Near Tingo
Maria, 2,400 meters, Seibert 2254 (det. A. C. Smith). Junin: San
Nicolas, Kittip & Smith 26109; 26026 (det. A. benensis). Dos de
Mayo, Killip & Smith 25836. Enenas, Kittip & Smith 25787 (det.
Croizat, A. Lechleri}. Chanchamayo Valley, Schunke 449; 451; 558;
562 (all det. A. benensis). Hacienda Schunke, 5651; Kittip & Smith
24641. Lore to: Florida, Klug 2165 (det. Standley, A. benensis;
Croizat, A. stachyura). Santa Rosa, Killip & Smith 28975. Forta-
leza, Williams 4184- Puerto Arturo, Wittiams 5185; 5251. Bolivia.
"Yana ocuera de oyada" (Williams), "ucutu f enana" (Klug, Huitoto).
Acalypha stenoloba M. Arg. in Flora 55: 41. 1872; 61. A.
capillaris Rusby, Mem. Torrey Club 4: 257. 1895. A. Baenitzii Pax,
Repert. Nov. Sp. 5: 227. 1908. A. ovata Pax & Hoffm. Meded.
Rijks Herb. Leiden 40: 23. 1921, fide Pax and Hoffmann as other
synonyms.
Shrub or small tree 4-5 meters high, the ligneous branchlets as
the remains of younger parts minutely pulverulent; petioles 1-7 cm.
long; leaves oblong- or lanceolate-ovate, cuneate to obtuse base,
cuspidate-acuminate, 9-15 cm. long, 2-7 cm. wide, sharply serrate,
142 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
nearly glabrous but barbate beneath in the axils of the 5-6 nerves,
rigid-membranous, pinnately nerved; flowers monoecious (always?),
the axillary male spikes sessile, 1-6 cm. long, the terminal female
6-12 cm. long, dense, shortly peduncled with flowering bracts 5-6
mm. long, deeply 5-9-parted, lobes linear-subsetaceous, gradually
long-acuminate, not papillose-asperous, fruiting to 8 mm. long,
rigid, 1-flowered, the female sepals ovate or lanceolate, acute or
acuminate, ciliate; ovary puberulent-pilose, finally verrucose; styles
4-8 mm. long, pinnately 7-15-lacinulate; capsules 3-4 mm. thick,
the subglobose seeds smooth, nearly 2 mm. thick. A. lucida Rusby,
61, probably the same, was thought to have dioecious flowers;
A. urostachya Baill. 60, country unknown but related here, is rather
tomentose and the 1-flowered auriculate female bracts are unequally
parted. This may well be an earlier name for A. stachyura. F.M.
Neg. 19517.
Cuzco: Idma, Prov. Convention, 1,500 meters, Vargas 8559.
Without locality, Haenke, type. Bolivia.
Acalypha stricta Poepp. in Poepp. & Endl. Nov. Gen. & Sp.
3: 21. pi. 225. 1845; 64.
Branchlets, petioles, these 1-4 cm. long, and leaves beneath
densely tomentose, the latter sparsely so above, ovate or oblong-
ovate, rounded-cordate at base, acuminate, a dm. or two long,
about half as wide, membranous or chartaceous, 3-5-nerved at base,
above pinnately; flowers monoecious, the axillary male spikes 5-10
cm. long, 3-5 mm. thick, sessile or nearly, dense, erect, the terminal
female 7-20 cm. long with flowering bracts 2-3, fruiting 7-8 mm.
wide, tomentose, 1-flowered, medially 13-15-dentate with linear
setose subequal teeth, the ovate acute ciliate sepals sparsely glan-
dular; ovary villous; styles 2-3 mm. long, pectinately 10-16-lacinu-
late; seeds ovoid, minutely puncticulate, 1.5 mm. long. My speci-
men from type locality from a compact shrub 1-2 meters high. F.M.
Neg. 7128 (Pavon).
Huanuco: Hacienda Exito, bank of Rio Ysabel, Mexia 8116 (det.
Standley, A. macrostachya). Pampayacu, 5051 (det. Johnston, A.
mapirensis); Poeppig, type. Puente Durand, Stork & Horton 9440
(det. Standley, A. benensis). Junin: Huacapistana, 1,600 meters,
Weberbauer 2329. Chanchamayo Valley, Schunke 323; 411; 1732;
1799; Isern 2391. La Merced, 5538 (det. Johnston, A. mapirensis);
Soukup 1524 (det. Ewan, A. macrophylla}; Killip & Smith 23744;
23743. Colonia Perene", Killip & Smith 25018; 25080. San Ramon,
FLORA OP PERU 143
Kittip & Smith 24749. Along Rio Perene*, Killip & Smith 25123;
25189 (both det. Standley, A. mapirensis). Rio Acre: Seringal
San Francisco, Ule 9533.
Acalypha subandina Ule, Verb. Bot. Ver. Brandenb. 50: 77.
1908; 20.
Similar to A. platyphylla but like it the male flowers unknown;
petioles 2-6 cm. long, minutely or obsoletely biglandular; leaves
subobovate-oblong, obtuse and lightly cordate at base, 13-22 cm.
long, 6-8.5 cm. wide, glabrous or nearly from the first except scaber-
ulous beneath; panicles 13-20 cm. long, diffuse, puberulent, the
lanceolate bracts scarcely 1 mm. long, the pedicels attaining 1-2
cm., the sepals pilose; ovary muricate; styles 2.5-4 mm. long, rather
stout below, 10-15-lacinulate. Not at all well defined from the
similar A. platyphylla M. Arg. (Pax & Hoffm.), with which it
apparently should be merged, but I have seen little material. F.M.
Neg. 5322.
Loreto: Cerro de Escaler, 1,200 meters, Ule 6840, type. Co-
lombia.
Acalypha subbullata Pax & Hoffm. Pflanzenreich IV. 147.
xvi: 67. 1924.
Shrub, the stipules caducous, the flowers monoecious, the pubes-
cence dense and soft on the petioles (to 5 cm. long), the branchlets
and the leaves beneath, these glabrescent and bullate above, charta-
ceous, orbicular-ovate, acute, about 9 cm. long, 6-7.5 cm. wide,
5-7 nerved from cordate base; male spikes axillary, subsessile, about
a dm. long, nearly 4 mm. broad, dense, the dense terminal female
with 1.5 cm. long peduncle, to 7 cm. long, the 1-flowered pubescent
ovate obtusely acuminate bracts in flower 3 mm. long, entire but
minutely glandular-denticulate, below strongly nerved, ovary hirsute;
styles about 1.5 mm. long, shortly around 10-lacinulate. Could
prove when better known to be A. bullata. F.M. Neg. 32507.
Junin: Pariahuanca, Mathews 1199, type. Ayacucho: Ccarrapa
between Huanta and Rio Apurimac, about 1,500 meters, Killip &
Smith 22379 (apparently, but specimen seen without flowers).
Acalypha subcastrata Aresch. Sv. Eugenies Resa Bot. 3: 137.
1910; 42.
Annual branching slightly pilose herb often several dm. high;
petioles about as long as the ovate acuminate serrate leaves, these
144 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
usually 8 or 10 cm. long, 5-9 cm. wide, 5-nerved at base; spikes
terminal and axillary, if binate, one male, the other female, the
former subsessile, often very short or not exceeding 6 mm., the
latter equaling or exceeding the leaves and with laciniate bracts
longer than the scabrous or nearly smooth capsules, the laciniae
subulate, ciliate; ovary nearly glabrous; seeds subrugose or, accord-
ing to Svenson, Amer. Journ. Bot. 33: 458. 1946, practically smooth
and only 1.5 mm. long. Perhaps too near A. setosa Rich., more
northern, ovary hirsute. Differs from the Mexican and Central
American A. polystachya Jacq. in pilosity, shorter male spikes and
smoother seeds (Svenson) and it seems probable that the Pavon
specimen referred to that species by Mueller actually was mis-
labeled, as is sometimes known to have been the case, and really
came from Mexico.
Piura: Amotape Hills, (Haught & Svenson 11511}. Tumbez:
Choza, Weberbauer 7699 (det. Svenson). Ecuador.
Acalypha tenuipes Pax & Hoffm. Pflanzenreich IV. 147. xvi:
122. 1924.
Slender-branched shrub a meter or 2 high, early rather densely
puberulent pilose with short straight and crisped trichomes, few if
any glandular, later glabrate especially the thin-membranous leaves,
these on slender petioles 1.5-3.5 (4.5) cm. long, the ovate or ovate-
lanceolate palmately nerved blades rather coarsely crenate, acumi-
nate, lightly cordate at base, 4-6 (8) cm. long, 2.5-3 (4) cm. wide;
flowers monoecious, the usually androgynous axillary spikes slender
and mostly with 2-4 female bracts at base, laxly disposed, the
peduncled male portion 4-12 mm. long or several cm. in the Peruvian
specimen; female bracts about 7 mm. wide, orbicular-reniform,
lightly repand-denticulate or merely repand; ovary densely muricate,
puberulent; styles about 5 mm. long, finally around 10-lacinulate.
Apparently very near to A. Weddelliana Baill. of southern Brazil,
that less pubescent, but I have not seen the type at this time. F.M.
Neg. 5325.
Tumbez: In deciduous bush wood on brooks, east of Hacienda
Chicama, Weberbauer 7662. Ecuador.
Acalypha villosa Jacq. Sel. Stirp. Amer. Hist. 254. pi. 188.
1763; 16.
Tall, more or less copiously villous or tomentose shrub; stipules
subulate; petioles usually several to many cm. long, minutely 2-4-
FLORA OF PERU 145
stipellate; leaves ovate, truncate or rounded to the somewhat
cordate base, acuminate, sometimes only around 5 cm. long, 3 cm.
wide, often 8-18 cm. long, 5-12 cm. wide, crenate, membranous,
glandular; inflorescence branches often more or less unisexual; spikes
axillary, male sometimes with female intermixed, sometimes with
these at base, 3-13 cm. long, the female spikes 3-11 cm. long, much
more laxly flowered, often simple; rachis pubescent, filiform, bracts
minute, the male flowers glomerulate, the 2-several female minutely
pedicelled, sepals 5, lanceolate, ciliate; ovary muricate, not pubescent;
styles 2-3 mm. long, lacinulate to base; capsules about 2.5 mm.
thick, the subglobose smooth seeds scarcely 1 mm. long. Said
typically (var. genuina M. Arg.) to have villosulous branchlets
and petioles, soon puberulent-tomentulose, leaves barely or not
cordate at the shortly contracted base, glabrate in age. Named
Peruvian variants include tomentosa M. Arg., leaves rounded at
base, pubescence soft, more lasting; intermedia M. Arg., leaves
cordulate, pubescence tomentulose; latiuscula Pax & Hoffm., leaves
subtruncate-cordulate, glabrate except nerves. F.M. Neg. 32504
(var. tomentosa).
San Martin: Tarapoto, Ule 6638; Williams 5487; 6024; 6270.
Chazuta, Klug 4114 (det. Standley). Without locality, Ruiz &
Pavdn, type, var. tomentosa (det. Boissier). Ayacucho: Estrella
between Huanta and Rio Apurimac, Killip & Smith 22661; 22662?
Central America to Bolivia and Paraguay.
Acalypha Wilkesiana M. Arg. in DC. Prodr. 15, pt. 2: 817.
1866.
Nearly glabrous bush or small tree except for some puberulence
on the branchlet tips and younger leaves, these becoming broadly
ovate or rotund and characteristically variegated or mottled with
red or purple in some degree, somewhat acuminate, rounded or
shortly cuneate at base, serrate, sometimes a dm. or two long and
more than half as wide; flowers monoecious in axillary spikes, only
the laxly flowered female much shorter than the leaves, these with
ovate 1-flowered bracts, the middle of the 9-13 dentations much the
longest; sepals 3-4, acuminate; ovary puberulent; styles 11-15-
lacinulate. Variable and long grown for its ornamental foliage in
greenhouses and in tropical regions where it may become established ;
its origin is thought to have been in the South Pacific. Illustrated,
Seemann, Fl. Vit. pi. 58.
Loreto: Bush in forest at Cajacuma on the Amazon River,
Williams 2010 (det. Standley). Widely cultivated in tropical regions.
146 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
23. ALCHORNEA Swartz
Reference: Pax & Hoffm., Pflanzenreich IV. 147. vii: 220-252.
1914.
Shrubs or trees often laxly branched, the indument stellate or
simple, the leaves more or less dentate, bistipuled, petioled, basally
and often on the face beneath maculate-glandular, the spikes
normally unisexual, simple or paniculately branched, the male often
axillary, the female often terminal, the former several to each
bract, the latter only 0-3. Flowers monoecious or dioecious, apeta-
lous. Male calyces valvately 2-5-parted; stamens 8 (or fewer), the
filaments connate at small disk, the anthers oblong; rudimentary
ovary none. Female sepals 4 (3-6); ovary 2-3(4)-celled; styles free
or basally connate, usually entire. Capsules composed of 2-3 cocci
or subglobose, smooth or tuberculate, the seeds nearly or quite
ecarunculate. Named for Alchorne, English botanist.
Leaves with 2 basal nerves besides midnerve.
Leaves with 1 or more secondary nerves besides the basal 2.
Leaves multiglandular at base, membranous A. glandulosa.
Leaves 2 (-4), glandular at base, coriaceous.
Secondary nerves besides basal 3-5 pairs; petioles stout,
many times shorter than blades.
Leaves 4-8 (10) cm. long, coriaceous.
Styles 2-3 mm. long; leaves oblongish, narrowed below
the middle A. brevistyla.
Styles 7-15 mm. long; leaves ovate, rounded at base.
A. acutifolia.
Leaves mostly 10 cm. long or longer, rigid, acuminate.
A. Pearcei.
Secondary nerves besides basal 1-2 (-4) pairs; petioles slender,
mostly about a third as long as blades. . . . A. triplinervia.
Leaves lacking secondary nerves besides the basal 2. A. acroneura.
Leaves pinnate-nerved from base, oblong A. castaneifolia.
Alchornea acroneura Pax & Hoffm. Pflanzenreich IV. 147. vii:
229. 1914.
Branchlets strict, slender, early with a few simple trichomes;
petioles 2-4 mm. long, canaliculate, sparsely pilose; leaves spathulate-
obovate or -lanceolate, shortly and rather abruptly narrowed to
base, obtuse and slightly emarginate at tip, 3-4.5 cm. long, 16-22
FLORA OF PERU 147
mm. wide, minutely biglandular above the base, denticulate, coria-
ceous, glabrate above, lightly stellulate beneath, 3-nerved from base
but with no other secondary nerves, the horizontal veins little
prominent; flowers monoecious but the male and female spikes on
diverse branchlets, the former simple, erect, slender, 2-8 cm. long,
bracts acute, about 3-flowered, the latter (one seen) 2 cm. long with
6-7 sterile bracts below, the upper 1-flowered; bracts 1 mm. long,
acute, strongly thickened at base; male calyx glabrous, subsessile;
stamens 8, anthers apiculate; female calyx pedicel 1 mm. long, sepals
4, lanceolate, acute, puberulent; ovary stellulate; styles 2, nearly
free, broad, margins narrowly revolute, smooth. Williams, Field
Mus. Bot. 15: 268. 1936, reduced this to A. triplinervia. Illustrated,
I.e. p. 224. F.M. Neg. 24413.
San Martin: Moyobamba, 1,500 meters, Weberbauer 4752, type.
Alchornea acutifolia M. Arg. Linnaea 34: 171. 1865; 226.
Subscandent shrub with terete fulvous tomentose tardily glabrate
branchlets; petioles 5-15 mm. long, deflexed; leaves ovate or ovate-
lanceolate, base lightly cordate, especially toward cuspidate-acumi-
nate apex sparsely serrate-denticulate, 4-10 cm. long, 2-4 cm. wide,
3-nerved, with also 3-5 secondary prominent as veins, coriaceous,
glabrous and papillose above, early stellate-pilose beneath on the
nerves and at base maculately 2-4-glandular; male spikes equaling
or exceeding leaves, the female simple, 5-15 cm. long, rachis striate,
puberulent, bracts small, those of the male several-flowered, of the
female 1-flowered, 1-2-glandular both sides; flowers sessile; male
calyx glabrous except at tip; stamens 8, anthers apiculate; female
sepals 4, 0.5 mm. long, acute, tomentose; ovary 2-celled, spreading
hispid-pilose, the purple styles very shortly connate, papillose
within, 7-15 mm. long; capsules 9 mm. thick, 5.5 mm. long, pubes-
cent, the obtusely tuberculate subglobose seeds 5.5 mm. in diameter.
Said to attain 5-9 meters. F.M. Neg. 5262.
San Martin: Tarapoto, Spruce 4269, type; Ule 6409. San Roque,
Williams 7532 (det. Mansfeld). Zepelacio, King 3272 (det. Standley,
A. triplinervia var.). In savannah woods, Moyobamba, Weberbauer
4481; 4482; 2S9.
Alchornea brevistyla Pax & Hoffm. Pflanzenreich IV. 147. vii:
227. 1914.
Similar in general character to the allied A. acutifolia; petioles
not deflexed, 1-2 cm. long; leaves oblong, 7-9.5 cm. long, 27-38 mm.
148 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
wide, the secondary nerves less prominent; female spikes 3-5 cm.
long, simple, solitary or fasciculate in the leaf-axils, the solitary
pedicels 1 mm. long and bibracteolate, sepals 0.75 mm. long, sub-
glabrous; ovary glabrous; styles 2, free, smooth, 2-3 mm. long;
male flowers unknown. As remarked by the authors, the known
characters seem to be very diverse from those of the species which
it so greatly resembles otherwise. Williams, however, Field Mus.
Bot. 15: 268. 1936, reduced it to A. triplinervia. F.M. Neg. 5265.
Loreto: Iquitos, Ule 6250, type; Killip & Smith 27301 (det.
Mansfeld). Yurimaguas to Balsapuerto, Killip & Smith 28157. \
Mishuyacu, Killip & Smith 29937?
Alchornea castaneifolia (Willd.) Juss. Euphorb. Tent. 42.
1824; 235. Hermesia castaneifolia Willd. Sp. PI. 4: 809. 1805.
Small nearly glabrous tree well-marked by the oblong-lanceolate
leaves often 8-16 cm. long, 1.5-4 cm. wide, acutish or rounded each
end, repand-serrulate or -spinulose, obscurely or obsoletely glandular
at base, minutely stellulate or glabrate beneath, finely and reticu- i
lately pinnate-nerved; petioles 5-10 mm. long; male spikes simple,
many-flowered, 1-2 dm. long; male perianth globose, filaments
eglandular; fruiting racemes about as long as leaves, bracts ovate,
acute, pedicels about 5 mm. long, the capsules as long or longer,
minutely stellulate or glabrate, depressed-globose, the shorter styles
spreading; seeds 7-8 mm. long. Belongs to section Hermesia (Willd.)
Croizat, marked by the pinnate-nerved obscurely glandular leaves
and small female flowers. In Brazil at water's edge with Salix
(Mexia) or on river bank (Williams). Illustrated, Pax, I.e. p. 236.
Loreto: Lower Rio Nanay, Williams 496. Yarina Cocha, Tess-
mann 3488. Colombia to Paraguay. "Yacochihua" (Williams), I
"ipurosa" (Tessmann).
Alchornea glandulosa Poepp. in Poepp. & Endl. Nov. Gen. &
Sp. 3:18. pi. 201.1845; 234.
Branches, petioles (1-7 cm. long) and spikes tomentulose, the
former tardily glabrescent; leaves oblong-elliptic, -ovate or rarely
broader, acute or rounded or narrowly subcordate at base, abruptly
caudate-acuminate or acute at apex, 9-16 cm. long, 5-9 cm. wide,
glandular-dentate, membranous, early finely stellulate both sides,
the trichomes a little longer on the nerves, glabrate above, basally
3-nerved and also with 3-6 secondary ones; basal glands 4-7; all
spikes axillary, the male panicled or simple, 2-11 cm. long, triangular
FLORA OF PERU 149
bracts about 7-flowered, the flowers 2.5 mm. broad, sepals 2-3,
ovate, acute, glabrous; stamens 8, anthers not or minutely apiculate;
female spikes simple, 3-6.5 cm. long, bracts ovate, subsaccate,
1-2-flowered, the 4 sepals triangular-lanceolate, acute, as long as
densely fulvous ovary; styles filiform, free, papillose, 5-13 mm. long.
Var. Pavoniana M. Arg. has the leaves subcordate at the narrowed
base, glands obscure. A. leptogyna Diels, Bibl. Bot. 116: 103. 1937,
Ecuador, is less pubescent, leaves to 2 dm. long, 7-10 cm. wide,
acumen to 2 cm. long, female spikes lax, the ovary less pilose. Deter-
minations by Mansfeld. According to Williams attains 15 meters.
F.M. Negs. 7151; 7150 (var.).
Lore to: Yurimaguas, Poeppig 2198, type; Williams 3899; 4621;
Killip & Smith 28209. Without locality, Pavdn (type, the var.).
To Ecuador, Colombia and Venezuela; Costa Rica?
Alchornea Pearcei Britton, Bull. Torrey Club 28: 305. 1901;
225.
Branches often stout, the younger more or less stellate-pilose;
petioles 1-2 cm. long, transversely rugose above, glabrous or pubes-
cent; leaves oblong or elliptic, acuminate, acute or narrowed or
broadly obtuse, even subcordate at base, 9-16 cm. long, 4-7.5 cm.
wide, maculately biglandular, 3-plinerved, crenate-dentate, rigid-
coriaceous, somewhat stellulate beneath and often barbate in the
axils of the prominent nerves; male inflorescences often about
2 dm. long, the simple pendulous female a third longer; male bracts
about 3-flowered, female 1-flowered, the flowers subsessile, the male
2.5 mm. broad, reddish-brown with 2 ovate glabrous sepals, 8
stamens, scarcely apiculate not emarginate anthers, the female
with 4 broadly triangular sepals, 2-celled ovary, entire subfree
styles 1-2 cm. long; capsules finely stellate, 11-12 mm. broad, 7-8
mm. long, the echinate-tuberculate or nearly smooth seeds 6-7 mm.
in diameter. There are two forms: var. sclerophylla Pax (the
typical) glabrescent, the inflorescence puberulent and var. coriacea
(Ule) Pax (A. coriacea Ule, Verh. Bot. Ver. Brandenb. 50: 76. 1908),
branchlets as petioles and inflorescence villous. F.M. Neg. 24414.
San Martin: San Roque, Williams 7506 (var. coriacea, det.
Mansfeld). Junin: La Merced, 5755 (det. Johnston). Loreto:
Cerro de Escaler, 1,300 meters, Ule 6836 (type, part, A. coriacea).
Puno: Chunchusmayo, Sandia, 300 meters, Weberbauer 1226 (var.
type, A. coriacea and sclerophylla). Bolivia.
150 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Alchornea triplinervia (Spreng.) M. Arg. in DC. Prodr. 15,
pt. 2: 909. 1866; 227. Antidesma triplinervia Spreng. Neue Entd.
2: 116. 1821.
Younger parts somewhat stellate-puberulent-tomentose but often
soon glabrate except the leaves beneath and the inflorescence;
petioles usually several cm. long; leaves usually ovate or elliptic,
ordinarily obtuse to subcordate at base and acute or acuminate at
apex, sometimes obtuse, 2.5-15 cm. long, 2-11 cm. wide, more or
less dentate, often coriaceous, reticulate-veined, elongately 3-nerved
with 1-2, rarely 3-4 additional secondary nerves, maculately 2-4-
glandular between the basal ones and sparsely stellulate, above
soon glabrate; spikes of both sexes axillary, simple or paniculate,
the solitary or fasciculate male 5-20 cm. long, the laxly flowered
female 3-12 cm. or often reduced; female bracts 1-2-flowered as
the male thickened at base, male pedicel short, female 1-2 mm. long;
male flowers with 2 orbicular glabrous sepals, those of female 4,
ovate, acute, puberulent; ovary pilose, styles 2, free or nearly,
7-15 mm. long, subsmooth; capsules 7-10 mm. broad, 5-7 mm. long,
finely pilose, the pedicels 2-3 mm. long; seeds 4 mm. thick, echinate-
tuberculate. The monographers follow Mueller in recognizing many
variants which mostly are not clearly defined and fail to indicate
the typical characters; to var. crassifolia M. Arg. they refer a Ruiz
specimen without locality, defining it as glabrate, petioles 2.5-6 cm.
long, leaves to 9 cm. long, subcoriaceous. Williams, Field Mus.
Bot. 15 : 269. 1936, found the wood of the variety with slightly offen-
sive odor and astringent taste, these qualities lacking in the material
determined as typical. F.M. Negs. 21503 (var.); 21504 (var.);
32553.
Loreto: Iquitos, Killip & Smith 27299; Williams 1527; 7988;
7972; 7980 (last two, var.). Upper Itaya, Williams 3540. San
Ramon, Williams 4589. Caballo-Cocha, Williams 2162; 2186 (both,
var.). Fortaleza, King 2824. Without locality, Ruiz & Pavdn.
Southern Brazil to Paraguay and Colombia. "Mojarra," "cocopano"
(both, Williams).
24. APARISTHMIUM Endl.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vii: 257-259.
1914.
Character in general that of Alchornea but stamens usually 4
(3-5), the filaments long-connate and, especially, styles basally
FLORA OF PERU 151
connate, bilobed, the sessile stigma nearly as broad as long and
coarsely papillose within. The ample leaves are deflexedly bistipel-
late at the tip of the petioles as well as bistipulate at their base.
The genus could be included in Alchornea as by M. Arg. Linnaea
34: 168. 1865.
Aparisthmium cordatum (Juss.) Baill. Adansonia 5: 307.
1865; 258. Conceveibum cordatum Juss. Euphorb. Tent. 43. pi. 13,
fig. 42a. 1824. Alchornea cordata (Juss.) M. Arg. in DC. Prodr. 15,
pt. 2: 901. 1866.
Slender tree sometimes about 15 meters high, the branchlets and
petioles, these 3-18 cm. long, somewhat puberulent; stipules cadu-
cous, subulate, 3 mm. long, the apical stipels 1.5-4 mm. long; leaves
usually broadly ovate, often rounded-cordate at base, abruptly shoj* t-
or long-acuminate, commonly 1-3 dm. long, a half to two thirds as
broad, denticulate to subentire, membranous or chartaceous, more
or less short-pilose or glabrate both sides and with 2 basal glands
beneath, the 7-11 secondary nerves prominent; male panicles and
terminal female racemes about as long as the leaves, bracts of both
biglandular, pedicels of latter 2-15 mm. long, flowers of former
2 mm. long, pilose without; stamens exserted, of latter, 1-2 mm.
long, ciliate and puberulent; capsules 8-11 mm. thick, 6-8 mm. long,
deeply 3-lobed, sparsely puberulent, the ellipsoid brown striolate
seeds 5-6 mm. long, 3.5-4 mm. broad. Williams, Field Mus. Bot.
15: 270. 1936, noted flowers as precocious (?), young bark yellowish,
older brown, smooth, wood soft but used for huts (Mexia). Deter-
minations mostly by Mansfeld. Illustrated, Pax, I.e. p. 259.
Junin: Puerto Yessup, Killip & Smith 26285. Loreto: La
Victoria, Williams 2582. Lower and upper Nanay, Williams 356;
645; 649. Near Iquitos, King 252; 1042; Killip & Smith 27298;
27313; 29904- Sierra del Pongo, 800 meters, Mexia 6273; 6274 (det.
Standley). Bolivia to Colombia, Guiana and Brazil (Minas Geraes).
"Rucurana" (Williams), "shamboquiro" (Mexia), "ukskakiro"
(Tessmann).
25. CONCEVEIBASTRUM (M. Arg.) Pax & Hoffm.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vii: 217. 1914.
In general like Conceveiba but male flowers unknown. Indument
stellate and simple. Stipules conspicuous, almost foliose. Female
flowers in stout terminal racemes with 10-12 broad imbricate sepals
152 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
notably biglandular without. Styles biparted to base, the circinate-
involute lobes coarsely papillose within. Seeds ecarunculate, trun-
cate. Probably this plant, apparently marked only by the sepals,
and several related ones should be brought into one group but until
completely known another disposition seems inadvisable, since only
by chance would it prove to be definitive.
Conceveibastrum Martianum (Baill.) Pax & Hoffm. Pflanz-
enreich IV. 147. vii: 217. 1914. Conceveiba Martiana Baill. Adan-
sonia 5: 221. 1865. Akhornea Martiana (Baill.) M. Arg. in Mart.
Fl. Bras. 11, pt. 2: 375. 1874.
Tree with stout angulate densely leafy reddish puberulent branch-
lets and lanceolate acute stipules retrorsely saccate-produced at
base, 3-4 cm. long, 6-7 mm. wide; petioles glabrate, 9-12 cm. long;
stipules auriculate-verruciform; leaves ovate, narrowly and deeply
cordate at base, obtuse, 1-3 dm. long, 1-2.5 dm. wide, crenate-
dentate, chartaceous, finally glabrate above, beneath especially on
the 7 basal nerves simply and stellately rusty pubescent; fruiting
racemes very stout, nearly 2 dm. long in type, bracts 2 mm. long,
acute, with 2 large patellate basal glands; pedicels 6-9 mm. long,
apically articulate; sepals 2-3 mm. long, 3-4 mm. wide, apiculate,
glandular; ovary globose, fulvous tomentose; capsules 18 mm. broad,
tomentulose; seeds 8 mm. long, 10 mm. broad, obtusely trigonous,
smooth. F.M. Neg. 5275.
Loreto: Mouth of Rio Santiago, Tessmann 1+1^9 (det. Dahlem).
Amazonian Brazil.
26. CONCEVEIBA Aublet
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vii: 214-216.
1914.
Trees soon glabrescent with rather ample alternate petioled bi-
stipuled reticulate-veined leaves and terminal puberulent panicles
of apetalous flowers, the male ample, the small flowers glomerulate,
the female more simple with biglandular basal bracts. Disk none.
Male calyx valvately 3-4-parted; stamens about 16, free on the
convex receptacle, the 6-8 outer fertile with short filaments; anthers
erect, connective broad; rudimentary ovary none. Female sepals
5-8, imbricate, with alternating glands; styles little or scarcely
connate, bilobed, thick-papillose. Ovary usually 3-celled. Capsules
large, smooth or echinate, the seeds carunculate.
FLORA OF PERU 153
Styles longer than calyx; capsules smooth, 3-ribbed. .C. guianensis.
Styles short; capsules subechinate-rugulose C. rhytidocarpa.
Conceveiba guianensis Aublet, PI. Guian. 2: 924. pi. 353.
1775; 214.
Straight-trunked tree, nearly glabrous except for the puberulent
tips of the stout branchlets; petioles 2-4.5 cm. long; leaves usually
distinctly glandular at obtuse or acutish base, obtusely short- or
caudate-acuminate, often remotely denticulate, 1-3 dm. long, about
half as wide, oblong- or oval-elliptic, sparsely stellate at least beneath,
puncticulate above, membranous or coriaceous; male flowers small,
subsessile in many fascicles on the spreading branches of the ample
panicles; female flowers in racemes, the basally bracteolate pedicels
elongate; ovary tomentose, 3-ribbed as capsule, the basally connate
styles recurved-spreading; capsules 2-2.5 cm. long, 2 cm. broad,
ligneous, the smooth seeds 1 cm. long, the caruncle small. Steyer-
mark, Field Mus. Bot. 17: 414-415. 1938, proposed three new
species from as near as Sao Paulo de Olivenca, Brazil, which may be
expected within Peru: C. magnified, sepals eglandular, C. Krukoffii,
style column longer, 3 mm. long and C. simulata, male inflorescence
shorter, fertile stamens usually 9, sterile 9-10, instead of 6-8 for
each; fruits are unknown for these and they may replace the typical
form in the Upper Amazon but the observed differences may be
included within the range of variation for C. guianensis. The seeds
are said to be edible. Illustrated, Mart. Fl. Bras. pi. 55 and Pax &
Hoffm. I.e. 215.
Peru (probably or one of the related forms as noted above).
Guianas.
Conceveiba rhytidocarpa M. Arg. in Mart. Fl. Bras. 11, pt. 2:
372. 1874; 216.
Resembles C. guyanensis; leaves obsoletely biglandular at base,
firm; male flowers unknown; female inflorescence spicate; ovary
4-celled; style very short, bilobed; capsules 13-14 mm. broad, sub-
globose, densely subechinate-rugose. Doubtful species apparently
aberrant in its 4-celled ovary. C. trigonocarpa M. Arg., 216, has short
erect styles on an acutely trigonous ovary. F.M. Neg. 24395.
Loreto: Yurimaguas, Poeppig 2485, type.
27. CARYODENDRON Karsten
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vii: 263-264.
1914.
154 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Trees, glabrous except for some simple pubescence in the inflores-
cences of dioecious apetalous flowers, the ample alternate entire
leaves bistipulate and maculately biglandular above the obtuse or
acute base. Male spikes terminal or axillary below the branchlet
tips, the flowers sessile, subglobose in bud, valvately 3-lobed at
an thesis, the lobes ovate; stamens 4-6, inserted about a thick broad
disk, sometimes with one in the center; filaments free, the dorsifixed
anthers ovoid. Female sepals ovate (disk annulate), imbricate under
the ovoid-globose hard dehiscent 3(2-4)-celled fruit (this unknown in
Peruvian species). Seeds solitary, ecarunculate.
Caryodendron grandifolium (M. Arg.) Pax in Pflanzenfam. 3,
Abt. 5: 52. 1890; 263. Centrodiscus grandifolius M. Arg. in Mart.
Fl. Bras. 11, pt. 2: 327. pi. 102. 1874.
Younger branchlets compressed toward tips, soon obtusely angled;
petioles stout, 1-5 cm. long; stipules ovate, subacute, rigid, ciliolate,
glabrate, about 3 mm. long; leaves elliptic-oblong, shortly and
obtusely acuminate, about 1.5-2 dm. long, to 1 dm. wide, cartilag-
inous-margined, pale green, reticulate-veined with 9-12 pairs of
secondary nerves; male spikes floriferous to base, stout, erect, 5-9
cm. long, rachis 1.5-2 mm. thick, the approximate very broad rounded
ciliate bracts many-flowered; pulvini of the sessile flowers densely
bracteolate, glabrous flowers 2 mm. long at anthesis; male sepals
broadly ovate, acute; stamens 6-7, connective entire, truncate; disk
ciliolate, thick, produced between the filaments and quasi 6-radiate;
female flowers and fruit unknown. In the similar type of the genus,
C. orinocense Karsten, to be expected within Peru and known by
the native name "Tagni" or "Palo de Nuez," with only 4 stamens,
the connective bicuspidate, the glabrous capsule is nearly 4 cm. wide
and subtended by the 5 or 6 obtuse sepals, the edible ovoid seed
about 3 cm. long; it is illustrated by Karsten, Fl. Columb. 1: pi. 45.
Simulates in general appearance species of Sapium; the white light
wood serves for the preparation of a coal for Sprengpulver (Peckolt,
Ber. Deutsch. Pharm. Gesellsch. 15: 242. 1905).
Rio Acre: Rio Macauhan, Krukoff 5553 (det. Steyermark).
Southern Brazil. "Pao podre" (Brazil).
28. ADENOPHAEDRA M. Arg.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vii: 261-262.
1914.
FLORA OF PERU 155
Nearly smooth shrubs, the caducous indument simple, with
cuneate-based shortly petioled alternate pinnate-nerved leaves and
apetalous dioecious flowers borne in very slender elongate spikes,
the male sometimes paniculately branched. Male perianth 3 (rarely
2) -parted, stamens 3 (2), the short stout filaments connate at base,
anthers ovate, cells distinct, connective glandularly enlarged and
produced; disk and ovary none or latter very minute; female perianth
with 6 biseriate imbricate sepals, the 3 inner smaller, disk annulate,
ovary 3-celled, styles sessile in the stigmatic 3-lobed disk. Capsules
depressed, seeds globose, smooth, ecarunculate.
Adenophaedra megalophylla M. Arg. in Mart. Fl. Bras. 11,
pt. 2: 386. pi. 101. 1874; 262.
Branchlets slender, early appressed strigose; petioles 3-30 mm.
long, in type soon glabrous, rugulose; leaves obovate-spathulate or
elliptic-lanceolate, acute, long-cuneate to base, repand-dentate or
subentire above, subcoriaceous, glabrous, spotted glandular above,
reticulate-veined beneath, often 12-22 cm. long, 4.5-9 cm. wide;
male spikes terminal and in the upper axils, equaling or exceeding
the leaves, the branches spreading, rachis pubescent, slender; bracts
distant, ovate, acuminate, many-flowered; pedicels 1-1.5 mm. long;
sepals minute, ovate, acute, puberulent; glands large, nearly as long
as anthers; larger outer female sepals 3 mm. long; ovary orange-red
pubescent; capsules 10-11 mm. broad; 7 mm. long, deeply lobed;
seeds 5 mm. long, subglobose, maculate-striate. The Peruvian
specimens are young; the petioles are densely pubescent and the leaf
glands few and obscure; the related species A. grandifolia (Klotzsch)
M. Arg. (but female flowers unknown to author) has eglandular
leaves, male inflorescence simple. A tree 2.5-6 meters high (Klug).
Loreto: Mishuyacu near Iquitos, Klug 61; 1383 (both det. Mans-
feld). Southern Brazil.
29. CLEIDION Blume
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vii: 288-298.
1914.
Shrubs or small trees with alternate usually dentate petioled
leaves and apetalous flowers, the male sessile in axillary spikes, the
female well-pedicellate in racemes or panicles, the pedicels some-
times apically clavate. Indument simple. Disk none. Male calyx
valvately 3-4-parted; stamens many (35-80) congested in alternate
156 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
verticils on a convex or conical receptacle, the younger anthers affixed
dorsally about medially to filaments, the 4 quasi-cruciate cells finally
confluent. Rudimentary ovary none. Female sepals 3-4 (5),
imbricate. Ovary 3(2)-celled, the elongate filiform styles often
connate basally, deeply bifid, densely and minutely papillose within.
Capsule-cocci sometimes by abortion 1, the subglobose seeds ecarun-
culate. Seems to be much like Akhornea except for the cleft styles,
a character which elsewhere in the family is not always generically
significant. However, Croizat, Journ. Arnold Arb. 24: 167. 1943,
remarks that "both tradition and an aggregate of characters have
rightly maintained them as distinct genera."
Petioles about 5 mm. long; female inflorescence branched.
C. amazonicum.
Petioles much longer; female inflorescence simple . .C. castaneaefolium.
Cleidion amazonicum Ule, Verh. Bot. Ver. Brandenb. 50: 76.
1908; 294.
Shrub or small tree 3-6 meters high, the terete slender branchlets
early appressed puberulent; petioles 5 mm. long, pubescent as the
subpersisting subulate 4 mm. long stipules; leaves oblong or obovate,
obliquely and narrowly cordate at base, shortly cuspidate-acuminate,
the apex itself obtuse or sometimes mucronulate, 10-23 cm. long,
5-8 cm. wide, denticulate, firm-membranous, basally maculate-
glandular beneath, the 6-8 nerves slightly pilose, the veins reticulate;
male spikes to 2 cm. long, densely flowered to base, the rachis
spreading pilosulous, the acuminate 1 mm. long bracts 3-5-flowered,
the 1 mm. long elliptic sepals acute; stamens about 50; female
panicles 9-12 cm. long, ovate acuminate bracts 1.5 mm. long, 1-3-
flowered, the pedicels apically enlarged, 4-14 mm. long, sepals 1.5
mm. long, lanceolate, medial nerve carinate pubescent; styles 5-8
mm. long, linear, reddish within; capsules unknown. Bark thin
and smooth (Williams).
Lore to: Paranapura, lower Huallaga, Williams 4600; 5272 (det.
Mansfeld). Florida, King 2025; 2367 (det. Standley). Rio Acre:
Varzea land, Rio Macauhan, Krukoff 5694- Amazonian Brazil;
Bolivia.
Cleidion castaneaefolium M. Arg. Linnaea 34: 184. 1865; 295.
Branchlet tips as the apically tumidulous petioles (these 1.5-4
cm. long) and inflorescences subvelutinous with short fulvous tri-
FLORA OF PERU 157
chomes; leaves lanceolate-elliptic, acute at base, shortly cuspidate-
acuminate, 18-22 cm. long, 9-12 cm. wide, coarsely and obtusely
serrate; male spikes many times shorter than the leaves, flowers
glomerately spicate, bracts broadly ovate, acuminate, sepals ovate,
acute, little puberulent; stamens about 70; complete female spikes
not seen, sepals narrowly triangular-ovate, acuminate, ovary 3-celled,
pubescent, styles firm, linear; capsule large with 6 alternately
unequal lobes, the cocci carinate dorsally. This is incompletely
known. F.M. Neg. 7159.
Peru: Without locality, Ruiz & Pavdn, type.
30. BERNARDIA [Houst.] Adans.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. vii: 21-44. 1914.
Usually shrubby and pubescent with simple or partly fasciculate
trichomes, alternate pinnate- or basally trinerved leaves, more or
less dentate and maculate glandular at base, stipules small. Male
flowers in axillary spikes sometimes short and capituliform with
numerous bracts, the female flowers ordinarily in terminal short
inflorescences or in the upper leaf -axils or in many-flowered racemi-
form spikes with firm or coriaceous concave bracts. Male calyx
valvately 3-4-parted at anthesis with 4-22 free stamens, the filaments
dilated toward base, the subglobose anthers quasi-cruciately 4-celled;
disk usually glandular; rudimentary ovary none. Female sepals 4-6,
hypogynous disk annulate with distinct glands, ovary 3-celled, the
short styles often continuous with carpels, the style lobes smooth
or lacerate. Capsules 3-lobed, endocarp crustaceous, seeds ecarun-
culate, more or less carinate. Genus commemorates one of a
famous family, Bernard de Jussieu (1699-1776). It is probable
that the following species or one of several from Brazil will be found
in southern Peru.
Bernardia Jacquiniana M. Arg. Linnaea 34: 173. 1865; 31.
Shrub or little tree, the pubescence somewhat spreading or mostly
on the branchlets retrorse, the subterete branches equally foliose
with lanceolate-oblong membranous crenate-denticulate leaves
usually 1-2 dm. long, about a third as wide, acute at biglandular
base and many-glandular toward the acuminate apex; petioles 2-10
mm. long; secondary nerves 5-10, the tertiary impressed reticulate
above; stipules subulate, 1.5-2 mm. long; male spikes 5-10 cm. long,
often floriferous to base, densely tomentose, bracts broadly ovate,
158 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
acuminate, carinate, 7-10-flowered, pedicels 1 mm. long; sepals 3,
oblong, acute, densely pubescent both sides; stamens 9-12; disk
glands small; female sepals 5, pubescent without, 3-4 mm. long;
disk urceolate; ovary sericeous; styles deeply bifid; capsules about
5 mm. long, tomentulose; seeds 4 mm. long, crenate-subdentate
below dorsally. F.M. Neg. 5253.
Peru (probably). Ecuador to Venezuela.
31. JATROPHAL.
Reference: Pax, Pflanzenreich IV. 147: 21-113. 1910, and later;
see I.e. xiv. Add. 6: 38. 1919.
Ligneous, sometimes trees or fleshy rhizomatous perennial herbs
with alternate usually palmately lobed or divided leaves (rarely
entire or pinnately lobed) often more or less glandular and small
or setaceously dissected stipules that are sometimes spinescent.
Pubescence never stimulose. Petiolar glands various, but not as in
Cnidoscolus. Flowers commonly monoecious, disposed in dichoto-
mously branched cymes. Sepals and petals 5, the former imbricate,
more or less connate at base or in male flowers forming a tube and
petaloid, the latter when present contorted-imbricate, free or some-
what coherent, thus seemingly sympetalous. Disk gland if annular
sessile, usually distinctly 5-glandular. Stamens biverticillate, 6-10,
more or less monadelphous, the outer 5 epipetalous; staminodia
usually none or filiform. Ovary 2-3(4-5)-celled, styles connate at
base, entire or bifid, the stigmas capitate or hooded; rudimentary
stamens rare, ovules solitary. Fruit capsular, seeds carunculate.
According to McVaugh pith is usually relatively small and solid in
contrast to that of the closely related genus Cnidoscolus.
Leaves eciliate, the lobes entire or with a few teeth.
Leaves shallowly lobed if at all J. Curcas.
Leaves deeply lobed.
Stipules glandiform, sessile; leaves epeltate J. macrantha.
Stipules dissected.
Leaves subbasally multi-lobed J. multifida.
Leaves medially 5-lobed, peltate J. Weberbaueri.
Leaves ciliate, often glandular and minutely denticulate.
Leaves shallowly lobed, medially peltate J. Humboldtiana.
Leaves deeply and acutely lobed, often epeltate.
FLORA OF PERU 159
Leaves broadly 3-5-lobed, never nearly to base.
Leaves all or mostly 5-lobed; stipules glandiform. . .J. ciliata.
Leaves all or mostly 3-lobed; stipules as petiolar trichomes
dissected J. gossypifolia.
Leaves narrowly 5-7-parted nearly to base; stipules dissected,
petiolar trichomes simple J. clavuligera.
Jatropha ciliata M. Arg. Linnaea 34: 209. 1865; 47. J. Hoff-
manniae Croizat, Journ. Arnold Arb. 24: 168. 1943. J. longipeduncu-
lata Pax & Hoffm. Pflanzenreich IV. 147: 85. 1910, and 147. xvii.
Add. 7: 191. 1924, not Brandg. 1920. J. Augusti Pax & Hoffm. I.e.
Glabrous, with stout cicatrose rather fleshy branches; petioles
6-12 cm. long; stipules reduced to 1-3 sessile glands; leaves 6-12
cm. wide, and long, openly cordate at base, or sometimes somewhat
peltate, more or less 5-lobed, the lobes cuspidate-acuminate and un-
equally ciliate-denticulate and glandular exceeding the early shortly
(often elongating) and stoutly peduncled corymbiform cymes; bracts
linear-lanceolate or ovate; sepals 2.5 mm. long, oblong-rhombic to
orbicular-ovate, erose-denticulate, acute (male) or acuminate; petals
scarlet, about 1 cm. long; male disk with ovate squamiform glands;
stamens 8 (9), shortly monadelphous; ovary glabrous or essentially,
the brown seeds 1 cm. long. At flowering nearly or quite leafless.
With regret I have concluded that J. Augusti is merely the state
with leaves shallowly peltate; my 2331 exhibits openly cordate leaves
with one or two peltate ones. Plants with peduncles to 15 cm. long
may be noted as var. longipedunculata (Pax & HofFm.) Macbr.,
comb. nov. J. longipedunculata Pax & Hoffm. I.e. not Brandeg.,
J. Hoffmanniae Croizat but the typical form grows in the same
region. Stork and Horton noted the long peduncles as "conspic-
uously red." F.M. Negs. 24382; 5379 (J. Augusti).
According to Ruiz and Pavon an infusion of the root was con-
sidered a powerful aphrodisiac, a reputation still persisting (West).
Huanuco: Near Huanuco, Ruiz & Pavdn, type, also Dombey
2331; Stork & Horton 9394; stony hillsides, Weberbauer (type,
J. Augusti); Soukup 2226; Ledig 12. Huancavelica: Rio Mantaro
below Succubamba, 1,500 meters, Weberbauer 6488 (type, J. longi-
pedunculata); Stork & Horton 10401. Apurimac: 2,500 meters,
Weberbauer 5850 (det. Mansfeld). Rio Pachachaca, West 3796;
Goodspeed Exped. 10530; Herrera 3053. "Huanarpo," "huanarpo-
maachou," "higos del duende."
160 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Jatropha clavuligera M. Arg. Linnaea 34: 209. 1865; 28.
Branchlet tips as the leaves both sides and shortly peduncled
cymes rather densely short-villous, this indument on the long
petioles mixed with some simple gland-tipped longer subsetose tri-
chomes; leaves 5-7-parted nearly to the cordate base, the divisions
lanceolate or narrowly elliptic, minutely denticulate, acute, 4-8 cm.
long, 8-16(-30) mm. wide; stipules dissected, 5-7 mm. long; bracts
lanceolate, glandular ciliate as the acuminate male sepals, these
4-5 mm. long, the glabrous obtuse purplish or blood-red petals
little longer; disk glands truncate, free; filaments monadelphous
nearly entire length; female sepals slightly larger, the petals acute;
capsules more or less hirsute, subglobose, about 1 cm. long, the pale
seeds spotted. The single Peruvian collection has smaller leaves.
F.M. Neg. 7160.
Cajamarca: Bellevista to union of Rios Chinchipe and Maranon,
600 meters, Weberbauer 6223 (det. Mansfeld). Bolivia.
Jatropha Curcas L. Sp. PL 1006. 1753; 77. Castiglionia lobi
R. & P. Prodr. 139. pi. 37. 1794.
Shrub or small tree sometimes several meters high with orbicular-
ovate acutely and shallowly 3-5-lobed leaves or these merely sinuate
undulate, openly cordate at base, and greenish-yellow flowers, the
petals cohering nearly medially, in slender-peduncled cymes shorte
than the foliage; leaves 6-15 cm. long and broad, early tomentulose,
later puberulent only on the nerves beneath where paler and reticu-
late-venose; bracts lanceolate, 4-8 mm. long, pubescent as also the
pedicels, the ovate subobtuse sepals sparsely so, nearly free, the male
3.5 mm. long, deciduous, the female acuminate and accrescent
beneath the large drupe-like (to 4 cm. long) capsule; petals lightl]
villous within, disk glands distinct, outer stamens nearly free; ovarj
glabrous, stigma bifid; seeds to 2 cm. long, oblong-ellipsoid, pale
with dark striae and prominently reticulate. Illustrated, Pax, I.e. 78.
Widely distributed or cultivated; the seeds contain an odorles
oil that is very active as a cathartic and used for soap and lubricating;
4-5 seeds are said to cause death but roasting destroys their harmful-
ness. Bark papery; latex translucent (Williams).
Piura: Serran, Huancabamba, Stork 11383. Cajamarca: (fide
Herrera). San Martin: Tarapoto, Williams 5855. Lima: Surco,
Ruiz & Pavdn. Loreto: Pinto-cocha, middle Nanay, Williams 816.
Near Iquitos, King 249. Pebas, Williams 1582. Leticia, Williams
FLORA OF PERU 161
3062. Cuzco: Valle de Santa Ana, Raimondi. Santa Rosa, Soukup.
South America to Mexico and the West Indies. "Pifion," "pinol."
Jatropha gossypifolia L. Sp. PL 1006. 1753; 26.
Tall herb or fleshy-ligneous below marked by the filiform-dis-
sected gland-tipped stipules, their divisions simulating the branched
gland-tipped trichomes that sparsely ornament the long slender
petioles; leaves slightly cordate at base, more than medially 3-lobed
or sometimes with 2 extra shorter lobes, somewhat pubescent or
glabrous, the lobes acute, entire or often glandular-denticulate;
flowers few, in shortly peduncled cymes with linear gland-margined
bracts; sepals 5-7 mm. long; petals obovate, the obovate purplish
petals little longer; stamens usually 8; ovary pubescent, the glabrate
3-sulcate capsules 1 cm. thick, the oblong seeds brown. As in a
number of other species, the oil of the seeds was at one time used
as a purgative. Illustrated, Bot. Reg. pi. 746.
San Martin: Clearing, Chazuta, King 3974. Tarapoto, Williams
5423; 5544- Loreto: Near Iquitos, Williams 1387. In forest,
Caballo-Cocha, Williams 2215. Leticia, Williams 3033. To Mexico
and the West Indies. "Pifion negro," "pinon."
Jatropha Humboldtiana McVaugh, Bull. Torrey Club 72: 35.
1945. J. peltata HBK. Nov. Gen. & Sp. 2: 104. 1817, vix Sesse"
ex Cerv., 1794.
In general like J. Weberbaueri; leaves 3-5 cm. across, 5-7-lobed,
glaucescent and glandular-ciliate; sepals of male flowers obtuse,
minutely denticulate, 1.5 mm. long; disk glands free; anthers
elongate; capsules 14 mm. long, 12 mm. thick, globose-ovoid.
The earliest described species in a group of five (or more) that are
much alike except for development of glands and slight differences
in flowers, especially in size of parts.
Cajamarca: Jae"n de Bracamoros at Tomependa, Humboldt &
Bonpland, type. Jae"n to Bellavista, 700 meters, Weberbauer 6211.
Jatropha macrantha M. Arg. Linnaea 34: 209. 1865; 48.
Spreading stockily branched glabrous shrub sometimes a meter
high, the more or less fleshy branches conspicuously marked by
the callus-margined scars of the fallen petioles; leaves 10-12 cm.
wide, 9-10 cm. long, deeply cordate at base, more than medially
3-lobed, the lobes broadly ovate, more or less imbricate, acute, entire;
stipules glanduliform; cymes shortly peduncled, the few large
162 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
scarlet-red flowers capitately congested, the lower bracts foliaceous,
ovate-lanceolate, acuminate, to 10 mm. long; sepals of male flowers
oblong-ovate, acute, glandular-dentate, free, 4-5 mm. long; petals
to 2 cm. long, oblong, obtuse, clawed, free; glands distinct; stamens
10, the outer shorter, the inner long-monadelphous; ovary glabrous.
Typical plant for the western middle slopes; the thick stems become
decumbent toward the base, the showy scarlet blossoms appearing
during the dry period tardily accompanied by the leaves which
early are brownish and sticky as the sap of the fleshy branchlets.
Illustrated: habital photograph Weberbauer, opposite p. 163; Pax
& Hoffm. I.e. (flowers).
Cajamarca: Santa Valley, 2,200 meters, Weberbauer, 173. Lima:
Matucana, 2,300 meters, 2915, also Weberbauer, 268 and others; 163,
164, 166; Stork & Norton 9136; Soukup 2135; Rose & Rose 18643.
Ancash: Lomas de La Chay,GoodspeedExped. 9217 (det. Johnston).
Huanuco: Near Huanuco, Ruiz & Pav6n, type. Arequipa:
Huasachi, Balls 5878 (det. Killip). Laspinas, Goodspeed Exped.
22145 (det. Johnston). Socabaya, 2,300 meters, West 7140 (det.
Wheeler). "Huancarpo de Canta" (Ruiz & Pavon), "barbasco"
(Balls).
Jatropha multifida L. Sp. PI. 1006. 1753; 40.
Slender-stemmed tall shrub or tree-like with long-petioled pal-
mately divided leaves, the many lanceolate acuminate sometimes
lobed divisions extending nearly or quite to base, often 12-15 cm. long,
1.5-2.5 cm. wide; stipules eglandular, setaceously dissected, 1-2 cm.
long; peduncles elongate, the cymes of small red flowers several cm.
wide, the male petals to 5, the female to 7 mm. long; capsules sub-
pyriform, smooth, yellow, nearly 3 cm. long. Illustrated, Mart.
Fl. Bras. 11, pt. 2: pi. 69.
This distinctive species widely cultivated in warmer and tropical
regions was apparently found established as noted by Williams.
The young leaves are said to be used as a vegetable while the seeds
at one time were employed medicinally.
San Martin: Morales, Tarapoto, cultivated, Williams 5692.
San Roque, weed, Williams 7349. Loreto: Near Iquitos, Williams
1528. Brazil to Texas and the West Indies. "Pinon."
Jatropha Weberbaueri Pax & Hoffm. Pflanzenreich IV. 147:
45. 1910.
Entirely glabrous shrub or small tree, the branches rather fleshy,
the leaves centrally peltate, the stipules finely dissected, gland-
FLORA OF PERU 163
tipped, 5 mm. long; petioles 6-8 cm. long, somewhat pruinose as
the long-peduncled cymes, these corymbiform and with scarious
lanceolate caducous bracts; leaves 8-9 cm. across, palmately 5-lobed,
truncately emarginate at base, the sinus between the short ovate
setaceous-acuminate eglandular entire (or few dentate) lobes nearly
rectangular-obtuse at base; male sepals medially connate, acuminate,
4 mm. long, petals obtuse, free, 18 mm. long; filaments 8, anthers
5-6 mm. long, disk glands free; female sepals free, 7-8 mm. long,
accrescent beneath the fruit; petals broader, 9-10 mm. long, disk
irregular; styles slender, connate at base. J. nudicaulis Benth. of
Ecuador has small stipules bearing very long caducous trichomes.
Illustrated, Pax & Hoffm. I.e. 44.
Piura: Valley of the Rio Quiros, 1,300 meters, Weberbauer 6355.
Cajamarca: Cutervo, Stork & Horton 26327 (det. McVaugh).
Huanuco: Luya, Tupen, valley of the Maranon, Weberbauer 4779,
type. Ecuador.
32. CNIDOSCOLUS Pohl
Reference: McVaugh, Bull. Torrey Club 71: 457-474. 1944.
Like Jatropha but apetalous and the pubescence at least in part
stimulose (often long and rigid) trichomes. Glands sometimes
present at top of petiole. Stem pith characteristically white and
divided into thin transverse plates. Styles 3-5 times dichotomous
with slender tips. Female flowers ordinarily with rudimentary
stamens and male with filiform staminodia. Annular gland usually
raised above base of staminal column. After McVaugh, from whose
thoughtful and clear synopsis the following key is compiled. He
has called attention to the fact that the list of species of Jatropha
given by Pax and Hoffmann in Pflanzenfam. ed. 2. 19c: 164. 1931
that are to be referred to this genus are not there validly transferred
since previous publication reference was omitted. Once I thought
to accept transfer in the case of their own Peruvian species, since
the implied publication source for these seemed obvious to me.
Apical petiolar glands large, depressed, solitary or geminate; stamens
usually 10, all monadelphous, the filaments biverticillate.
Leaves broadly 5-lobed about one-third, the lobes rounded-
apiculate or merely acute C. pyrophorus.
Leaves 3-7-lobed at least medially, the lobes shortly acuminate.
C. jaenensis, C. tubulosus.
164 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Apical petiolar glands usually clustered, papilliform, usually more
or less unequally elongate, if lacking, anthers many or if 10,
outer filaments distinct.
Petioles and leaf-blades subequal, the latter lobed or dentate.
Leaves at least medially lobed; stamens 8-10, only inner of
2 whorls of filaments coherent C. urens.
Leaves merely lobate if at all; stamens 15-19 in 3-4 whorls.
Stamens 18 or 19 in 4 verticils; petioles usually basally bicu-
leate C. diacanthus.
Stamens 15-17 in 3 verticils. .C. basiacanihus, C. peruvianus.
Petioles much shorter than subentire leaf -blades. . .C. hypoleucus.
Cnidoscolus basiacanthus (Pax & Hoffm.) Macbr., comb. nov.
Jatropha basiacantha Pax & Hoffm. Pflanzenreich IV. 147: 90. 1910.
Younger branches glabrous and estimulose; petioles glabrous but
stimulose (as the leaves), including a ring of trichomes 8 mm. long
at their base that, becoming indurate, persists after the petiole
falls; leaves 6 cm. long and broad, rotund-ovate with cordate base,
obtuse, elobate but irregularly denticulate; stipules not seen; cymes
unknown; male flowers (young) about 8 mm. long, scarcely to one-
half connate, puberulent or glabrate, obtuse, fertile stamens 15 in
3 verticils, the column sparsely pilose at base, disk glands nearly
free, glabrous; female flowers with lanceolate glabrous free sepals
12 mm. long, glabrous lobulate disk, squamate within, appressed
pubescent ovary, the styles palmately cleft. Unless by the "ring
of trichomes" at petiole-base this seems scarcely separable from
C. peruvianus. F.M. Neg. 5381.
Lima: Matucana, 1,700 meters, Weberbauer 1694, type.
Cnidoscolus diacanthus (Pax & Hoffm.) Macbr., comb. nov.
Jatropha diacantha Pax & Hoffm. Pflanzenreich IV. 147. vii: 399.
1914.
A 3-meter shrub with stout subfleshy estimulose branches;
petioles soon 3-5 cm. long, glabrous, stimulose with 2 indurate
spines persisting at base; leaves rotund-ovate, cordate, 10-12 cm.
long, 9-10 cm. wide, glabrous but sparsely stimulose above, mem-
branous, acute or acuminate, irregularly sinuate-dentate; cymes
capitate-congested, many-flowered; male sepals connate to two-
thirds into a cylindrical tube, velvety pubescent, 11-12 mm. long,
the obovate lobes obtuse; fertile stamens 18-19, 4-verticillate, the
FLORA OF PERU 165
column ciliolate; disk glands glabrous; female flowers unknown;
ovary densely velutinous. Allied to C. basiacanthus but petioles
biaculeate basally and stamens 4-verticillate (authors).
Apurimac: Between mouth of Rio Pachachaca and Rio Pampas
on the Rio Apurimac, 1,000 meters, savannah, Weberbauer 5903,
type. Pachachaca Valley, Goodspeed Exped. 10562 (det. Standley).
Rio Pampas, West 3684 (det. Johnston). Huancavelica: Mantaro
Valley, Stork & Horton 10402 (det. Standley). "Mula-huanuchi,"
"huanarpo."
Cnidoscolus hypoleucus (Pax) Pax & Hoffm. Pflanzenfam.
ed. 2. 19c: 164. 1931. Jatropha hypoleuca Pax, Pflanzenreich IV.
147: 96. 1910.
A shrub with unarmed but stoutly stimulose branches; petioles
about 1 cm. long, densely stimulose, otherwise nearly glabrate;
leaves broadly ovate or subobovately rounded, rounded-obtuse or
subcuneate at base, acute, subentire or lightly denticulate, opaque
above, densely white tomentose beneath, subcoriaceous, margins
and especially midnerve of the pinnate nerves stimulose, 6-10 cm.
long, 6 cm. wide; cymes depauperate, sessile, glabrous; male sepals
18 mm. long, medially connate, white, puberulent, linear; stamens
to 17, glabrous, monadelphous, 3-verticillate, staminodia 2-3,
filiform; disk annulate; female flowers unknown; capsules verruculose,
about 1 cm. long, the lustrous brown applanate seeds 7 mm. long
with appressed bilobed caruncle. F.M. Neg. 5392.
Amazonas: Stony outcrops, Tupen, 800 meters, Weberbauer 4797 >
type.
Cnidoscolus jaenensis (Pax & Hoffm.) Macbr., comb. nov.
Jatropha jaenensis Pax & Hoffm. Pflanzenreich IV. 147. vii: 400.
1914.
A 2-meter shrub, the shortly puberulent branchlets stimulose as
the petioles (to 2 dm. long), cymes and leaves especially beneath on
the nerves; apical petiolar glands large, depressed, binate; leaves
more or less deeply cordate, thin-membranous, nearly glabrous
above, pubescent beneath, 9-28 cm. long, 10-33 cm. wide, more
than medially 3-5-parted, the ovate or oblong shortly acuminate
lobes entire or repand-crenate; cymes with short peduncle 6-8 cm.
long; buds pyriform, opening to 1.5 cm. long, 2 cm. broad, white,
the oblong lobes 8-10 mm. long, 5-6 mm. wide; stamens 10, biverticil-
late, monadelphous; staminodia 3, filiform, glabrous; column pubes-
166 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
cent; disk urceolate, crenulate; female calyx patellately persisting;
ovary densely stimulose; capsules about 8 mm. long. Affine J. tubu-
losa but cymes shorter, flowers large and staminal column pubescent
(authors). These differences seem scarcely of specific value.
Cajamarca: In bushes, Valle de Shumba, Ja&i, 800 meters, Weber-
bauer 6234, type.
Cnidoscolus peruvianus (M. Arg.) Macbr., comb. nov. Jatro-
pha peruviana M. Arg. in DC. Prodr. 15, pt. 2: 1096. 1866; 90.
Younger branchlets, petioles and leaf-nerves more or less hispid
with stinging trichomes; petioles not aculeate, as long as the broadly
ovate leaves, these 8-14 cm. long, 9-12 cm. wide, entire or 3-lobed,
cordate at base, membranous, early softly villous, finally glabrescent;
stipules triangular, glandular-denticulate; cymes small, shorter than
the petioles, corymbiform, bracts subulate-acuminate; male sepals
nearly glabrous, two-thirds connate, 15 mm. long, fertile stamens,
about 15 in 3 verticils, the long stamen column pubescent at base,
the anthers 2 mm. long, longer than broad; female flowers unknown.
My specimen from an open spreading plant with 3-5 stems woody
below, rubber-like above, less than 1 meter high. As noted by
McVaugh see my remark at end of generic description the listing
of this species without indicating its original publication did not
constitute transfer by Pax and Hoffmann. F.M. Neg. 8495.
Lima: Acotama, Pavdn, type. Rocky hillside, Chosica, 2865.
"Huanarpo."
Cnidoscolus pyrophorus (Pax) Macbr., comb. nov. Jatropha
pyrophora Pax, Pflanzenreich IV. 147: 101. 1910.
Stimulose but otherwise glabrous; leaves 2 dm. wide, 16 cm.
long, cordate at base, 5-lobed one-third, membranous, stimulose,
especially above the short rounded obtuse lobes, lightly denticulate
and ciliate; cymes densely flowered, corymbiform, shortly peduncled;
male sepals to 1 cm. long, connate to middle, ovate, subobtuse,
merely stimulose, white; stamens 10, monadelphous, in two verticils;
staminodia filiform, 3 or 4, stamen column villous at base; disk
annulate, glabrous; capsule 10 mm. long, densely stimulose; seed
caruncle bilobed. A meter shrub allied to J. urens but with totally
different androecium (Pax). Listed by McVaugh, I.e. 472, with
query, as allied to C. tubulosus. F.M. Neg. 5400.
Cajamarca: Open formation, cacti, etc., below Santa Cruz, 1,300-
2,000 meters, Weberbauer 4129, type.
FLORA OF PERU 167
Cnidoscolus tubulosus (M. Arg.) Johnst. Contr. Gray Herb.
68: 86. 1923. Jatropha tubulosa M. Arg. Linnaea 34: 212. 1865;
108.
Sometimes a tree, pubescent as well as stimulose, including the
well-peduncled and many-flowered cymes; petioles 10-15 cm. long;
leaves 18 cm. long, 20-24 cm. wide, cordate at base, 3-7-parted to
the middle or deeper, the ovate or obovate shortly acuminate lobes
entire to minutely or spinescent dentate, puberulent above, ashy
pubescent beneath; male calyx at anthesis pyriform, 5 mm. long,
later 7-8 mm. long, tomentulose; fertile stamens 10 in 2 verticils,
sterile 5, column glabrous, female calyx shortly 5-lobed, the persisting
basal part cupulate, capsule 10 mm. long, verruculose, stimulose.
The original author designated vars. septemloba, quinqueloba and
triloba, the last with entire but minutely denticulate lobes. To
avoid futile reference work it may be noted that there is no explana-
tion by the second authority justifying his transfer.
San Martin: Near Moyobamba, Woytkowski 35301 (distr. as
J. jaenensis). Bolivia to Mexico.
Cnidoscolus urens (L.) Arthur, Torreya 21: 11. 1921. Jatropha
urens L. Sp. PL 1007. 1753; 97.
Herbaceous or suffrutescent, to 1.5 meters, the petioles, leaves
and cymes more or less pilose and stimulose; stipules small, laciniate-
dentate; leaves often broader than long, usually about 12 cm. long,
truncate to cordate at base, 3-5-lobed to the middle or deeper, the
lobes ovate-lanceolate, acute, angulate-dentate, sinuate or pinnatifid;
cymes corymbiform, shortly peduncled, rather few-flowered; male
calyx to 1 cm. long, slender, constricted at throat, tomentulose and
stimulose, lobed to middle; stamens 8-10, outer nearly free, villous
at base; staminodia none or 2-3; ovary acute, stimulose-puberulent;
styles slender, 2-3-cleft above, the divisions sometimes divided; cap-
sules 10-12 mm. long, seeds 8 mm. long. Variable in shape of foliage.
Most of the following material has been determined in herbaria as
C. tubulosus but it does not have the entire depressed apical petiolar
glands of that species; perhaps the character is unimportant.
San Martin: Juanjui, Klug 3898. Tarapoto, Williams 5906.
Junin: La Merced, brushy river flat, 5330. Cuzco: Hacienda Pabel-
lon, Valle" de Lares, Herrera 779; Diehl 2487; Weberbauer 7936.
Pumachaca, Valle de Santa Ana, 1,400 meters, Herrera 3322 (Diehl).
Echarate, Goodspeed Exped. 10476. Valle del Rio Villcanota, Mexia
168 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
8051A. Warm America to Mexico and the West Indies. "Angel-
tauna" (Herrera), "vara del Angel" (Mexia).
33. CUNURIA Baillon
Reference: Pax, Pflanzenreich, IV. 147: 16-17. 1910.
Glabrous trees with alternate entire coriaceous pinnate-nerved
leaves, biglandular above at base, and dioecious apetalous flowers,
sessile and ternate in small congested cymules lateral or apical.
Male flowers with 5 strongly imbricate sepals, no disk, 10 stamens,
the short free filaments affixed to obviously elevated receptacle,
anthers dorsifixed, ovary rudiment villous; female disk cupulate,
glandular-lobate or none, ovary 3-celled, style short, bifid, reflexed,
ovules solitary. Capsules large, subglobose, exocarp fleshy, endocarp
ligneous, dehiscing into bivalved cocci with large ovoid seeds.
Baillon published his name, based on a native one noted by Spruce
in Brazil, a few months before Mueller proposed another in reference
to the great resemblance of the foliage to that of Clusia.
Cunuria Spruceana Baill. Adansonia 4: 288. August 1864; 16.
Stoutish branchlets apically clothed with stipules 5-6 mm. long;
petioles 1-3.5 cm. long, less than 2 mm. thick; leaves broadly elliptic,
or somewhat oblong-obovate, obtuse or subobtuse both ends, 6-14
cm. long, 4-8 cm. wide, rigid, slightly lustrous, the many parallel
nerves prominent, especially beneath; cymes to about half as long
as leaves; bracts minute; male calyx to 2 mm. long, subglobose,
scarcely 5-parted more than medially, the broadly ovate lobes obtuse;
rudimentary ovary 3-parted; capsules 4 cm. long, seeds 22 mm. long,
17 mm. broad, brown, lustrous. Buttressed trees to 35 meters tall,
the trunk to 8 dm. in diameter, fide Schultes, Bot. Mus. Leaflets
Harvard 13: 269. 1949. The similar C. crassipes M. Arg., perhaps
occurring within Peru, has petioles 2 mm. thick or stouter, much
smaller capsules. Illustrated, Baldwin, Jr. & Schultes, I.e. 12.
pi. 45.
Loreto: Mishuyacu, (King 1312; 1325; 340, det. Schultes).
Colombia; Amazonian Brazil. "Cunuri" (Spruce).
34. MANIHOT Adans.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. ii: 1-99. 1910
and I.e. xiv. Add. 6: 44-49. 1919.
Usually smooth glaucescent and more or less pruinose shrubs
with alternate petioled deeply lobed leaves or these rarely sessile
FLORA OF PERU 169
or undivided. Stipules sometimes foliaceous. Flowers monoecious,
apetalous, racemose or paniculate, the racemes terminal or from the
upper axils, the female often few or solitary near the base and long-
and stoutly pedicellate, the shortly pedicelled male much more
numerous, their more or less campanulate 5-parted calyx often
colored, the lobes imbricate or contorted. Stamens 10, biverticillate
between lobes or glands, the filaments free, the anthers dorsifixed,
longitudinally dehiscent. Ovary 3-celled, the basally shortly connate
styles variously lobed or dilated ; ovules solitary. Capsules separating
into bivalvate cocci, the seeds carunculate. The name is of native
origin.
At least two species, notably M. esculenta Crantz and M. dulcis
(J. F. Gmel.) Pax are widely cultivated in tropical regions for the
tuberous roots that yield a starchy meal known as Cassava, or,
particularly in Peru, as Yuca; from it is prepared the Tapioca of
commerce. Cutler and Cardenas in their important account of the
Andean beverage "Chicha," Bot. Mus. Leaflets, Harvard 13: 33-60.
1947, give M. esculenta as sometimes employed in its preparation.
Leaves deeply parted, at least some of them; anthers elongate (un-
known in type of M. Pavoniana) except M. esculenta.
Bracts 6 mm. long or longer; calyces in types puberulent even
without, somewhat globose, unless M. Weberbaueri.
Leaf-divisions oblong-lanceolate, 3-5 cm. wide, glaucous be-
neath; calyx about 1.5 cm. long M. peruviana.
Leaf -divisions obovate, 5-7 cm. wide, concolor; calyx about
1 cm. long M. Weberbaueri, M. Pavoniana.
Bracts small, caducous; calyces in types glabrous without, usually
campanulate, pubescent in var. of M. dukis.
Leaves or their divisions lanceolate-spathulate.
Anthers short, scarcely two times longer than wide; capsules
wing-angled, sometimes slightly M. esculenta.
Anthers elongate; capsules merely angulate above. M. dulcis.
Leaves or their divisions broadly ovate M. Weberbaueri.
Leaves all entire, linear; anthers elongate M. linearifolia.
Manihot dulcis (J. F. Gmel.) Pax, Pflanzenreich IV. 147. ii:
71. 1910. Jatropha dulcis J. F. Gmel. Onomat. Bot. 5: 7. 1772-78.
M. amazonica Ule, Verh. Bot. Ver. Brandenb. 50: 83. 1908. M.
leptophylla Pax & Hoffm. (?) I.e. M. palmata var. ferruginea M. Arg.
in DC. Prodr. 15, pt. 2: 1063. 1866.
170 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Resembles the closely related M. esculenta but the roots said to
be always sweet, not poisonous, the anthers many times longer than
broad, ovary subterete and the capsule only lightly angled above;
leaves deeply 3-13-parted; panicles many-flowered; calyx 12-14 mm.
long; capsule smooth, drying rugose. Said to comprise several forms,
at least the subscandent more or less pubescent Peruvian (var.
diffusa (Pohl) Pax) perhaps a distinct species (Pax). Flowers
usually greenish-white or yellowish.
Variable or there are a number of distinct entities; cf. Croizat,
Rev. Arg. Agron. 10: 221-222, 225. 1943, who allies M. amazonica
Ule, Verh. Bot. Ver. Brandenb. 50: 83. 1908, with M. brachyloba M.
Arg., 74, and M. tristis M. Arg., 59, Pax, I.e. 59, both however with
shorter anthers, and who thought the former to be distinct by the
pendulous flowers, calyx completely glabrous even within (at least
the male) and distinguished yet another form (M. leptophylla Pax
& Hoffm., I.e. 57) in Ecuador, similar but sepals ciliate and the leaves
pilosulous beneath. Much investigation is necessary before the
significance of these characters which appear in bewildering maybe
ecotypic variations (see Pax and Hoffmann key) can be determined;
it seems probable that in this genus the same thing is happening as
has been shown to occur in the case of Hevea. The race or species
M. amazonica is said to be common in the Alto Acre where scandent,
the flowers purple. The var. ferruginea (M. Arg.) Pax, rusty pubes-
cence existing to branchlets and petioles, seems readily placed within
the soon glabrate var. diffusa. F.M. Neg. 5432 (M. amazonica).
Huanuco: Cuchero, Poeppig 1523 (var. diffusa, a little puberulent,
but short anthers, glabrous calyx); also Poeppig 1410 (type, M.
palmata, var. ferruginea M. Arg.). San Martin: Juanjui, flowers
yellowish and violet, Klug 3861 (det. Standley, M. esculenta).
Pongo de Cainarachi, flowers cream and lilac, Klug 2662. Loreto:
Gamitanacocha, Rio Mazan, flowers scarlet, Schunke 291 (det.
Croizat, M. esculenta or M. tristis). Aguaitia, 170 meters, Woytkow-
ski 34448 (var. diffusa). Cuzco: Cosnipata, Vargas 007387 (var.
diffusa). Rio Ucayali, Skutch 5009, det. Croizat. Without locality,
Ruiz & Pavdn (var. diffusa). Colombia to the Guianas and southern
Brazil. "Sacharuma" (Schunke).
Manihot esculenta Crantz, Inst. Herb. 1: 167. 1766; 67. M.
utilissima Pohl, PL Bras. Icon. 1: 32. pi 24. 1827; 67.
Slender-stemmed glaucous-pruinose shrub usually 2-3 meters
high from elongate tubers; petioles finally longer than the leaf-
FLORA OF PERU 171
blades, these glabrous or minutely puberulent on the nerves beneath
and, except for the uppermost, deeply 3-7-parted with spathulate-
or linear-lanceolate acutely acuminate divisions 8-17 cm. long, 1-5
cm. wide, gradually attenuate to base where confluent into a disk
about 2 cm. broad; stipules 5-7 mm. long, lanceolate, setaceous-
acuminate, entire or 1-2-lacinulate, puberulent; peduncles usually
as long as the petioles, slender, equaling the panicle branches; bracts
linear-lanceolate, entire, articulately deciduous above the base;
male pedicels 4-7 mm. long, the deflexed-spreading female attaining
2 or more cm. ; calyx retuse at base, campanulate, glabrous without,
puberulent within above, more than medially 5-parted, about 10 mm.
long; disk and filaments glabrous; anthers 1.25-2 mm. long, apically
hispidulous; ovary glabrous, 6-wing-angulate; capsules 1.5 cm. long,
globose-ellipsoid, rugulose-asperous, the 6 wings undulate-crenulate;
seeds marble-spotted. Said to be native to Brazil but long cultivated
in nearly all tropical regions for its starchy tubers, and in Peru
between 1,500 and 2,200 meters (Weberbauer). According to Pax
the tubers exceptionally attain 6 dm. and weigh 4 kg. while Peckolt
recorded a much larger example; ordinarily however they are about
half as large. They are prepared by boiling or roasting and only
then are free from a poison that is present in the raw state in greater
or lesser degree. Illustrated, Pflanzenreich I.e. p. 68; Mart. Fl. Bras.
11, pt. 2: pi. 65.
Huanuco: Weed in forest clearing, Pampayacu, 5074- Junin:
Thicket, sandy valley floor, La Merced, 5472; Killip & Smith 23791.
Ayacucho: Escaped near Kimpitiriki, Killip & Smith 22911;
22964. Aina, vine, Killip & Smith 22712 (det. Croizat, M. peruviana).
Loreto: Mishuyacu, near Iquitos, Klug 863. Trail to San Juan,
principal root crop, Mexia 6494- La Victoria, Williams 2797; 3102.
Caballo-Cocha, Williams 2491 ; 2492. Lower Rio Nanay, Williams
361. Puerto Arturo, Williams 5095. Yurimaguas, Poeppig 2178.
Cuzco: Valle de Santa Ana, 2,000 meters, Valles de Marcapata and
Lares (Herrera). Brazil. "Yuca," "r'umu" (Herrera), "yuca
amarillo," "yuca blanca."
Manihot linearifolia M. Arg. Flora 55: 43. 1872; 96.
Entirely glabrous, the branches slender, the internodes 3-5 cm.
long, the subsessile entire linear leaves attenuate to the acutely
acuminate mucronate tip, 10-14 cm. long, 6-9 mm. wide; stipules
subulate-setaceous; racemes spiciform, many-flowered, about 8 cm.
long, the solitary basal female flower on pedicel 10 mm. long, the
172 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
male pedicels 1-2 mm. long; bracts lanceolate-subulate-acuminate;
calyces glabrous, the male 7 mm., the female 11 mm. long; filaments
short, glabrous; anthers three and a half times longer than broad;
disk hypogynous, undulately crisped; ovary ellipsoid, glabrous.
F.M. Neg. 19507.
Peru(?): Without locality, Haenke, type.
Manihot Pavoniana M. Arg. Linnaea 34: 205. 1865; 65.
Stems lax or possibly scandent, the younger parts including the
leaf -nerves beneath even in age more or less puberulent or pubescent;
petioles 5-9 cm. long; leaves membranous, concolor, deeply 3-parted
or entire, the obovate-rhombic acutely cuspidate or acuminate lobes
13 cm. long, 5-7 cm. wide, narrowed toward base; smaller undivided
leaves rounded at base or somewhat cordate, ovate; stipules decid-
uous; racemes peduncled, compact, the lanceolate-subulate entire
bracts 6-10 mm. long, 1-1.5 mm. wide, rusty pubescent as the
pedicels; male calyx finally globose-ellipsoid, 8 mm. long, puberulent
without, pubescent within; filaments as connective pubescent; ovary
subangled. The Indians attribute aphrodisiac properties to the
root of this plant (Ruiz & Pavon). F.M. Neg. 5464.
Huanuco: Muria, Ruiz & Pavdn, type. Bolivia? "Simayuca/
Manihot peruviana M. Arg. Linnaea 34: 206. 1865; 29. M.
heterandra Ule, Verh. Bot. Ver. Brandenb. 50: 84. 1908.
Stems lax or possibly scandent, apically puberulent, obtusely
angled; petioles as long as the leaf blades, these early appressed
puberulent above but glabrous in age, softly pubescent and glaucous
beneath, all except the ovate acuminate upper ones 3-5-parted nearly
to base, the oblong- or lanceolate-ovate acutely acuminate divisions
8-12 cm. long, 3-5 cm. wide; stipules 5-6 mm. long, biparted into
linear-lanceolate acuminate puberulent lobes; racemes sessile, 3 cm.
long, the lanceolate entire bracts 8-12 mm. long, to 3 mm. wide;
calyces finally 1.5 cm. long, ovoid and 5-angled above where puberu-
lent without, tomentulose within, the male 5-parted only medially
and with glabrous disk; filaments dimorphic densely villous especially
those alternate with the sepals; connective hispid; anthers elongate;
ovary glabrous, prominently costate. The Spruce specimen is the
type for both names. F.M. Neg. 5465.
San Martin: Tarapoto, Spruce 4287, type; Ule 6635 (det. Ule,
M. heterandra}. Loreto: Lower Rio Huallaga (Williams 5323, det.
Croizat). Brazil?
FLORA OF PERU 173
Manihot Weberbaueri Pax & Hoffm. Pflanzenreich IV. 147.
xvii. Add. 7: 194. 1924.
Branchlets apically hirtellous, soon glabrate; petioles 3.5-10 cm.
long; stipules caducous; leaves entire or 3-parted, the former about
12 cm. long, 5-8 cm. wide, ovate, cuspidate, truncate-rounded at
base, the latter 8-15 cm. long, 14-16 cm. wide, the short entire elliptic
lobes setaceous-acuminate, sparsely pubescent near base and on
nerves; secondary nerves about 6; inflorescence few-flowered, puberu-
lent; bracts to 1 cm. long, 1-1.5 cm. wide, linear, remotely lacinulate,
acuminate; male calyx 5-parted, glabrous except for 2 pilose lines
within; filaments glabrous, disk pilose; female calyx completely
glabrous. Type from a 4-meter shrub.
Huancavelica: Valle de Rio Mantaro, near Pampas, grass steppes,
1,300 meters, Weberbauer 6517, type. "Keytayuca."
35. PAUSANDRA Radlk.
Reference: Lanjouw, Meded. Bot. Mus. Rijksuniv. Utrecht 38:
758-769. 1936.
Small or medium glabrate lactiferous trees with bistipulate
petioled glandularly dentate often cuneate-spathulate leaves 2-4-
glandular at base and glomerulate dioecious or monoecious flowers
arranged in slender spikes or sometimes somewhat panicled, bibracte-
olate in the axils of biglandular bracts. Calyx of the male with 5 (3)
unequal lobes, imbricate as those of the more or less gamopetalous
corolla, this barbate inside at the throat, the extra-staminal urceolate
disk sometimes deeply 5-lobed; stamens 3-7, usually 6 (1 central)
with oblong anthers; rudimentary ovary none. Female flowers
similar but the apparently free petals medially barbate; ovary
pubescent, 3-celled, the ovules solitary; styles 2-lobed. Capsules
with 3 cocci, loculicidally dehiscent, the somewhat marbled seeds
oblong, caruncled. After Lanjouw, who remarks: perhaps several
of the old and new species can be united, as one can find often only
small differences, but for the present I think it advisable to keep
them separate.
Flowers many, usually 15-40, in each glomerule, the male spikes
often as long as or longer than the leaves.
P. densiflora, P. quadriglandulosa.
Flowers few in each glomerule, the male spikes often much shorter
than the leaves.
174 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Leaf -serrations prominent, incurved; lateral nerves 25-30; basal
glands usually 4 P. Trianae.
Leaf -serrations little or not incurved; lateral nerves 15-21 or
rarely more; basal glands 2 P. Morisiana.
Pausandra densiflora Lanj. Meded. Bot. Mus. Rijksuniv.
Utrecht 38: 766. 1936.
Branchlets stout, the younger as the lanceolate acute stipules
(about 8 mm. long) and petioles, these 1-3 cm. long, appressed
pubescent; leaves spathulate-oblanceolate, long-cuneate to base,
shortly and acutely acuminate at apex, rigid-membranous, shortly
and sparsely appressed pubescent, especially beneath on the 26-29
lateral nerves, about 3-4 dm. long, a third as wide, coarsely serrate
with glandular more or less incurved teeth, the 2 basal glands about
1.5 mm. long, 1 mm. wide; male spikes 4-5.5 dm. long, the many-
flowered glomerules sparsely short-pubescent (both bracts and
sepals); petals barbate within at throat; stamens mostly 6; female
flowers unknown. Distinguished from all species by the very long
spikes and the dense glomerules (Lanjouw); however, cotype speci-
men matches closely in these respects type of P. quadriglandulosa.
Illustrated, Lanjouw, I.e. p. 765.
Rio Acre: Mouth of Rio Macauhan, Krukoff 5464, type.
Pausandra Morisiana (Casaretto) Radlk. Flora 53: 92. pi. 2.
1870; 761. Thouinia Morisiana Casaretto, Nov. Stirp. Bras. Dec. 9:
75. 1845.
Young branches, petioles, these 1-6 cm. long, rugulose and
slightly thicker at apex, and leaves beneath sparsely appressed
pubescent, soon glabrescent; leaves oblanceolate or -oblong, long-
cuneate to base, shortly acuminate, usually 1.5-3 dm. long, about
half as wide, chartaceous to coriaceous, glabrous above, nerves 15-21,
basal glands 2, small, subsessile, subpatelliform; male spikes 5-10
(26) cm. long, densely appressed pubescent rachis with few-flowered
glomerules at short distances; bractlets and buds pubescent but
unequally 5-lobed; calyx glabrous at anthesis as the basally connate
petals without except the ciliolate margins but these barbate within
about one-fourth above the base; disk urceolate, slightly undulate;
stamens 6, anthers ovate; female spikes 7-8 cm. long, sepals pubes-
cent, 4 mm. long, petals free, ovary pubescent; fruits about 22 mm.
high, the obscurely marmorate brownish seeds about 16 mm. long,
11 mm. broad. Similar Amazonian species include P. macrostachya
FLORA OF PERU 175
Ducke, 764, the type with larger leaves with about 23 lateral nerves,
spikes 2-4 dm. long, petals long-connate, and P. macropetala Ducke,
768, collected as near as Sao Paulo de Olivenga, the leaves glabrous
both sides, the disk deeply lobed. Illustrated, Mart. Fl. Bras. 11,
pt. 2: pi. 99, and Pax and Hoffmann in Pflanzenreich IV. 147. iii:
43. fig. 14.
Loreto: Mishuyacu near Iquitos, Killip & Smith 29968 (det.
Standley, P. quadriglandulosa). Balsapuerto, Killip & Smith 28607
(det. Standley, P. quadriglandulosa). Rio Acre: Mouth of Rio
Macauhan, Krukoff 5455 (det. Steyermark). To southern Brazil.
Pausandra quadriglandulosa Pax & Hoffm. Pflanzenreich IV.
147. xiv. Add. 6: 43. 1919.
Type from a shrub or a tree 3-18 meters tall, the stout floriferous
branchlets soon glabrate; petioles (uppermost) 1.5-2 cm. long;
leaves spathulate, long-attenuate to base, subabruptly acuminate,
to 3 dm. long, a third as wide, many-nerved, coarsely serrate with
incurved teeth, nearly glabrous above, lightly villosulous and paler
beneath with 4 glands above the base, the 2 inner smaller, the outer
conical; male spikes nearly or quite as long as the leaves, appressed
pubescent, the glomerules many-flowered; male flowers finally 5 mm.
long, pale yellow; calyx puberulent, lobes broadly ovate; petals
connate below, the lobes reflexed; stamens 5; female flowers and
fruit unknown. Included by Lanjouw in P. Trianae and perhaps
rightly but that as to type at least has few-flowered glomerules,
much shorter male spikes. F.M. Neg. 5406.
Rio Acre: Seringal San Francisco, Ule 9538, type. Brazil?
Pausandra Trianae (M. Arg.) Baill. Adansonia 11: 92. 1873;
763. Pogonophora Trianae M. Arg. Flora 47: 434. 1864.
Young branchlets and leaves, both surfaces, densely appressed
pubescent, soon glabrescent; petioles 1.5 cm. long or longer; leaves
ample, often 3-6 dm. long, scarcely a third as wide, oblanceolate,
narrowed to base, shortly acuminate, coriaceous to chartaceous,
coarsely serrate with incurved glandular teeth, usually 4-glandular
at base (Lanjouw), lateral nerves many; male spikes in type 3 times
shorter than leaves with small few-flowered glomerules; calyx short-
pubescent with 5 short lobes; petals connate below, tomentose
within at throat; stamens 5. According to Lanjouw disk long-
cupulate, slightly undulate, stamens 5-7, but he included here
176 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
P. quadriglandulosa and material from Central America. Illustrated,
Lanjouw, I.e. p. 763 (leaf margin). F.M. Neg. 24574.
Peru (possibly; cf. note above). Colombia; Nicaragua?
36. POGONOPHORA Miers
Reference: Pax & Hoffm. Pflanzenreich IV. 147. iii: 108-110.
1911.
Trees with alternate petioled typically entire leaves and short
axillary panicles of small glomerulate 5-merous petaliferous flowers,
the male subsessile, the female shortly pedicelled. Sepals and petals
imbricate, the latter longer and typically barbate; stamens 5, affixed
beneath thick disk, the short filaments free, the long linear anthers
apiculate; rudimentary ovary trifid, linear. Disk shortly cupulate.
Ovary 3-celled; styles nearly free, irregularly bilobed, fimbriately
stigmatose; ovules solitary. Capsules separating into bivalved cocci.
Seeds ovoid, lustrous; endosperm fleshy, cotyledons plane.
Pogonophora Schomburgkiana Miers ex Benth. in Hook.
Journ. Bot. Misc. 6: 373. 1854; 108.
Tall shrub or becoming a large tree, the branchlet tips minutely
puberulent with malpighiaceous trichomes; petioles 1-3 cm. long,
early pubescent like the branchlets, especially beneath, canaliculate,
slightly tumidulous at tip; leaves oblong-elliptic or elliptic-lanceolate,
acute both ends or shortly acuminate, often about 1 dm. long, half
as wide, sometimes 2 dm. long, three times longer than wide, cori-
aceous, glabrous or nearly, entire, the lateral nerves 7-8, slender,
the veins reticulate, little prominent; inflorescences 2-5 cm. long,
sericeous; bracts ovate, 2 mm. long; calyx lobes orbicular-ovate,
obtuse, the 2 outer shorter; petals narrowly elliptic, coriaceous,
conspicuously and rigidly barbate within, 2.25 mm. long, the male
acute, the female obtuse; interstaminal disk urceolate, 5-crenate,
thick; ovary tomentulose; capsules ellipsoid, 8 mm. long, nearly
as thick, the seeds fuscous. Collected by J. Murca Pires and G. A.
Black at Tabatinga, Brazil, near the boundary and so undoubtedly
also within Peru.
Peru (see note above). Amazonian Brazil to the Guianas.
"Aracapuri" (Pires and Black).
37. DODECASTIGMA Ducke
Trees, glabrous except the canescent tomentulose inflorescences
of 3-4- or 5-merous green flowers, the male many, fasciculate in
FLORA OP PERU 177
panicles, the female solitary and scattered in few-flowered racemes.
Calyx lobes both sexes with firm glabrous margins, imbricate as the
externally pilose petals. Disk extra-staminal, the 14-16 (or in one
species 9) stamens with free erect filaments. Ovary 3-celled, the
ovules solitary, the 3 styles bifid, often with many dentate branches.
Capsules elastically dehiscent, bivalved with large endocarp. Seeds
ecarunculate. Genus of two species; Sandwith, Kew Bull. 134-136.
1950, D. integrifolium (Lanj.) Lanj. & Sandw. of Guiana, stamens 9.
Dodecastigma amazonicum Ducke, Notizbl. Bot. Gart.
Berlin 11: 343. 1932.
Bark brown, minutely laminate; petioles 3-12 cm. long, enlarged
base and apex; leaves more or less elliptic-oblong, acute at base,
shortly and abruptly acuminate, 10-26 cm. long, 4-12 cm. wide,
margin beneath nerviform-revolute, coriaceous, lucid above, opaque
beneath, lateral nerves 10-12, prominent only beneath; panicles
3-4 dm. long, male lax, the pedicels 4-5 mm. long, flowers 6-7 mm.
long, 8-11 mm. wide; female in simple racemes, pedicels 2-2.5 cm.
long (-4 in fruit); calyx 6-7 mm. long, 10 mm. wide, 2 inner sepals
broadly appendaged; petals obtuse; capsules tomentose; seeds 2 cm.
long, 14 mm. thick. Has been collected at Sao Paulo de Olivenca
and so to be expected within Peru. Leaves entire; stamens 14-16.
Peru (probably). Amazonian Brazil.
38. TETRORCHIDIUM Poeppig
Reference: Pax & Hoffm. Pflanzenreich IV. 147. iv: 29-32. 1912.
Small slender trees with alternate stipulate entire or dentate
leaves borne on apically or subapically biglandular petioles and
axillary racemes or panicles of small glomerulate (the female some-
times solitary) apetalous flowers. Floral bracts with one conspicuous
gland. Male calyx with 3 broad lightly imbricate sepals, no disk,
3 episepalous subsessile barely free stamens, the broad 4-celled
anthers introrsely peltate; female disk cyathiform or divided into
3 petaloid scales; ovary 2-3-celled, style short, stout, bifid, stellately
spreading or nearly connate, the ovules solitary. Capsules 2-3-
parted into bivalved cocci, the globose seeds deeply foveolate.
The name refers to the 4-celled stamens. Croizat has noted, Journ.
Arnold Arb. 24 : 170. 1943, that the author of this genus is as above,
as indicated on the title page of Poepp. & Endl. Nov. Gen. & Sp.
3: 23. pi. 227. 1845 (not 1842).
178 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Disk urceolate, 3-lobed; leaves prominently reticulate.
T. macrophyllum.
Disk glands free, lingulate; leaf -reticulation fine or obscure.
T. rubrivenium.
Tetrorchidium macrophyllum M. Arg. in DC. Prodr. 15,
pt. 2: 1133. 1866; 30.
Resembles T. rubrivenium but imperfectly known; branchlets
stout, appressed pubescent as the petioles, leaf-nerves and racemes
with pale reddish malpighiaceous trichomes; leaves 1.5-3 dm. long,
6-10 cm. wide, shortly cuspidate-acuminate, entire, prominently
reticulate-veined; female racemes 10-15 cm. long, laxly flowered,
slender, naked below, the pedicels 5-7 mm. long; sepals ovate, acute,
2 mm. long, rusty pubescent without and within; disk glands connate
into a 3-lobed pubescent cup; ovary 3-celled, sericeous; styles scarcely
connate at base, deeply bifid, the branches slender. The two knob-
like glands at petiole tip are not quite opposite. Male racemes and
flowers unknown. Said to have a milky latex. Illustrated, Mart.
Fl. Bras. 11, pt. 2: pi. 71; also Pax, I.e. (female flowers).
San Martin: In woods near Tocache, Poeppig 2034, type.
Loreto: Pongo de Manseriche, Tessmann 4188. Ecuador.
Tetrorchidium rubrivenium Poepp. in Poepp. & Endl. Nov.
Gen. & Sp. 3: 23: pi. 227. 1845; 31.
Weak tree, to 12 meters high, the younger parts early minutely
appressed pubescent with simple and malpighiaceous trichomes
mixed, finally glabrescent; petioles 2-4 cm. long; leaves 10-18 cm.
long, 4-8 cm. wide, elliptic- or ovate- or obovate-lanceolate, acute
or subobtuse, cuneately narrowed to base, entire or widely dentate,
finely or not at all prominently reticulate, membranous, lustrous;
secondary nerves about 7-8 each side, slender; male panicles many-
flowered, much-branched, about equaling the leaves, the female
racemes short, commonly half as long as the petioles; male glom-
erules many-flowered; female pedicels stout; flowers fulvous pu-
berulent, the male 1.5 mm. wide, their sepals not at all or scarcely
exceeding the appressed anthers, the female sepals triangular-
ovate; disk glands free, lingulate, glabrous, sometimes as long as
calyx; ovary 2-3-celled, pubescent or finally glabrescent; style
branches subpetaloid, ovate. The apical or subapical petiolar glands
in specimens seen are opposite or one lower on same side. The
FLORA OF PERU 179
obscure T. andinum M. Arg., 30, of Ecuador, has glabrous ovary,
entire or lobulate disk. Illustrated, Pax, I.e. p. 30.
San Martin: Woods near Tocache, Poeppig 1915; 1951 (types).
Zepelacio, King 3717 (det. Standley and Croizat). Huanuco: Rio
Huallaga above Cayumba, Mexia 8322 (det. Standley). Colombia;
Brazil.
39. OMPHALEAL.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 14-22. 1912.
Shrubs, in Peru scandent, with alternate stipulate leaves borne
on petioles biglandular at apex, either oblong and pinnately nerved
or cordate and 3-5-nerved and with small monoecious apetalous
flowers, the female, if present, disposed in the center of the male
cymules. Cymule bracts often elongate, glandular-petiolate and
subfoliaceous. Disk none or obsolete. Male sepals broad, 4-5,
strongly imbricate; stamens 2-3, very shortly connate at base, the
stout connectives peltately and lobately connate. Ovary 2-3-
celled, the styles connate into a stout column, the ovules solitary.
Fruit large, fleshy without, indehiscent or finally separating into
2-valved cocci. Seeds large, nearly subglobose, ecarunculate but
apparently sometimes arillate. The name was suggested by the
centrally connate stamens.
Omphalea diandra L. Sp. PL ed. 2: 1377. 1763; 20.
High climbing, the younger terete branches appressed scabrius-
culous; petioles elongating to approximately length of blades, the
lustrous apical glands 2.5-3 mm. wide; leaves elliptic or orbicular-
ovate, openly cordate at base, shortly cuspidate-acuminate, 15-18
cm. long, 10-13 cm. wide, coriaceous, the younger softly fulvous
pubescent beneath, 3-5-nerved at base, the 2-3 secondary nerves
prominent beneath; stipules acuminate, 2.5 mm. long, sometimes
lobed; panicles very long, simple or with a few long branches;
cymules short, usually subbiparted; bracts 1-2 (7) cm. long, narrowly
lanceolate, basally biglandular; male flowers 2.5 mm. long, sepals
fulvous pubescent, orbicular-ovate, concave; stamens 2; ovary
pubescent; seeds of the large fruits 4.5 cm. long, 3 cm. thick, dorsally
obtusely convex, the ventral side obtusely carinate. It seems
strange that this conspicuous plant apparently has been found in
northern Peru only once. It is marked by the several dm. long
narrow or laxly branched remotely floriferous panicles with occasional
long bracts sometimes several cm. long. Illustrated, Pax, I.e. p. 21.
180 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
San Martin: Near Tarapoto, Spruce 2415. Rio Acre: Seringal
San Francisco, Ule 9382. Bolivia to Central America, West Indies
and Brazil.
40. NEALCHORNEA Huber
Reference: Pax & Hoffm. Pflanzenreich IV. 147. xiv. Add. 6:
51. 1919.
Nearly glabrous tree with petioled pinnate-nerved leaves and
dioecious (or monoecious, Ducke) flowers, the male in puberulent
panicles, the female in racemes, all with 4-merous lightly imbricate
or subentire subcupulate calyx. Disk none. Stamens 10, sessile,
the outer alternate with the sepals, connective longer than anthers
or completely adnate. Rudimentary ovary none. Staminodia in
female flowers few; ovary 2-celled; styles 2, connate only at base,
simple, stout; ovules solitary. Capsules depressed. Seeds ecaruncu-
late, cotyledons reniform, radicle short. Unknown to the monog-
raphers who suggest relationship with Gelonieae, separating it
from Cunuria Baillon by the lack of a rudimentary ovary. Ducke,
Archiv. Jard. Bot. Rio Jan. 4: 107. 1925, recorded his observation
made in the field that the species is monoecious or dioecious.
Nealchornea yapurensis Huber, Bol. Mus. Goeldi 7: 298.
1913; 51.
Medium-sized tree; petioles 2-10 cm. long; leaves ovate-elliptic,
acute at base, abruptly and shortly caudate-acuminate, remotely
crenate-dentate, 1-2 dm. long, 4-6 (8) cm. wide, subcoriaceous,
reticulate-venose; male panicles to 15 cm. long, many-flowered,
much-branched; bracts minute, triangular; pedicels 7 mm. long,
clavate; female flowers yellowish-green, fragrant, 3 mm. broad in
racemes 5-7 cm. long, the spreading pedicels 6 mm. long; sepals
with large hemispheric glands at base; ovary ovoid, reddish tomen-
tose; styles arcuately spreading, sulcate; male sepals orbicular,
fleshy at base, margins scariose; capsules 3 cm. broad, half as long,
seeds transverse-ovoid. There are sometimes one or two glands on
leaves beneath at base. The Rio Acre material has stouter, more
fulvous tomentulose pedicels than type. Illustrated, Ducke, Archiv.
Jard. Bot. Rio Jan. 4: pi. 10 (flower parts). F.M. Neg. 5407.
San Martin: Pongo de Cainarachi, King 2701. Loreto: Yuri-
maguas, Ule 6292 (det. Ducke). Rio Acre: Near mouth of Rio
Macauhan, Krukoff 5278; 5607 (both det. Steyermark). Brazil;
Colombia.
FLORA OF PERU 181
41. SENEFELDERA Mart.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 23-25. 1912.
Shrubs or trees with alternate stipulate petioled entire leaves,
crowded at tips of branchlets, and small monoecious apetalous flowers
in spikes panicled at the branchlet tips, the male sessile or nearly,
few to each bract, the female in lower part of spikes or panicle,
usually shortly pedicelled, solitary in each bract. Male calyx open
well before anthesis, 3-5-lobed; stamens 3-12, the receptacle elevated,
filaments short, anthers erect, extrorse; ovary rudiment none. Calyx
of female flowers deeply 3-parted, the lobes imbricate. Styles
connate into a short thick column, recurved-spreading above,
undivided. Capsule tridymous, the endocarp of the cocci ligneous,
the pyriform or irregularly subglobose seeds with small caruncle.
The male calyx ruptures irregularly, the pedicel turning toward the
rachis in S. inclinata M. Arg. of the upper Amazon. Named for
Alois Senefelder (1771-1834), lithographer of Vellozo's Fl. Flum.
and said to have invented the process.
Inflorescence with many spicate branches, simulating a compound
panicle S. Skutchiana.
Inflorescence open, the branches few.
Male pedicels erect, the calyx regular; petioles and leaves slightly
pubescent S. triandra.
Male pedicels somewhat or in part inclined; petioles and leaves
glabrous S. inclinata.
Senefeldera inclinata M. Arg. in Mart. Fl. Bras. 11, pt. 2:
530. 1874; 25.
Glabrous tree with smooth terete branchlets, their apically
crowded leaves on very unequal petioles, these typically 1.5-4 cm.
long, apically subglandular-tumidulous; leaves broadly obovate-
lanceolate, narrowed to base, acuminate or obtusish and cuspidately
acuminate, usually 1.5-2 dm. long, 5-7 cm. wide, coriaceous, lustrous,
prominently 12-14-nerved and reticulate, minutely maculate-bi-
glandular nearly at the base beneath; male calyces strongly inclined
at least in type, subirregular, 2-3-parted; stamens 7-10, receptacle
moderately elevated; capsules 15 mm. long, smooth, black, the
seeds 8-10 mm. long and broad. S. macrophylla Ducke, Archiv.
Jard. Bot. Rio Jan. 4: 113. 1925, in view of the Peruvian collections
may be designated S. inclinata var. macrophylla (Ducke) Macbr.,
comb, nov., differing typically as noted by the author in the larger
182 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
leaves, these usually 1.5-3 dm. long, 7-12 cm. wide and the much
longer petioles (5-14 cm. long); these characters vary greatly in
Peru material determined S. macrophylla by Mansfeld, who noted
"toward S. inclinata." F.M. Negs. 5412; 5414 (S. macrophylla}.
Loreto: Mishuyacu near Iquitos, Klug 109; 876. Amazonian
Brazil.
Senefeldera Skutchiana Croizat, Journ. Wash. Acad. Sci. 33:
18. 1943.
Entirely glabrous medium-sized tree; petioles 2.5-8 cm. long,
slightly enlarged apically, drying blackish; leaves entire, broadly
elliptic, broadly cuneate to subrotund at base, short-acuminate to
apiculate, slightly glandular at end of midrib where a little reflexed,
chartaceous to subcoriaceous, drying greenish, glands obsolete but
base of blade sometimes obscurely spotted above; nerves 10-14
pairs, prominent both sides; inflorescence apical with numerous
spiciform rigid axes, about 2 dm. long and wide, simulating a com-
pound panicle; male flowers many, the fewer female basal or scattered,
the former usually in 3's in axil of ovate bractlet 1.5-2 mm. long, the
margin erose-hy aline; in upper axils only middle flower maturing;
female flowers with 3 lanceolate entire imbricate bracts, glandular
at base within, 1.5-2.5 mm. long, with an occasional lateral male
flower; ovary subfusiform to 2 mm. long; styles fleshy, scarcely
divided. Distinctive in inflorescence (Croizat).
Huanuco: Tingo Maria, Skutch 4967, type; also 4961.
Senefeldera triandra Pax & Hoffm. Pflanzenreich IV. 147.
xiv. Add. 6: 55. 1919.
A 2-8-meter shrub with glabrous branchlets but the apically
and basally tumidulous petioles (1.5-4 cm. long) sparsely malpigh-
iaceous pubescent; stipules caducous, subulate, 2-3 mm. long;
leaves elliptic, acuminate, base acute, 13-15 cm. long, 5-6.5 cm.
wide, membranous, finely reticulate on both sides, sparsely pilose
and glandular beneath near margins; inflorescence glabrous or nearly,
4 cm. long, branches nearly 1.5 cm. long, lower female flowers 2-3,
sessile, the calyx 1.5 mm. long; male part about 1 cm. long, broad
bracts acuminate, biglandular, male pedicels nearly 1 mm. long,
erect; calyx 0.5 mm. long, glabrous, 3-lobed; stamens 3; styles 3,
barely connate at base, arcuate-recurved; ovary glabrous. S. multi-
flora Mart, (or apparently correctly S. verticillata (Veil.) Croizat,
Journ. Wash. Acad. Sci. 33: 18. 1943) of the upper Amazon, has
FLORA OF PERU 183
glabrous coriaceous leaves, longer inflorescences, 5-8 stamens. F.M.
Neg. 5413.
Rio Acre: Seringal San Francisco, Ule 9547, type.
42. MABEAAublet
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 26-42, 282.
1912.
Lactescent trees or shrubs sometimes scandent or the branches
so, the leaves alternate, about oblong, entire or denticulate, stipulate,
shortly petioled, with the flowers numerous in terminal sometimes
paniculate racemes, the bracts usually biglandular; female flowers
few at base of raceme, with solitary bract. Petals and disk none.
Male flowers with young calyces globose opening long before anthesis,
the 3-5 lobes lightly imbricate, 10-70 stamens, rarely only 2 or 3,
affixed to convex receptacle, anthers nearly sessile, extrorse, longi-
tudinally dehiscent, ovary none. Female flowers with 5 (rarely 6
or 3) imbricate often unequal sepals, 3-celled ovary, the styles free
above, undivided, ovules solitary. Capsules globose, breaking into
bivalved follicles with carunculate ovoid smooth seeds. Specific
definition within this group is not yet established; the presence or
absence of sepalar glands, for example, unless a concomitant
character, is surely open to question as taxonomically significant.
Leaves often multiglandular along margin beneath; male flowers
umbellate in narrow panicles.
Leaves firm, the marginal glands often obscure; anthers 12-15.
M. nitida.
Leaves membranous, the marginal glands prominent; anthers
15-25 M. Macbridei.
Leaves eglandular or glands few, scattered along margin; male
flowers if umbellate in panicles several cm. wide.
Male flowers umbellate in panicles several cm. wide.
Pedicels conspicuously globose-glandular at base . . M. Standleyi.
Pedicels if conspicuously glandular clearly so above the base;
species not proved.
Plants glabrous to the panicles, these about 3 cm. wide.
M. maynensis.
Plants more or less furfuraceous, at least the branchlets and
panicles, these mostly much wider than 4 cm. .M. elata.
184 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Male flowers racemose or spicate in racemiform panicles about
1 cm. wide, or, rarely, racemose.
Male flowers racemulose on short branchlets M. anadena.
Male flowers 1-3 in axils of bracts on the rachis. .M. subsessilis.
Mabea anadena Pax & Hoffm. Pflanzenreich IV. 147. xiv. Add.
vi: 55. 1919.
Branchlets compressed and, as the 8-12 mm. long petioles, reddish
scurfy, becoming terete and glabrescent; stipules caducous; leaves
lanceolate, acute at base, abruptly acuminate, 8-11 cm. long, 22-
33 mm. wide, serrate, membranous, lustrous above, opaque beneath
and scurfy on the 10-14 veins; panicles 6-10 cm. long including the
1-2 cm. long furfuraceous peduncles; male branchlets dense, to 8 mm.
long, racemes 4-6-flowered with ovate eglandular paucidenticulate
bracts, the flowers 2-2.5 mm. wide with 13-15(-19) stamens; sepals
of female flowers 6, ovate-acuminate, about 3 mm. long; ovary
reddish pubescent, style column 4. 5 (-9) mm. long, free part to 7 mm.
long. In section Spiculigerae but bracts eglandular (authors). In
Krukoff specimen stamens to 19, style column twice as long, sepals
3.5-4.5 mm. long (Steyermark). Tree, 10-30 meters high. F.M.
Neg. 5415.
Rio Acre: Seringal San Francisco, Ule 9548, type. Mouth of
Rio Macauhan, Krukoff 5430. Bolivia.
Mabea elata Steyerm. Field Mus. Bot. 17: 418. 1938. M.
rhynchophylla Diels, Notizbl. Bot. Gart. Berlin 14: 335. 1939.
Branchlets brownish furfuraceous; stipules wanting; leaves ob-
long-elliptic, obtuse at base, abruptly caudate-acuminate, 8.5-20
cm. long, 2.5-6.5 cm. wide, subentire, subcoriaceous, glabrous or
the midrib above and near tip scurfy puberulent, superficially
puberulent beneath, lateral nerves 15-17 pairs; panicles 11-17 cm.
long, 4-5.5 cm. wide, rachis pubescent; sepals all puberulent; male
pedicels in racemose umbels, with bracts 4-5 mm. long, 1.5-2 cm.
long, biglandular 2-3 mm. above the base, the glands black, 3-4 mm.
long, half as thick, ashy puberulent; sepals round, stamens 50-60;
female sepals 4.5-7 mm. long, interior longer than exterior, acuminate,
all glandular both sides, black without; style column 8 mm. long,
tomentulose; styles 12 mm. long. The glandular inner surface of
the female sepals places this in the section Intermediae. But the
character is not clear and its constancy may be questioned; certainly
the species otherwise resembles M. eximia Ducke, Archiv. Jard.
FLORA OF PERU 185
Bot. Rio Jan. 4: 107. 1925, to which the careful Mansfeld referred
King 43, "vel valde aff." and which in turn as noted by Ducke is
near M. pulcherrima M. Arg., 32, of French Guiana; M. eximia
however as to type has 100-120 stamens, M. pulcherrima about
half as many, that is as to type. Klug 43 seems to be intermediate
and until the validity of these several characters is further proved
may as well be included here. Another closely related species, M.
speciosa M. Arg., 31, recently is said to have been found as near as
Sao Paulo de Olivenca and would be sought here; it has leaves
fuscous-glaucous beneath, panicles 3.5-4 cm. wide, sepals all eglan-
dular, stamens about 65. These forms seem to be very narrow
indeed and perhaps will prove to comprise at most two species.
M. Klugii Steyerm. I.e., 416, of the upper Putumayo, Colombia,
has persisting sepals, 25-30 stamens. Type a 10-meter tree and is
the same fide Croizat, Caldasia 2: 360. 1944, who has pointed out
that short branchlets present differences of leaves and inflorescences
not apparent on adult branches.
Loreto: Between Balsapuerto and Moyobamba, Klug 3206, type
(also for M. rhynchophylla). Mishuyacu, Klug 48? Ecuador;
Colombia?
Mabea Macbridei Johnst. Contr. Gray Herb. 75: 27. 1925.
A small tree with early pulverulent slender subangled branchlets;
petioles 9-12 mm. long; leaves oblong or elliptic-oblong, 12-18 cm.
long, 4.5-6 cm. wide, glabrous, lightly crenate, lustrous above,
glaucous beneath, the veins conspicuous and with about 30 minute
lineately marginal glands toward the abruptly acuminate and obtuse
tip, rounded or obtuse at the eglandular base; panicles terminal
(sometimes branched), ferrugineous-tomentulose, about 1 dm. long,
10-12 mm. wide; bracts lanceolate; pistillate branchlets 1 mm. long,
1-flowered, staminate 1-2 mm. long, 2-4-flowered, obscurely glandu-
lar at base, the pedicels of female flowers 7 mm. long, canescent-
strigose, those of the male 5-10 mm. long, sepals of former orbicular
or acutely ovate, puberulent, 1-1.5 mm. long, eglandular, the ovary
merely tomentose with style column 5-7 mm. long, styles as long;
male flowers with calyx 2 mm. wide, deltoid lobes, 15-25 stamens,
sparsely pubescent anthers. The peculiar submarginal row of glands
distinguishes this from all South American species except M. nitida
Benth., 39, of the upper Amazon with different inflorescence;
M. maynensis also differs in the panicles (Johnston). I have not
seen authentic material of M. nitida but ex char, and photograph
186 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
it appears to have firmer more lustrous leaves and the 12-15 anthers
are described as pulverulent.
Junin: Hacienda Schunke above La Merced, 5702, type; Schunke
A106.
Mabea maynensis M. Arg. in DC. Prodr. 15, pt. 2: 1150.
1866; 32.
Glabrous to the panicles; branchlets early compressed, slender;
petioles 12-15 mm. long; leaves lanceolate-elliptic, subacute at
base, abruptly and shortly acuminate, 9-14 cm. long, 3.5-7 cm. wide,
repandly crenate, reticulate-veined, secondary veins 10-15; panicles
about a dm. long, 3 or 4 cm. wide, male branchlets 4-5 mm. long,
umbellately 2-3-flowered, coarsely biglandular well above the base,
pedicels 1-1.5 cm. long, calyx pulverulent, nearly 3 mm. wide, with
semiorbicular lobes; stamens 33-40, connective apically crenulate,
pubescent; female pedicels 2-3 cm. long, sepals acuminate, 3 mm.
long; bracts reddish-puberulent, eglandular, as the ovary, this smooth,
tomentulose; style column nearly 2 cm. long, the free parts about
two and a half times longer. M. caudata Pax & Hoffm. I.e., 282,
of British Guiana, to which Mansfeld referred Tessmann 4275, has
rather ovate leaves 11-15 cm. long, 5.5-9 cm. wide, about 30 stamens,
style column 22-25 mm. long; it seems to be doubtfully distinct, at
least specifically. F.M. Neg. 5424.
San Martin: Tarapoto, Spmce 4888, type. Pongo de Cainarachi,
Klug 2721 (det. Croizat). Loreto: Lower Rio Huallaga, Williams
3903 (det. Croizat). Mouth of the Rio Santiago, (Tessmann 4275).
Amazonian Brazil; British Guiana?
Mabea nitida Spruce ex Benth. in Hook. Journ. Bot. Misc. 6:
367. 1854; 39.
Glabrous to the pulverulent or glabrate panicles, these only 1 cm.
wide or narrower, the umbellately 3-flowered male branchlets ver
short, typically scarcely longer than the calyces, eglandular bracted
a little above the base; branchlets slender, lustrous, dark; petioles
7-10 mm. long; leaves oblong-elliptic, obtuse or subacute at base,
subabruptly and shortly caudate-acuminate, (8) 12-17 cm. long,
3-5 cm. wide, lustrous above, glaucous-opaque beneath, the marginal
glands several or nearly lacking; male pedicels unequal, the longer
middle one scarcely more than 2 mm. long, the calyx not that broad
and with 12-15 pulverulent anthers; sepals of female flowers eglandu-
lar; ovary pulverulent, the carpels geminately bimuricate; capsules
FLORA OF PERU 187
globose, intensely rufous-pulverulent, smooth or minutely muricate,
16 mm. large, the obscurely olive-black seeds 1 cm. long, 8.5 mm.
broad. Trees 10-20 meters tall with abundant latex (Klug). F.M.
Neg. 5425.
Loreto: Mishuyacu near Iquitos, Klug 1442. Amazonian Brazil.
Mabea Standleyi Steyerm. Field Mus. Bot. 17: 417. 1938.
Glabrous except the panicles, these puberulent; branchlets olive-
brown; petioles 8-10 mm. long; leaves oblong or elliptic-oblong,
obtuse at base, abruptly acuminate, 12-20 cm. long, 5.5-6.5 cm.
wide, subentire, the 11-12 pairs of lateral nerves conspicuous be-
neath; stipules of flowering branches persisting, linear-setaceous,
about 1 cm. long; panicles about 1 dm. long, 2.5 cm. broad, male
pedicels 10-13 mm. long with 2 globose glands at very base 2-2.3
mm. in diameter, sepals rounded, purplish-puberulent; stamens 60-
65; female sepals 2.5-3.5 mm. long, acuminate, eglandular within;
style column puberulent, about 2 cm. long, the recurved styles 6 mm.
long. Forest shrub 2 meters high of section Umbelluliferae, near
M. speciosa M. Arg., 31, but that as all other species of the section
without the orbicular glands next to the panicle rachis (the author).
Loreto: Florida, mouth of the Rio Zubineta, Klug 2064, type.
Mabea subsessilis Pax & Hoffm. Pflanzenreich IV. 147. v: 282.
1912.
Small tree, the rusty puberulent slender branchlets tardily
glabrescent; petioles puberulent, 1-2 mm. long; leaves oblong-
lanceolate or in type broadly lanceolate, cordate or subcordate at
base, abruptly caudate-acuminate, in type 6-10 cm. long, 2-3.5 cm.
wide, mucronulate, coarsely uncinate-serrate, lustrous above, sub-
glaucescent beneath, firm-membranous, reticulate-veined both sides;
stipules subpersisting, glabrous, 5 mm. long, subulate-attenuate;
panicled inflorescence composed of about 4 racemes 3-4 cm. long,
only the 2-3 basal flowers female; rachis and bracts pubescent, the
broadly ovate acuminate male minutely glandular at base, the
narrower female eglandular, the former with 3 flowers on pedicels
1-1.5 mm. long, the solitary pedicels of the latter 3-4 mm. long;
male flowers 1 mm. wide or smaller; stamens about 10; female sepals
6, acuminate, unequal, 1.5-2 mm. long, at least one denticulate;
ovary smooth, reddish-furfuraceous; style column to 7 mm. long,
pubescent; capsules puberulent, 15 mm. long. Section Apodae Pax
& Hoffm. characterized by the male pedicels ternate in bracts on
188 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
the rachis and according to the authors nearest M. crenulata Sp.
Moore, 39, also from southern Brazil but "very diverse"; it and
M . indorum Sp. Moore, 40, similar but male pedicels to 4 mm. long
have crenulate leaves acute or little rounded at base, obtuse to
shortly cuspidate. Determinations by Mansfeld as M. subsessilis
but as the Peruvian form is remote geographically and differs slightly
it may be worth while to note it as M. subsessilis var. peruviana
Macbr., var. nov., foliis oblongis 1-2.5 cm. latis minute serrulatis
subtus obscure reticulate- venosis; pedicellis circa 2 mm. longis;
capsulis 18-20 mm. longis. Type, King 555. Williams, Field Mus.
Bot. 15: 279. 1936, gives the anatomy, noting the wood as light and
soft, suitable for carpentry; sometimes 15 meters tall. F.M. Neg.
5428.
Loreto: Mishuyacu near Iquitos, Klug 269; 555; 701; 847.
Lower Rio Nanay, Williams 653. Tierra Doble, Williams 896.
Brazil. "Amaquillo," "manchinga blanca" (both, Williams).
43. SEBASTIANIA Spreng.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 88-153. 1912,
and Add. 6: 58. 1919.
Usually more or less shrubby, sometimes arborescent, rarely
completely herbaceous and annual, the shortly petioled leaves
ordinarily alternate, small, rigid and serrulate; stipules small.
Flowers small, apetalous, usually monoecious and in slender often
entirely male spikes commonly borne in leafy terminal branchlets
or opposite the leaves; bracts basally biglandular; female flowers
most frequently few or solitary at the base of the male. Disk none.
Male calyx often 3-lobed or -parted, the somewhat larger female
with 3 sepals. Stamens 3, 2 or exceptionally 4, free or connate
basally, the anthers longitudinally dehiscent. Ovary 3 (2) -celled,
ovules solitary, styles spreading or revolute, undivided, rarely con-
nate. Capsules tridymous or subglobose, smooth or armed, separat-
ing from central column into bivalved cocci with oblong subglobose
or cylindric carunculate seeds. Commemorates Antonio Sebastiani,
professor of botany and physician of Rome (1782-1821).
The "brincadores" or Jumping Beans are often the seeds of a
Mexican species, similar to the Peruvian S. obtusifolia, that have
become inhabited by a larva which by its movements causes the
seeds to stir or even jump; Pax and Hoffmann, Pflanzenreich I.e.
147-148, give a detailed account (with bibliography) of this popular
subject.
FLORA OF PERU 189
Annual, sometimes lignescent at base; spikes opposite leaves.
S. corniculata.
Shrubs or trees; spikes axillary or terminal.
Petioles 3-5 mm. long; obtuse or barely acute.
Branches terminating in long sharp spines S. haploclada.
Branches not modified into spines S. obtusifolia.
Petioles about 10 mm. long; leaves shortly acuminate.
S. huallagensis.
Sebastiania corniculata (Vahl) Pax, Pflanzenreich IV. 147. v:
96. 1912. Tragia corniculata Vahl, Eclog. Amer. 2: 55. pi. 19. 1798.
Annual herb or sometimes becoming more or less lignescent at
base, the branching stems often 1-several dm. high, pilose-hispid or
glabrate; petioles slender, often 1 cm. long or longer; leaves ovate-
lanceolate, rounded-cordate at base, acuminate, usually 3-4 cm.
long, 1-2 cm. wide, subentire or with small incurved whitish teeth
and with a few marginal glands near the base beneath where often
more or less pilose; male spikes opposite the leaves 1-2 cm. long, the
slender rachis pilose or glabrate; bracts minute, 3-5-flowered, 3-lobed;
larger female flowers usually solitary at base of male spikes or below
and opposite leaves, their sepals denticulate and basally stipitate
glandular; ovary subglobose, rather irregularly 6-muricate, the
capsule about the truncate base and apex with 6 wing-like ap-
pendages, finally 5 mm. long; seeds 2 mm. long, subcylindric, dark
gray. Illustrated, Pax, I.e. p. 97. F.M. Neg. 21566.
Peru (probably). Mexico and the West Indies to Colombia and
eastward.
Sebastiania haploclada Briq. Ann. Cons. Jard. Bot. Geneve 4:
231. 1900; 131.
Shrub or tree with stout branches terminating in a sharp spine
as much as 5 cm. long; petioles 3-5 mm. long; leaves elliptic-obovate,
cuneate to short petioles, obtusely rounded or subobtuse, about 1 cm.
wide, 3 cm. long, entire, hard-coriaceous, green and somewhat
lustrous, glabrous but glandular, with 5-6 little or scarcely prominent
lateral nerves; male spikes 2-3 cm. long terminating short leafy
branchlets, the rachis stout; bracts ovate, entire or nearly, 3-flowered;
male and female sessile or subsessile, the parts minute, ovate;
anthers large, orange, long-separated; styles elongate. Not seen
by Pax, who does not distinguish it from S. pachystachys M. Arg.,
190 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
129, of southern Brazil; Briquet allied it to the more widely dis-
tributed S. Klotzschiana M. Arg.; except for the spinescent branches
its resemblance to S. obtusifolia is striking. F.M. Neg. 24565.
Amazonas: Chachapoyas, Mathews, type.
Sebastiania huallagensis Croizat, Journ. Arnold Arb. 24: 177.
1943.
Entirely glabrous tree, even the young shoots; petioles 8-10 mm.
long; leaves elliptic-lanceolate, base shortly cuneate and at juncture
with petiole with 2 dark-colored glands beneath, rather abruptly
acuminate, entire, 6-9 cm. long, 2.5-3.5 cm. wide, with about 10-14
pairs of slender nerves; inflorescences slender, bisexual, to 8 cm. long,
the male flowers 6-8, about 1.5 mm. wide, perianth minute, sub-
sessile, 3-lobulate, the 3 stamens alternate, lobes basally connate;
female flowers solitary on pedicels about 5 mm. long; ovary smooth,
4 mm. long, 1.5 mm. broad, basally enclosed with 3-5 imbricate
bracts, the 2-3 scarcely slender styles divergent. Distributed as
Alchornea sp. which it certainly is not; the sum of the characters
suggests Sebastiania but better material is needed to confirm this
disposition (Croizat). Compare however S. brasiliensis Spreng., 138,
which it resembles greatly at least in general character, except that
in that species typically the male bracts are 3 (16) -flowered and the
leaves are eglandular. Bolivian material, however, so determined
seems to be the same as S. huallagensis but of course may be incor-
rectly named.
San Martin: Juanjui, Alto Rio Huallaga, Klug 4%43, type.
Sebastiania obtusifolia (HBK.) Pax & Hoffm. Pflanzenreich
IV. 147. 5: 149. 1912. Sapium obtusifolium HBK. Nov. Gen. &
Sp. 2: 63. 1817.
An entirely glabrous 2-meter shrub with rigid terete branches
but the densely leafy branchlets angled in drying; petioles 3-5 mm.
long; leaves usually lanceolate-obovate or -spathulate, acute or
cuneate at base, rounded-obtuse or subacute, 2-4.5 cm. long, 10-18
mm. wide, entire or scarcely crenulate, eglandular, subcoriaceous,
the very fine secondary nerves hardly conspicuous; stipules broad
at base, denticulate, long subulate-acuminate, minute; spikes
terminal, 3-4 cm. long, dense, rigid, with one sessile female flower
at base or none; bracts ovate, acute or acuminate, lightly denticulate,
with smooth orbicular glands at each side, the 3 male flowers sub-
sessile; male sepals 3, narrow, subulate, the acuminate female
FLORA OF PERU 191
'glandular within and longer than the smooth glabrous ovary;
stamens 3; styles columnar ly connate; capsules 15 mm. long, 2 cm.
vide, the globose-ovoid seeds 6 mm. thick. The latex is used
nedicinally (Vargas).
Cajamarca: In woods at Querecotillo, Humboldt, type. Huari,
3havin de Huanter in the valley of the Rio Pucha, Weberbauer
1739. Libertad: Angasmarca, 3,300 meters, Weberbauer 7088
funin: Huasahuasi, Ruiz & Pavdn. Tarma, Weberbauer 1752.
3hanchamayo, Isern 2332. Apurimac: Andahuaylas, Vargas 8803.
3urahuasi, Vargas 9613 (det. Standley). "Vinagrillo" (Vargas).
44. STILLINGIA Garden
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 180-199.
L912.
Like Sapium but, especially, the base of the pericarp persisting
between the bivalvate cocci, these usually 3, leaving a 3-radiate
receptacle, the columella, if developed, more or less winged. Leaves
glandular-serrulate, often biglandular at tip of short petioles, some-
times opposite. As remarked by the monographers the group is
usually not distinguishable from Sapium unless the species are in
fruit; Otto Kuntze treated it as a section and his interpretation
sould well be followed, at least in floristic works. The name com-
memorates Benjamin Stillingfleet, an English botanist (1702-1771).
Stillingia salpingadenia (M. Arg.) Huber, Bull. Herb. Boiss.
3e"r. 2. 6: 452. 1906; 188. Sapium salpingadenium M. Arg. Linnaea
32: 121. 1863. Excoecaria salpingadenium M. Arg. in DC. Prodr.
15, pt. 2: 1209. 1866.
Branches short, densely leafy; petioles 2-5 mm. long, leaves
4-9 cm. long, 1-2 cm. wide, elliptic, lanceolate or linear-lanceolate,
acute or subobtuse both ends, densely and acutely callous serrulate,
minutely puncticulate, rigid-coriaceous, slightly lustrous above, paler
beneath, cartilaginous-margined, nerves prominent both sides;
stipules subulate, caducous; spikes terminal, 4-5 cm. long, all male
or female at base, broadly acuminate, 3-9-flowered, with large
cupuliform glands, flowers sessile, calyx of male 2-lobed, the female
with three ovate cuspidate subdenticulate sepals; stamens 2; styles
very shortly connate; capsules smooth, 10 mm. long and broad,
cocci 8-9 mm. wide; seeds smooth, subglobose, 6 mm. long, caruncu-
late. The determination of the Peruvian material is doubtful but
192 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
placed here by Standley; for identification it may be designated
var. peruviana Macbr., var. nov., petiolis 2.5 mm. longis; foliis
2-3.5 cm. longis, 7-10 mm. latis, margine obscure calloso-serrulatis;
bracteis non valde glanduligeris; styli recurvo-patuli. The spikes
too are shorter than described for this species and according to
collectors deep blood-red in color; if Standley is correct in placing
the shrub in this genus it may well prove to be new. Fruit said to
be edible. Illustrated, Pflanzenreich I.e. p. 189.
Huancavelica: South of Mejorada in river shrubland, 2,500
meters, Stork & Horton (type, var.). Bolivia; Paraguay. "Cabra-
cabra" (Stork & Horton).
45. SAPIUM P. Brown
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v. 199-258.
1912, and Add. 6: 60-61. 1919.
Glabrous trees or shrub with actually alternate entire or denticu-
late or often marginally glandular leaves and commonly terminal
spikes of small usually monoecious apetalous flowers, the female
ordinarily few at the base. Disk none. Calyces 2-3-lobed or
dentate or the female sometimes 2-3-parted or tubular. Stamens
2-3, free or partly connate, exserted; anthers 2-celled, longitudinally
dehiscent. Ovary 2-3-celled; ovules solitary; styles free or shortly
connate, recurved-spreading, deciduous. Capsules globose, pyriform
or composed of 3 cocci, ligneous or subdrupaceous, variously
dehiscent or often indehiscent, sometimes 1-seeded. Acknowledg-
ment with appreciation is made to Croizat, Journ. Arnold Arb. 24:
173-177. 1943, and Caldasia 2: 131-134. 1943, for his careful observa-
tions in this group the species of which as proposed will not be
understood until much more study of complete materials.
As indicated below, various species are a source of rubber.
Petioles all or mostly 2-3 cm. long (1-5 cm.).
Leaves entire or essentially, about half as wide as long.
Leaves plane or nearly even to tip, large (often 6 cm. wide);
petioles elongate (2-5 cm.) S. marmieri.
Leaves mostly more or less cucullate or inflexed at tip; petioles
often less than 3 cm. long.
Leaves as to type described as metallic lustrous above.
S. aerium.
Leaves dull, apparently never metallic lustrous above.
S. taburu, S. glandulosum.
FLORA OF PERU 193
Leaves entire, about one-third as wide as long, inflexed at tip.
S. Poeppigii, S. aereum.
Leaves somewhat serrulate, sometimes obsoletely, about one-third
as wide as long, plane.
Petiolar glands linear, immersed S. utile.
Petiolar glands lacking or obscure S. eglandulosum.
Petioles all or mostly 1 cm. long (7-12 mm.) S. Martii.
Sapiutn aereum Klotzsch ex M. Arg. Linnaea 32: 119. 1863;
233.
Similar to S. marmieri but the smaller lustrous leaves a sub-
metallic greenish color above in type, paler beneath, oval-oblong,
subcuneate at base, rounded at apex where thicker and cucullately
introflexed, somewhat glandular and revolute marginally, about
10-12 cm. long, 4-4.5 cm. wide, the slender secondary nerves arcuate;
petioles 1.5-3 cm. long, the apical glands cylindric, conspicuous or
in type conic-ovoid, small; flowers unknown in type but in following
material like S. Poeppigii; ovary enlarged at base, crowned with the
3-lobed deciduous style; capsules about 1 cm. long, 3-celled; seeds
subglobose, compressed, tuberculate, 5-6 mm. broad. Croizat has
suggested that the fruit may be larger and slightly different in shape
than that of S. glandulosum, but information as yet is lacking on
this character; he also says that it differs from S. marmieri primarily
in its leaves on the whole being smaller and narrower, in which case
my first key character (after Pax) is not valid, but ex negative the
second is and thus the species name is probably the earlier for
S. Poeppigii. S. taburu could also be included here. The petiolar
glands are evidently variable in development. Croizat has referred
from neighboring regions of Brazil Krukoff 6158; 6296; 6307 and
8311. Illustrated, Hook. Icon. pi. 2682. F.M. Neg. 5519.
The white latex is used as an adulterant of Hevea rubber
(Seibert).
Huanuco: Pampayacu, Kanehira 28 (det. Johnston, S. glandu-
losum). Junin: Vitoc, Ruiz & Pavdn, type. La Merced, Weber-
bauer 1937. San Martin: Zepelacio, Klug 3374. San Roque,
Williams 7351 . Loreto : Santa Rosa, Williams 4771 (det. Steyer-
mark, S. Poeppigii). La Victoria, Williams 2865 (det. Standley,
S. glandulosum). Rio Ucayali, Seibert 1886 (det. A. C. Smith,
S. Pavonianum). Amazonian Brazil. "Shiringa rana" (Seibert),
"guta percha" (Williams).
194 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Sapium eglandulosum Ule, Tropenpfl. 9, Beih. 6: 14. 1905,
and Bot. Jahrb. 35: 673. pi 2. 1905; 206.
Tree 5-15 meters high with rigid flowering branchlets; petioles
2-3 cm. long sometimes glandular apically; leaves oblong-lanceolate,
acute both ends, the terminal gland not inflexed, entire, lustrous,
10-16 cm. long, 4 cm. wide, the subhorizontal numerous secondary
nerves inconspicuously reticulate; spikes unisexual, the stout
terminal male 1-2 dm. long, with much shorter (but only young
known) female (about 3) borne at their base; bracts (male) broadly
ovate, obtuse, biglandular, 10-15-flowered; calyx bilobed; stamens 2;
ovary 3-celled; seeds corrugate. Not clearly distinct from S.
marmieri unless in leaf shape, which resembles more that of S. utile.
Illustrated, Ule, I.e. and Pax, I.e. p. 207.
The sap is said to provide a rubber.
Huanuco: Huamalies, between Monzon and the Huallags
Weberbauer 3578 (probably but sterile, Pax). Without data, Weber-
bauer, fide Pax. Brazil.
Sapium glandulosum (L.) Morong, Ann. N. Y. Acad. Sci. 7:
227. 1893; 231. S. Hippomane G. F. W. Mey. Prim. PL Esseq.
275. 1818 as to authors, the name fide Croizat, Journ. Arnold Arb.
24 : 176. 1943, being a new name for Hippomane biglandulosa L.
Shrub or tree attaining 20 meters; petioles canaliculate, 7-30 mm.
long, biglandular a little below the apex with straight, recurved or
incurved glands 2-3 mm. long; leaves elliptic or oblong-lanceolate,
rounded or cuneate at base, abruptly acuminate, the tip itself more
or less cucullately inflexed, subentire, subcoriaceous, commonly
4-15 cm. long, 3.5-8 cm. wide, or those on sterile branches much
larger, acute, serrulate and somewhat glandular; secondary nerves
very numerous, slightly arcuate and as the veins prominent; spikes
terminal, 7-25 cm. long, slender, androgynous or entirely male;
bracts greatly reduced with an oval gland both sides, the male
6-9-flowered, the calyx of these purplish, bilobed, that of the female
3-parted; stamens 2; styles 3, connate at base; capsules sessile, glo-
bose, 7-9 mm. long; seeds sublenticular, 5-6 mm. long and broad
with red membranous cover and deciduously tuberculate. Illus-
trated, Hook. Icon. pi. 2650 (as S. aucuparium).
The lactescent sap according to Ule provides a high class rubber.
Williams found the tree attaining 40 meters with spreading crown and
straight cylindrical trunk 5 dm. or more in diameter, the yellowish-
white or brown bark with coarse ridges, the latex sweet, pinkish,
FLORA OF PERU 195
the wood not durable (Field Mus. Bot. 15: 282. 1936). Probably
not in strict sense in Peru as suggested by Croizat but the several
names may represent variants of less than specific importance.
Junin: La Merced, Chanchamayo Valley, Weberbauer 1929 (det.
Pax). Huanuco: Upper Amazon, Kanehira 12 (det. Johnston).
Loreto: Fortaleza, Williams 4403. Yarina Cocha, Tessmann 3404
(det. Mansfeld). Puerto Mele"ndez, Tessmann 4790 (det. Mansfeld).
To the West Indies. "Gutapercha" (Williams), "milk tree,"
"steringarana' ' (Tessmann) .
Sapium marmieri Huber, Bot. Mus. Paraense 3: 397. 1902; 256.
A 20-meter tree with smooth gray bark, stout branches, nigrescent
branchlets; stipules broadly ovate, acute; petioles 3-5, usually 4 cm.
long, slender, terete but lightly canaliculate above, faintly bi-
glandular below the middle; leaves oblong-elliptic (apparently some-
times shortly so), rounded base and apex or the latter obtusish or
slightly emarginate, 10-12 or sometimes 8 or 15 cm. long, 5-7 often
6 cm. wide, entire or little undulate, lustrous above, opaque beneath;
secondary nerves 15-20, arcuate; male bracts 3-flowered, triangular
or orbicular, the large glands obovate or subreniform; male calyx
bifid; capsules 8 mm. in diameter; seeds rugose. Illustrated, Hook.
Icon. pi. 2899. FM. Neg. 24577.
The abundant milky sap is very poisonous and produces a less
valuable rubber for which it is used in Ecuador (Huber). Determina-
tions by Croizat. Most of the specimens have dried a reddish-brown
color.
Loreto: Alluvial soils of the Ucayali and Huallaga, (Huber,
type), Tessmann 3234. Alto Rio Itaya, Williams 3490; Killip &
Smith 29285. Lower Rio Huallaga, Williams 4904; 4898; 4789;
4626; Killip & Smith 29265. Rio Acre: Mouth of Rio Macauhan,
Krukoff 5717 (det. Steyermark). Brazil; Colombia. "Seringa
rana," "seringueria," "shiringa masha," "caucho masha," "pampa
caucho," "gutapercha."
Sapium Martii (M. Arg.) Huber, Bull. Herb. Boiss. se>. 2, 6:
448. fig. 42. 1906; 210. Excoecaria Martii M. Arg. in Mart. Fl.
Bras. 11, pt. 2: 614. 1874.
Much-branched; stipules fulvous, triangular hastate at base,
lacerate-dentate, scarious except the nerves; petioles to 12 mm. long,
thick, glabrous, biglandular at tip; leaves oblong-elliptic, shortly
and acutely acuminate both ends, concolorous, chartaceous, 5-8 cm.
196 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
long, 2-3.5 cm. wide; secondary nerves 12-15, prominent as the
reticulate veins; spikes terminal, bracts semi-orbicular, male 10-
flowered, with a large orbicular gland both sides; female calyx sessile,
tubular, obtusely 3-lobed, the short lobe semi-orbicular, style shortly
3-parted; capsules about 7 mm. long, seeds tuberculate, subglobose,
4 mm. thick. Perhaps a variant of the Brazilian S. obovatum (M.
Arg.) Klotzsch with coriaceous stipules and leaves. The Peruvian
shrub which furnishes description of fruit and is placed here ex
char, seems to have thinner leaves slightly cucullate at tip and for
convenience may be noted as var. peruviana Macbr., var. nov.,
foliis chartaceis apice obscure incurvis. F.M. Neg. 19538.
San Martin: Tarapoto, Williams 6658. Juan Guerra, Williams
6909. Loreto: Iquitos, Williams 8032. Brazil; Ecuador?
Sapium Poeppigii Hemsley in Hook. Icon. pi. 2678. 1901; 229.
Sapium biglandulosum (Aubl.) M. Arg. var. hamatum [Poeppig] M.
Arg. Linnaea 32: 116. 1863. S. hamatum ([Poeppig] M. Arg.) Pax
& Hoffm. Pflanzenreich IV. 147. v: 229. 1912.
Flowering branchlets stout; stipules squamiform, persisting;
petioles to 2.5 cm. long with 2 subglobose glands at the apex; leaves
oblong or elliptic-oblong, rounded both ends or shortly contracted
just at base, the introrsely cucullate tip with a large gland, 9-12 cm.
long, 4-5 cm. wide, lustrous above, paler and opaque beneath, merely
subcoriaceous, obscurely crenulate and glandular, the many second-
ary slender nerves arcuate-ascending; spikes terminal, androgynous
or all male, 8-10 cm. long, usually with 12-15 basal female flowers;
bracts broadly triangular with oblong glands, the male 5-7-flowered,
these with bilobed calyx, that of the female 3-parted; stamens 2;
ovary 3-celled, sessile; styles 3, nearly free, not persisting. Croizat
notes that the leaves are narrower, longer and thinner than are those
of S. marmieri. It seems to be the same as S. aereum, or very similar.
The white latex is used as an adulterant of Hevea rubber (Seibert).
Illustrated, Pax, I.e. p. 227.
Huanuco: Cuchero, Poeppig 1782, type.
Sapium taburu Ule, Tropenpfl. 9. Beih. 6: 13, 16. fig. 3 D,E.
1905, and Bot. Jahrb. 35: 671. fig. 2E, G. 1905; 232.
Apparently scarcely distinguishable from S. glandulosum, the
only definite contrast in description relating to the seeds, which are
described as subtrigonous, compressed, 6-7 mm. in diameter, corru-
gated. Croizat thinks it will be shown that this is properly a part
FLORA OF PERU 197
of S. aereum and suggests that Klug 1668 from Putumayo, Colombia,
may belong to it. Illustrated, Pax, I.e. p. 207; Hook. Icon. pi. 2898.
The milky sap is often mixed with that of Hevea (Ule).
San Martin: Moyobamba, Weberbauer 4759 (det. Pax); 288.
Brazil. "Tapuru."
Sapium utile Preuss, Exped. Centr. & Siidamer. 386. pi. 11.
1901; 204. S. biglandulosum (L.) M. Arg. var. Pavonianum M. Arg.
Linnaea 32: 116. 1863? S. Pavonianum (M. Arg.) Huber, Bull.
Herb. Boiss. ser. 2, 6: 356. fig. 14. 1906?
Becoming a tree 12-15 meters high, the rather stout branches
erect, the branchlets densely leafy; petioles 1.5-3.5 cm. long with
2 linear immersed sometimes obsolete glands at apex; stipules ovate;
leaves oblong, lanceolate or oblanceolate, cuneate or rarely rounded
at base, shortly and obtusely acuminate, subcoriaceous, plane or
nearly so, minutely serrulate or subentire, often 1.5-2 dm. long,
5.5-7 cm. wide, the secondary nerves very many and prominent
especially beneath; spikes androgynous or all flowers male, terminal,
1-2.5 dm. long; bracts broadly ovate, subentire, with an oblong-
peltate gland base each side, the male flowers 7-9, these with bifid
calyx; stamens 2; ovary 3-celled; capsules globose, mature un-
described. The type was from Juan de Oro, Ecuador. The Peruvian
specimen may not be the same; the plate of S. Pavonianum shows
an entire leaf. Illustrated, Hook. Icon. pi. 2896.
The sap is said to provide a rubber of medium quality and accord-
ing to Preuss the fast-growing tree is easily adapted to use as a
shade tree for cacao and other plantations.
Peru: Without locality, (Pavon). Ecuador; Colombia. "Palo
de Leche."
46. ACTINOSTEMON Klotzsch
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 57-80.
1912.
Shrubs or trees, glabrous or nearly, with alternate shortly petioled
entire pinnate-nerved leaves often more or less glanduliferous be-
neath, and ordinarily if not always monoecious apetalous flowers in
solitary or few terminal or axillary racemes from (in bud) coriaceous
imbricated scales, the male flowers 2-7 in the axils of often biglan-
dular bracts, the solitary female often well-pedicelled. Disk lacking.
Male perianth (as female) commonly with 1-3 sepals, 2-17 free
198 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
stamens, anthers ovoid, rudimentary ovary wanting; female flowers
(as male) sometimes esepalous; ovary 3-celled, styles connate at
base, recurved, undivided; capsules with 3 armed or smooth cocci,
the subglobose seeds carunculate.
Actinostemon imbricatus M. Arg. Linnaea 34: 216. 1865; 80.
Ultimate branches elongate, compressed above, glabrous as the
entire plant; leaves obovate, rounded-obtuse, shortly cuneately
narrowed into the short (4-6 mm.) petiole, rigid, prominently reticu-
late, eglandular beneath, 5-12 cm. long, 3-8 cm. wide; secondary
nerves nearly all starting between the base and the lower third of
the blade, all ascending, subflexuose; male spikes aggregate, short,
strobiliform at first by the many imbricate brown ciliolate scales;
flowering bracts minute, 3-flowered, early caducous; stamens 3;
receptacle between the bases of the filaments tumid, accrescent;
female flowers unknown. Mueller remarked : recedes from this genus
in the nature of the male receptacle and the deciduous bracts. Pax:
probably to be excluded. However, neither authority suggested
another relationship. P.M. Neg. 32495.
Huanuco: Cuchero, Poeppig 1308, type. Brazil?
. ' .
47. MAPROUNEA Aublet
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 175-180.
1912.
Entirely glabrous trees or shrubs with petioled entire leaves
often maculate-glandular near the base and subglobose usually
bisexual terminal spikes of small usually monoecious apetalous
flowers, the 1-3 female usually pedicellate below, the congested
sessile male, these often 3 in each imbricate glandular bract. Disk
none. Calyces 3- or the male 2-lobed and membranous, urceolate,
the segments imbricate. Stamens 1-3, exserted, the slender filaments
nearly connate. Ovary 3-celled; styles apically free, undivided;
ovules solitary. Capsules globose, lightly tridymous, the cocci
bivalved. Seeds obovoid, foveolate or smooth with large fleshy
caruncle.
Baillon, Adansonia 5: 332. 1865, referred this to Stillingia
Garden, a genus only recently found in Peru, characterized by the
pericarp persisting radiately between the cocci; later, Hist. PI. 5:
133. 1874, he placed it in Excoecaria L., more reasonably probably
from a broadly conservative viewpoint.
FLORA OF PERU 199
Maprounea guianensis Aublet, PI. Guian. 2: 895. pi. 342.
1775; 177.
Small tree sometimes 15 meters high; petioles slender, 5-15 mm.
long; stipules ovate, obtuse; leaves ovate or oblong-elliptic, sometimes
lanceolate, firm-membranous or more rigid, acute or subacute both
ends or obtuse at base where usually maculate-biglandular, lustrous
above, the nerves and reticulation slender, commonly 2.5-7 cm. long,
1.5-3.5 cm. wide; spikes peduncled, cylindric-ovoid, 3-6 mm. long,
rarely longer, 3 mm. thick; bracts (both sexes) broadly ovate, thin,
shortly acute, eglandular; female pedicels 5-6 mm. long; calyces
with broadly ovate denticulate subobtuse lobes; capsules 4 mm. long,
5 mm. thick, the coarsely foveolate dark brown seeds 3-3.5 mm.
long and thick with the caruncle. Mueller Argoviensis in Mart.
Fl. Bras. 11, pt. 2: 543. 1874 noted three variants. Williams, Field
Mus. Bot. 15: 279. 1936, described the wood which he found used
"rather extensively for general carpentry," the tree, sometimes 15
meters high, being characteristic in the Tarapoto plain and in the
hills around 500 meters. Illustrated, Mart. Fl. Bras. 11, pt. 2. pi. 81.
San Martin: Tarapoto, Ule 6504; Williams 5427; 5899; 5900.
Lomas, Williams 6880; 6470. Moyobamba, Weberbauer 4477; 290.
Zepelacio, King 3744- To the Guianas, Brazil and Bolivia.
"Airana," "arean" (Williams).
48. HURAL.
Reference: Pax & Hoffm. Pflanzenreich IV. 147. v: 271-274. 1912.
Spiny-trunked tall glabrous tree with abundant bistipulate alter-
nate petioled more or less cordate bright green leaves and terminal
reddish often cone-shaped spikes of sessile male flowers, each finally
irregularly rupturing bract enclosing one flower, the stoutly pedicelled
female borne solitary in the upper axils or at the base of the male
spike. Calyces and bracts eglandular, the male membranous, cupu-
late, truncate-denticulate, the female coriaceous, entire, laxly en-
closing 5-20-celled ovary, the high-connate fleshy styles radiately
spreading at tip. Stamens often many, indefinite, the filaments
continuous with the connectives into a fleshy apically dilated
column, the adnate distinct extrorse anthers 2-many-verticillate.
Hura crepitans L. Sp. PI. 1008. 1753; 272.
Branchlets sometimes aculeate; stipules linear-lanceolate, pubes-
cent, 1.5 cm. long; petioles as long as or longer than the basally
200 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
rounded-cordate orbicular-ovate leaves, these often 1-2 dm. wide
and as long or longer, abruptly short-acuminate, remotely repand
dentate or subentire, often pilose on the prominent nerves
beneath, dull both sides; peduncles 5-8 cm. long, the cylindric-ovoid
spikes finally to 6 cm. long, about 2 cm. thick; anthers usually
2-verticillate; fruiting pedicels about equaling petioles; ovary with
style 3-5 cm. long, the stigma reddish-violet; capsules concave both
ends, deeply sulcate, 8 cm. broad, half as long at maturity, expelling
the wafer-like seeds with considerable force. Williams, Field Mus.
Bot. 15: 277. 1936, gives the anatomy of the wood, which is used
in general carpentry, and describes the tree as erect, the columnar
trunk to 18 dm. in diameter, to 40 meters high, armed below with
stout conical spines, the pale bark with shallow ridges, the latex
said to be injurious to eyes and employed locally as a fish poison
as observed by Martius. See Pax, I.e., for variously reported uses
of the sap and bark in medicine. Said to be cultivated in most
tropical regions. Illustrated, Mart. Fl. Bras. 11, pt. 2: pi. 86; also
Pax, I.e. p. 272.
San Martin: Tarapoto, Williams 6318. Loreto: Lower Itaya,
Williams 111. Caballo-Cocha, Williams 2303. Yurimaguas,
Williams 4243- West Indies to Bolivia and Central America.
"Catahua" (Williams).
HIPPOCRATEACEAE Juss.
Reference: A. C. Smith, Brittonia 3: 341-555. 1940.
Glabrous or lightly puberulent lianas, shrubs or trees, the
branches and simple petioled leaves usually opposite, the stipules
if present interpetiolar, minute. Flowers small, hermaphrodite,
actinomorphic, bracteolate, usually pedicellate, in compound in-
florescences that are axillary or on short branches below the leaves
or pseudoterminal. Sepals and petals usually 5, the former imbricate,
persisting, as often also the latter, these alternate with sepals,
suberect to spreading, inserted beneath or without the edge of the
disk, this various in development. Stamens ordinarily 3 (2, 4 or 5),
inserted within the disk, the filaments often recurved, the anthers
basifixed, 2-celled, never introrsely dehiscent, usually laterally
contiguous, the clefts confluent at least apically, rarely dorsally
connate, the vertical or oblique clefts distinct. Ovary superior,
commonly adnate to or within the disk, usually 3-celled, 2-14 ovules
in a cell, these rarely massed, the style short or obsolete with 3 (-5)
FLORA OF PERU 201
entire or bifid stigmas. Fruit capsular or drupaceous, if the former
composed of 3 separate or connate capsules, if the latter indehiscent.
Seeds few to many, endosperm lacking, cotyledons large, radicle
small, basally winged in capsules.
The position of the stamens distinguishes the family from the
similar Celastraceae which has the stamens without the disk or
fused with it, and introrse dehiscence. This compilation follows
closely or is copied (with much appreciation) from Smith's scholarly
revision, except that it is adapted to my feeling that taxonomy
may endeavor, especially in floristic work, to serve greater numbers
of students, who often have need for only generic indications. The
argument of Smith that "it is by no means difficult to distinguish
the 12 genera recognized" introduces a new factor in determining
criteria for drawing generic lines; in admitting more broadly defined
genora to include subgroups as sections many will find the taxonomy
simplified, without having the relationships, so ably discerned or
redefined by Smith, obscured.
Disk discontinuous, forming staminal pockets; stigmas sessile; fruit
a drupe 1. Cheiloclinium.
Disk continuous; style present; fruit drupaceous or capsular.
Anther clefts vertical or oblique but not confluent; fruit a drupe.
2. Peritassa.
Anther clefts horizontal or oblique, confluent.
Fruit drupaceous; ovary if angled not deeply 3. Salacia.
Fruit capsular; ovary deeply sulcate-angled.
Capsules attached separately; petals entire or merely erose.
4. Hippocratea.
Capsules connate laterally; petals finely serrate. 5. Anthodon.
1. CHEILOCLINIUM Miers
Kippistia Miers, Trans. Linn. Soc. 28: 416. 1872, not F. Muell.
1858.
Various in habit but usually at least the upper branches scandent,
glabrous or essentially and much-branched with axillary thyrsoid,
corymbose or cymose inflorescences, dichotomously branched, the
flowers often crowded and paired or ternate at the tips. Sepals
usually erose or fimbriate, the petals entire, ordinarily thin-carnose.
Disk broken into as many saccate depressions as the stamens which
they enclose and which protrude slightly, usually 3, sometimes 5,
202 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
rarely 4. Anthers broader than long, extrorsely mitant, dehiscing
by horizontal confluent clefts. Ovary subglobose, trigonous or some-
times pentagonous, truncate or depressed, the 3-5 stigmas sessile,
entire to bilobed; ovules 2-4 per cell. Fruit subglobose, coriaceous,
soon 1-celled with 2-6 oblong- or ovate-angled seeds imbedded in
a mucilaginous pulp. Miers, as Smith, stresses the position of the
stigmas opposite the stamens and alternate with the ovary cells
as sometimes in Salacia, sens. lat. and the latter suggests that this
is a derivative condition. Besides the following there are a number
of species from the Acre and Solimoes basins that may occur within
Peru, mostly with narrow not lobed stigmas. Genus placed in
Celastraceae by Loesener.
Inflorescence thyrsoid; shrub or slender tree; leaves acuminate,
venation above obscure C. cognatum.
Inflorescence dichotomous; usually lianas; leaf venation obvious.
Stamens normally 3; leaves entire or obscurely crenate; leaves
acuminate except C. obtusum.
Peduncles 1-3 cm. long; stigmas sub truncate C. Klugii.
Peduncles 4-8 mm. long; stigmas minutely lobed.
Inflorescence branched 3-6 times; stigma lobes obscure.
C. obtusum.
Inflorescence branched 6-9 times; stigma lobes obvious.
C. hippocrateoides.
Stamens normally 5; leaves usually crenate-serrate, acuminate;
peduncle usually to 13 mm. long C. anomalum.
Cheiloclinium anomalum Miers, Trans. Linn. Soc. 28: 420.
1872; 548. Salacia anomala (Miers) Peyr. in Mart. Fl. Bras. 11,
pt. 1:144. 1878.
Much like C. hippocrateoides but the stamens, stigmas and ovary
cells 5, the stigmas minutely lobed, the leaves often serrate but
subentire in Peru, usually smaller, 7-15 cm. long, 3-5.8 cm. wide,
peduncle usually to 13 mm. long; pedicels 0.8-4 mm. long; sepals
entire; petals carnose, 1.3-1.8 mm. long, 1-1.5 mm. wide, usually
erose and conspicuously swollen within near the base; fruit to 3 cm.
long, 2.5 cm. broad, coriaceous, lustrous, the seeds about 15 mm.
long, 10 mm. wide. Illustrated, Miers, I.e. pi. 31. F.M. Neg. 19584.
Loreto: Mouth of the Santiago, Tessmann 4207. Brazil; British
Guiana.
FLORA OF PERU 203
Cheiloclinium cognatum (Miers) A. C. Sm. Brittonia 3: 529.
1940. Kippistia cognata Miers, Trans. Linn. Soc. 28: 417. 1872.
Slender, sometimes tree-like and as much as 12 meters high,
younger branchlets terete or subterete; petioles rugose, 3-8 mm.
long; leaves chartaceous, olive-green, midnerve prominent both
sides, the venation usually obscure, subentire or crenate, slightly
revolute, oblong, elliptic- or ovate-oblong, 8-20 cm. long, 2.5-8 cm.
wide, obtuse or subacute at base, obtusely long-acuminate; thyrse
shortly peduncled, to 10 cm. long, the branches swollen in fruit,
rachis and branchlets straight; pedicels very short; sepals broad,
obtuse, entire or nearly; petals suberect, thick, 1.5-2.6 mm. long,
1.2-2 mm. wide, opaque or obscurely lineate; fruits spherical or
oblong-ellipsoid, to 3.5 cm. long, 3 cm. broad, often lenticellate,
the seeds ovate, to 15 mm. long. C. lineolatum A. C. Sm. of Ama-
zonian Brazil has angled branchlets, leaf venation conspicuous,
petals lineate. F.M. Neg. 19585.
San Martin: Chazuta, Klug 4084. Huanuco: In forest, Rio
Cayumba at Rio Huallaga, Mexia 8287. Junin: Chanchamayo
Valley, Killip & Smith 24005; Schunke A42. Loreto: Mouth of
Rio Santiago, Tessmann 3955; 4393. Bolivia and Rio de Janeiro
to Panama and Tobago.
Cheiloclinium hippocrateoides (Peyr.) A. C. Sm. Brittonia
3: 546. 1940. Salacia hippocrateoides Peyr. in Mart. Fl. Bras. 11,
pt. 1: 142. 1878.
Liana with elongate slender slightly nodally swollen branches,
rugose petioles 7-16 mm. long and elliptic-oblong leaves, 8.5-20 cm.
long, 3.5-11 cm. wide, obtuse to subattenuate at the decurrent base,
cuspidate and obtusely acuminate, chartaceous, drying greenish
or olivaceous, entire or undulate crenulate, more or less conspicuously
venose both sides; inflorescence branched 6-9 times, peduncle
obsolete to 8 mm. long, branches slender, bracts papyraceous, flowers
subsessile or pedicels to 2 mm. long; sepals membranous, usually
ciliolate; petals thin-carnose or submembranous, 0.8-1.5 mm. long
and broad, glabrous or puberulent, subentire to fimbriolate; stigmas
broadly oblong, deeply lobed; immature fruits ellipsoid, smooth.
Name originally in herbaria by Richard, probably from French
Guiana. Illustrated, Smith, I.e. pi. 12. F.M. Neg. 35944.
Loreto: Mishuyacu, Klug 154- Bolivia and Brazil to the Gui-
anas.
204 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Cheiloclinium Klugii A. C. Sm. Brittonia 3: 543. 1940.
Liana with terete branchlets slightly swollen at the nodes and
chartaceous entire or obscurely crenulate leaves, dark green when
dried, paler beneath, conspicuously reticulate-veined both sides,
elliptic-oblong, 6.5-11.5 cm. long, 3-5 cm. wide, acute at base on
the 6-10 mm. long petioles, broadly acuminate and cuspidate;
inflorescence 3-7 cm. long, 5-7 times dichotomously branched, the
branchlets slender, as the peduncle, this 1-3 cm. long, pedicels very
slender, to 3 mm. long; sepals membranous, subentire; petals thin-
carnose, 1.3 mm. long, 1.1 mm. wide, minutely erose, obscurely
glandular-lineate; stigmas broadly oblong, obscurely emarginate
or truncate.
San Martin: In forest near Moyobamba, Klug 3689, type.
Cheiloclinium obtusum A. C. Sm. Brittonia 3: 545. 1940.
Tree, the type to 11 meters, with stout probably scandent
branches swollen at the nodes, coriaceous leaves, drying brownish,
broadly elliptic, abruptly cuneate at base and decurrent on petiole
8-12 mm. long, rounded or obtuse at apex, entire and often revolute,
usually plane above, the nerves slightly raised beneath; inflorescence
rigid, 3-5 cm. long, branched 5-6 times, peduncle stout, 4-6 mm.
long; flowers subsessile; sepals thin-carnose, the inner scarious and
erose-fimbriate-margined; petals carnose, 0.8-1 mm. long and broad,
subentire or minutely fimbriolate, obscurely puberulent without;
stigmas minutely bilobed. C. Krukovii A. C. Sm., 544, Amazonian
Brazil, has obtusely acuminate leaves, longer and glabrous petals.
Loreto: Mishuyacu near Iquitos, Klug 381, type.
2. PERITASSA Miers
Similar to Salacia from which it is scarcely distinguishable
especially in fruit but the stamens distinctive, the cells being vertical,
the connective more or less developed and the dehiscence by longi-
tudinal or oblique not confluent clefts; from the more typical species
of Salacia it is also characterized by its free short-tubular disk,
erect stamens, and petals. The stigmas are always obscure, which,
however, is a character recurring in species-groups in the allied
genus. There is one species known which approaches Salacia, P.
retusa (Miers) A. C. Sm., as the anther cells diverge at base, the con-
nective obscure; nevertheless these similar plants, it seems to me,
may be grouped together separately for convenience and, apparently,
naturally.
FLORA OF PERU 205
Leaves obviously crenate-serrate.
Anther connective long-extended; branchlets lenticellate. P. dulcis.
Anther connective not extended; branchlets tuberculate.
P. Killipii.
Leaves entire or obscurely crenate.
Leaves narrowly long-acuminate, distinctly bullate .... P. bullata.
Leaves acute or broadly and shortly acuminate, ebullate.
Inflorescence glabrous.
Anther connective extended; leaves obscurely nerved, obtuse
or rounded at base P. laevigata.
Anther connective included; leaves rather prominently
nervose, acute at base P. peruviana.
Inflorescence somewhat tomentulose, glabrate in age.
P. huanucana.
Peritassa bullata A. C. Sm. Brittonia 3: 513. 1940.
Liana with lenticellate branchlets, opposite or subopposite con-
spicuously bullate leaves and much-branched sessile or subsessile
inflorescences not more than 2 cm. long at anthesis; petioles 3-6 mm.
long; leaves obscurely crenulate, chartaceous or thin-coriaceous,
drying dark brown, narrowly elliptic, acute at base, caudate (acumen
1-2 mm. wide, 10-20 mm. long, obtuse), secondary nerves prominent
both sides, 7-14 cm. long, 1.7-3.7 cm. wide; bracts ovate, less than
1 mm. long; flowers apparently solitary at tips of ultimate branchlets,
the slender pedicels 1-1.5 mm. long; sepals papyraceous, narrowly
scariose and erose-fimbriolate; petals thin-carnose, about 1 mm.
long and half as wide, obscurely erose and sometimes a little farinose
within; disk undulate-crenulate; anther clefts slightly oblique, the
connective not prolonged; fruiting inflorescence swollen especially
the pedicels, the few developing fruits glaucous when young, smooth.
Loreto: In forest near Florida, mouth of Rio Zubineta, Klug
2298, type.
Peritassa dulcis (Benth.) Miers, Trans. Linn. Soc. 28: 403.
1872; 506. Salacia dulcis Benth. in Hook. Journ. Bot. Misc. 4: 9.
1852. S. Colasi Benoist, Bull. Soc. Bot. France 80: 335. 1933, fide
Smith.
Shrub or small tree with slender usually conspicuously lenticellate
branchlets (the lenticels soon tuberculate) and opposite or sub-
opposite prominently crenate-serrate leaves, their rugose petioles
206 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
5-12 mm. long, their blades narrowly elliptic, acute at base, rather
abruptly and shortly acuminate, the acumen obtuse or callose-
apiculate, 10-15 cm. long, 3.5-5 (7) cm. wide, chartaceous, drying
brownish, the secondary nerves prominent beneath, the veinlets
usually obscure; inflorescence compact, 1-3 cm. long, divided to
base or with short peduncle, dichotomous, the slender branchlets
usually rusty short-hirsute, the usually connate bracts 1.5-2 mm.
long, the pedicels scarcely 1 mm. long; flowers in clusters of 3-5,
bractlets minute; sepals papyraceous, rounded, narrowly scarious and
subentire; petals submembranous, often glandular-lineolate, 2-2.8
mm. long, about 1 mm. wide, the scariose margins erose; disk mem-
branous, crenulate; anther connective acute, produced 0.1-0.2 mm.;
fruit depressed-globose to 4 cm. thick, chartaceous, the 6 seeds oval,
to 13 mm. long, 1 cm. broad, the mucilaginous pulp said to be sweet,
edible. Type of S. Colasi not seen, reported as from Iquitos. Illus-
trated, Trans. Linn. Soc. 28. pi 26. F.M. Neg. 13353.
Loreto: Near Iquitos, Tessmann 3703a; Killip & Smith 27202;
King 904; Williams 3671; 1501. Rio Mazan, Jose Schunke 349.
Rio Itaya, Williams 16. Brazil.
Peritassa huanucana (Loes.) A. C. Sm. Brittonia 3: 521.
1940. Hippocratea huanucana Loes. Repert. Nov. Sp. 1: 163. 1905.
Liana with closely (soon obscurely) brown-tomentulose compactly
branched inflorescences and glabrous thin-coriaceous entire leaves
on stout petioles 6-10 mm. long; leaf blades brownish when dried,
elliptic- or ovate-oblong, 9-18 cm. long, 3.5-7.5 cm. wide, rounded
to acute at base, cuspidate or short-acuminate, the obtuse acumen
3-10 mm. long, the secondary nerves nearly plane above, spreading,
elevated beneath; inflorescence dichotomously branched several
times, finally glabrate, the bracts conspicuous, 1-2 mm. long, as
often the bractlets, the flowers paired or in ultimate dichotomies;
pedicels about 1 mm. long, usually swollen; sepals rounded or obtuse,
erose or fimbriolate, scarious-margined; petals suberect, 1.3-1.9 mm.
long, 0.7-1 mm. wide, narrowly scariose and entire, sometimes
glandular-punctate; disk undulate-crenulate; anther connective not
produced; ovules 4 (probably sometimes 2) in each cell. Like P.
perumana but with spreading instead of sharply ascending basal
secondary leaf nerves. F.M. Neg. 13339.
San Martin: Near Moyobamba, 800 meters, Weberbauer 4575,
type. Huanuco: Near Monzon, 900 meters, Weberbauer 3483; 285.
Loreto: Mouth of Rio Santiago, Tessmann 4303. Pongo de Man-
FLORA OF PERU 207
seriche, Tessmann 4870. Mishuyacu near Iquitos, King 1361.
Brazil? Guiana?
Peritassa Killipii A. C. Sm. Brittonia 3: 516. 1940.
With much resemblance to P. dulcis but the brownish branchlets
conspicuously tuberculate-scabrous, the leaves papyraceous or thin-
chartaceous, sometimes drying olivaceous, the inflorescence com-
paratively few-flowered, only twice branched, scabrous, the petals
only 1.3-1.6 mm. long, the anther connective not prolonged beyond
the cells. Differences slight but unless intermediate states are
developed they may be significant.
Loreto: In dense forest near Puerto Arturo below Yurimaguas,
Kittip & Smith 27777, type.
Peritassa laevigata (Hoffmsg.) A. C. Sm. Brittonia 3: 508.
1940. Tonsella laevigata Hoffmsg. ex Link, Jahrb. Gewachsk. 1,
pt. 3: 68. 1820. Salacia laevigata (Hoffmsg.) DC. Prodr. 1: 570.
1824. S. amazonica Loes. Verh. Bot. Ver. Brandenb. 48: 179. 1907.
Glabrous, at least the branches scandent, the branchlets notably
lenticellate; petioles stout, 8-18 mm. long, opposite or nearly so;
leaves elliptic-oblong, sometimes black-punctate beneath, obtuse
to rounded at base, subobtuse to shortly acuminate, 10-19 cm. long,
4.5-9.5 cm. wide, coriaceous, entire or somewhat crenulate and
thickened at margins, the nerves little if at all impressed above,
raised beneath; inflorescence 2-8 cm. long, sessile or nearly, much
enlarged in fruit, the papyraceous bracts minute, the obscurely brac-
teolate flowers laxly borne on pedicels 1-2 mm. long; sepals papyra-
ceous, often very unequal, scarious and subentire; petals thin, 2-2.8
mm. long, 1-1.5 mm. wide, usually erose, sometimes faintly glandular-
punctate or lineate; disk rather thin, crenulate; anther connective
obtuse, produced 0.1-0.2 mm.; fruits subglobose, to 3.5 cm. thick,
stiped, coriaceous, with 2-4 seeds about 2 cm. long, 13 mm. wide.
Apparently somewhat variable in shape and texture of leaves;
twice illustrated but, according to Smith, not accurately. F.M.
Negs. 13346 (S. amazonica); 13366; 35945 (synonyms).
Moore reported the pulp surrounding seeds greatly esteemed,
probably the case for all species, including those of Salacia.
Loreto: Rio Mazan, Jose Schunke 227. Brazil; Venezuela.
"Gogo" (Krukoff).
Peritassa peruviana (Miers) A. C. Sm. Brittonia 3: 519. 1940.
Sicyomorpha peruviana Miers, Trans. Linn. Soc. 28: 411. 1872.
208 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
More or less scandent, the branchlets lenticellate, the slender
petioles 6-8 mm. long; leaves chartaceous, drying brownish, promi-
nently reticulate both sides, entire or obsoletely crenulate, obovate-
elliptic, acute to subattenuate at base, obtusely short-cuspidate at
apex, 10-15 cm. long, 4-6.5 cm. wide; inflorescence 2-3 cm. long,
dichotomously or pseudodichotomously branched, the peduncle 10-
13 mm. long, bracts less than 1 mm. long, pedicels of the laxly
arranged flowers about 2 mm. long; sepals rounded, fimbriolate at
margins as the oblong petals; disk entire; ovary depressed, trigonous.
In spite of the name the type came probably from Guayaquil,
Ecuador, this information being with the Madrid specimen. F.M.
Neg. 29390.
Peru(?), cf. remark above. Ecuador.
3. SALACIAL.
Tontelea Aublet, PI. Guian. 1: 31. 1775. Thermophila Miers,
Trans. Linn. Soc. 28: 398. 1872, fide A. C. Smith.
Lianas or if shrubby or slender trees the upper branches usually
more or less scandent, glabrous or nearly, the opposite or subopposite
branches and branchlets often somewhat swollen at the nodes and
also a little compressed. Leaves sometimes alternate, the usually
grooved petioles often rugulose. Inflorescence axillary or on defoliate
branchlets, thyrsoid paniculate, cymose, or more or less dichoto-
mously branched, the flowers congested or lax at the ends of the
bracted branchlets. Sepals 5 (4-6), usually rounded, often unequal
and erose; petals 5 (6), rather similar. Disk various, annular-pulvi-
nate, truncate-conical, flattened, cupulate or short-tubular, free or
adnate, entire to crenulate. Stamens 3 (4), erect to reflexed, the
anthers dehiscing by apical confluent or rarely merely contiguous
clefts. Styles short to rather long, the stigmas 3 (4) obscure or
developed, alternate or opposite to the stamens, entire to lobed
sometimes so deeply that the stigmas appear to be 6 and the position
is thus obscured. Ovules 2 (rarely 6 or 8), the fruits subglobose or
ellipsoid, the few to many angled seeds imbedded in a mucilaginous
pulp. The genus Tontelea is maintained by Smith chiefly on the
basis of its short- tubular disk; in view of the variability in develop-
ment of the disk everywhere it seems to me more reasonable and
convenient to regard it at most as a section as by Triana and
Planchon and by Loesener.
Thermophila cordata Miers, Trans. Linn. Soc. 28: 401. 1872,
omitted here, based on a single leaf and a detached flowerless panicle
FLORA OF PERU 209
n the British Museum; not seen by Smith, the identity not estab-
ished; the type labeled by Pavon from Peru could be from Ecuador,
is, for example, was the case for Peritassa peruviana! The key,
argely after Smith here as elsewhere, needs material for comparison
is observed by him.
Stigmas obscure; inflorescences dichotomously branched or thyrsoid
paniculate, often elongate, rarely compact (S. spectabilis);
fasciculate in S. macrantha, S. impressifolia and S. juruana.
Style short; disk not higher than thick; petals smooth.
Disk flat, thicker than high, uniform; leaves often acute or
narrowly rounded at base; inflorescence fasciculate except
S. opacifolia.
Inflorescence 2-5 cm. long.
Petioles 2-7 mm. long; flowers 5-6 mm. wide. S. opacifolia.
Petioles 7-20 mm. long; flowers 12-30 mm. wide.
Flowers 24-30 mm. wide S. macrantha.
Flowers less than 22 mm. wide.
Lateral nerves obscure; leaves to 8.5 cm. wide.
S. impressifolia.
Lateral nerves obvious; leaves mostly wider .S. juruana.
Inflorescence 2-2.5 dm. long S. insignis.
Disk erect, thicker above; leaves rounded at base. S. myrsinoides.
Style long; inflorescence thyrsoid-fasciculate, disk higher than
thick, lobed between stamens; petals tuberculate; inflorescence
compact, to 2.5 cm. long S. spectabilis.
Stigmas obvious or conspicuous; inflorescence various, but not
fasciculate, more than 5 (at most about 8) cm. long; disk
cupulate or short-tubular.
Stigmas entire; petals glabrous.
Leaves three to four times as long as broad S. attenuata.
Leaves two to three times as long as broad S. fluminensis.
Stigmas bilobed or emarginate; petals farinose or puberulent, at
least within or without, sometimes obscurely.
Stigmas more or less emarginate or bilobed but clearly distinct.
Stigmas barely emarginate, opposite stamens; petals obviously
farinose within S. emarginata.
Stigmas lobed, alternate with the stamens, petals puberulent
both sides or only within.
210 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Leaves thin, translucent, obtusely acuminate. S. corymbosa.
Leaves coriaceous, opaque, obtusely cuspidate. .S. coriacea.
Stigmas so deeply lobed as to appear like 6; petals glabrous
within.
Leaves chartaceous, acuminate; inflorescence not ceriferous.
S. divergens.
Leaves coriaceous, rounded or obtusely cusped; inflorescence
ceriferous S. Weberbaueri.
Salacia attenuata (Miers) Peyr. in Mart. Fl. Bras. 11, pt. 1:
149. 1878; 474. Tontelea attenuata Miers, Trans. Linn. Soc. 28:
384. 1872.
Slender, often scandent; petioles 6-14 mm. long; leaves elliptic-
oblong, 7-18 cm. long, 2.5-6.5 cm. wide, acute or attenuate at base
and decurrent, gradually acuminate (acumen obtuse, 8-15 mm.
long), entire, narrowly re volute, usually sparsely black-punctate
beneath, thin-coriaceous or chartaceous, brownish or dark green
when dried, secondary nerves 7-10, arcuate-ascending, often rather
prominent both sides, sometimes subimmersed, the veins usually
obscurely reticulate, inflorescence thyrsoid paniculate, much-
branched from base, 1.5-3 cm. long, bracts and bractlets less than
1 mm. long; sepals broader than long, scariose and erose or fimbriate,
the entire petals to 2.2 mm. long, 1.6 mm. wide or smaller; disk
erect-spreading, undulate or crenulate; stamens suberect, anther
clefts horizontal, confluent; stigmas deltoid, spreading, subacute,
alternate with stamens; ovules 2 per cell; flowers subsessile, the
pedicels or branchlets thick in fruit, the fruit at least 1 cm. in diam-
eter, smooth or minutely rugulose. Illustrated, Miers, I.e. pi. 28.
F.M. Neg. 13348.
Loreto: Rio Mazan, Jose Schunke 251. Balsapuerto, Klug 2900.
British Guiana; Brazil; Colombia?
Salacia coriacea (A. C. Sm.) Macbr., comb. nov. Tontelea
coriacea A. C. Sm. Brittonia 3: 480. 1940.
Liana with terete often conspicuously lenticellate branchlets;
stout petioles 9-17 mm. long; thick coriaceous leaves, opaque and
brownish when dried, elliptic-oblong, obtuse to acute at base, obtusely
cuspidate at tip, 7-30 cm. long, 4-10.5 cm. wide, sometimes black
punctate beneath, the 6-12 spreading secondary nerves elevated
beneath, the veins immersed or faint; inflorescence 1-3.5 cm. long,
FLORA OF PERU 211
minutely ceriferous-farinose; flowers subsessile, congested, to 3 mm.
wide; sepals 0.5-1 mm. wide and long, entire or obscurely fimbrio-
late, puberulent without, the petals so on both sides, to 2.2 mm.
long; disk erect, undulate at margin; stamens suberect, anther clefts
horizontal, confluent; stigmas spreading, deeply emarginate or
bilobed, alternate with stamens; ovules 2 per cell, fruiting branchlets
swollen, the young fruits obscurely rugose.
Loreto: Mishuyacu, Klug 234. To the Guianas.
Salacia corymbosa Huber, Bol. Mus. Goeldi 4: 580. 1906; 485.
Tontelea corymbosa (Huber) A. C. Sm. Brittonia 3: 485. 1940.
Scandent or erect or probably with scandent branches; petioles
5-9 mm. long; leaves oblong or narrowly elliptic-oblong, 8-15 cm.
long, 3-6.5 cm. wide, acute or cuneate at base, decurrent, caudate
or long-cuspidate at tip (slender acumen obtuse, 7-15 mm. long),
papyraceous, translucent, olivaceous when dried, sometimes sparsely
black punctate on either side, secondary nerves 6-9, typically strongly
curved, arcuate-ascending, acutely elevated both sides, veins promi-
nently reticulate; inflorescence 3-4.5 cm. long, thyrsoid paniculate,
divided from base; flowers solitary, 3-4 mm. wide, sepals obtuse,
entire or erose as the thin petals, these to 2.4 mm. long; disk suberect,
subentire; style stout, stigmas alternate with stamens obviously
bilobed but the lobes clearly united in pairs; ovules 2 per cell.
Illustrated, Smith, I.e. page 466 (flower). F.M. Neg. 1433.
San Martin: Pongo de Cainarachi, Klug 2707. Loreto: Cerro
de Canchahuaya, Rio Ucayali, Huber 1433, type. Mouth of Rio
Santiago, Tessmann 4720 (leaves broader, secondaries spreading).
Salacia divergens (A. C. Sm.) Macbr., comb. nov. Tontelea
divergens A. C. Sm. Brittonia 3: 491. 1940.
Liana with dark brown lenticellate branchlets and scarcely dif-
ferent from S. Weberbaueri but petioles 6-8 mm. long, leaves chart-
aceous, elliptic-oblong, 8-15 cm. long, 3-5.3 cm. wide, obtuse at
base, short-acuminate or conspicuously cuspidate (acumen stout,
6-10 mm. long, obtuse) and the inflorescence not ceriferous; flowers
about 3 mm. wide; pedicels 1.5-2 mm. long; disk prominently undu-
late.
San Martin: Chazuta, 260 meters, Klug 4120, type.
Salacia emarginata (A. C. Sm.) Macbr., comb. nov. Tontelea
emarginata A. C. Sm. Brittonia 3: 494. 1940.
212 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Liana; petioles stout, 15-25 mm. long; leaves broadly elliptic or
elliptic-oblong, 10-18 cm. long, 4.5-10.5 cm. wide, obtuse or cuneate
at base, obtusely callose-mucronate or short-cuspidate (acumen 3-5
mm. long, callosed), thin-coriaceous, sparsely black-punctate be-
neath, secondary nerves 7-9, interspersed with others of equal
prominence, spreading, elevated both sides, veins laxly reticulate,
prominent or immersed; inflorescence 3-8 cm. long, often lax, farinose-
ceriferous; flowers 4-5 mm. wide, pedicels 3-6 mm. long, ceriferous
as often the calyces; sepals entire, rounded; petals yellow, to 2.3 mm.
long, rounded, erose, conspicuously farinose within; disk suberect
or slightly spreading, obscurely crenulate; stigmas minutely emargi-
nate, opposite the stamens; ovules 2 per cell.
San Martin: Zepelacio near Moyobamba, 1,100 meters, King
3269, type.
Salacia fluminensis Peyr. in Mart. Fl. Bras. 11, pt. 1: 149.
1878; 477. Tontelea fluminensis (Peyr.) A. C. Sm. Brittonia 3: 477.
1940.
Allied to S. attenuata; petioles 7-25 mm. long; leaves olivaceous,
dried, 11-22 cm. long, 4.5-9 cm. wide, obtuse to acute at base,
cuspidate or shortly acuminate, secondary nerves 7-13, veins rather
prominent beneath, sometimes obscure above; inflorescence 3.5-4.5
cm. long; ovules 4 per cell. S. ovalifolia (Miers) Macbr., comb. nov.
Cuervo, ovalifolia Miers, Trans. Linn. Soc. 28: 371. 1872; T. ovali-
folia (Miers) A. C. Sm. I.e., 476 of Colombia and adjacent Brazil
has leaves rounded or obtuse at apex, the veins immersed, the in-
florescence short to 8.5 cm. long. F.M. Neg. 13354.
Loreto: Rio Mazdn; Schunke 232. Bolivia; Brazil.
Salacia impressifolia (Miers) A. C. Sm. Bull. Torrey Club 66:
247. 1939; 447. S. grandiflora (Benth.) Peyr. in Mart. Fl. Bras.
11, pt. 1: 157. 1878, not Kurz, 1872. Raddia impressifolia Miers,
Trans. Linn. Soc. 28: 392. 1872. Anthodon grandiflorus Benth. in
Hook. Journ. Bot. Misc. 4: 10. 1852.
Liana or scandent shrub related to S. macrantha, but the leaves
rarely decurrent, usually rounded or obtuse at base, usually gradually
acuminate, 8-21 cm. long, 3.5-8.5 cm. wide, the nerves if elevated
beneath not prominent; sepals to 2.6 mm. long, 2.6-4 mm. wide,
sometimes ciliate, often punctate or variegated as the petals, these
5.5-10 mm. long, 5-8 mm. wide; disk nearly of the other species;
anther clefts oblique; ovules 4 per cell. Illustrated, Vellosia 3,
FLORA OF PERU 213
pt. 1. pi. 8 (as S. polyanthomaniaca). F.M. Negs. 19587; 23265
(S. grandiflora) .
Junin: Cahuapanas, Rio Pichis, Killip & Smith 26720. Loreto:
Rio Itaya, Tessmann 5202. Rio Acre: Vie 9555. Amazonian
Brazil; Bolivia.
Salacia insignis A. C. Sm. Brittonia 3: 435. 1940.
Character in general that of S. opacifolia, but the petioles stout,
2-4 mm. thick, 10-16 mm. long, leaves pale green when dried, elliptic-
oblong, 25-28 cm. long, 9.5-10 cm. wide, with 14-18 secondary nerves
and elongate inflorescences from the branchlets below or between
the leaves, the panicles 2-2.5 dm. long on peduncles 3-5 cm. long
with many flowers 6-8 mm. wide at anthesis; disk thinner toward
outer margin, not at all elevated. A liana, the branchlets with loose
brown bark.
Loreto: In forest at Mishuyacu near Iquitos, Klug 74, type.
Salacia juruana Loes. Verh. Bot. Ver. Brandenb. 48: 181. 1907;
449.
Resembles the allied S. impressifolia and S. macrantha, especially
the former, from which it differs in having the 10-13 secondary
nerves very prominent beneath, the leaves 12-27 cm. long, 5-15 cm.
wide, rounded or obtuse at both ends; flowers 13-14 mm. wide,
the sepals 1-1.5 mm. long, 2-3 mm. wide, entire or a little ciliolate,
the petals 5-7 mm. long, 4-5 mm. wide, sometimes sparsely glandular;
anther clefts oblique; ovules 6-8 per cell. Pedicels are 8-15 mm.
long, in the similar S. gigantea Loes., 3-3.5 mm. long, flowers 15-18
mm. wide, style obsolete, ovules 2 per cell. As Smith remarks, the
species in this group are apparently closely related. F.M. Neg.
13358.
Loreto: Yurimaguas, Killip & Smith 29072. Mouth of Rio
Santiago, Tessmann 4672. Rio Acre: Mouth of Rio Macauhan,
Krukoff 5731. Adjacent Brazil.
Salacia macrantha A. C. Sm. Bull. Torrey Club 66: 245. 1939;
445.
Slender tree marked by the large (24-30 mm. wide) flowers borne
in few-flowered fascicles or glomerules in the axils or on defoliate
branchlets; petioles stout, 7-20 mm. long; leaves elliptic, oblong or
obovate-oblong, usually attenuate and decurrent at base, shortly
and obtusely acuminate, 15-32 cm. long, 4.5-11.5 cm. wide, cori-
214 FIELD MUSEUM OP NATURAL HISTORY BOTANY, VOL. XIII
aceous, olivaceous or greenish dried, often glaucous beneath, the
secondary nerves 7-11, often arcuate, more or less prominent beneath;
veins obscurely reticulate; sepals 3-4 mm. long and broad, entire,
narrowly scariose; petals 5 (6), thin, yellow or greenish, 10-13 mm.
long, 7-10 mm. wide; disk elevated at inner margin, conspicuously
flattened and membranous at the outer and concealing the petal-
bases; anthers dehiscing by horizontal apical confluent clefts; ovules
apparently 6 per cell; fruiting pedicels much swollen, the subglobose
fruit 3-4 cm. thick, very rugose and lenticellate.
San Martin: Chazuta, Klug 4047. Loreto: Mishuyacu, Killip
& Smith 29893; Klug 184. Adjacent Colombia and Brazil.
Salacia myrsinoides (A. C. Sm.) Macbr., comb. nov. Tontelea
myrsinoides A. C. Sm. Brittonia 3: 499. 1940.
Liana with terete striate branches; petioles 6-8 mm. long;
leaves elliptic- or ovate-oblong, rounded or broadly obtuse at base,
cuspidate (acumen about 1 cm. long, obtuse), 9-14 cm. long, 4-5.5
cm. wide, chartaceous, brownish when dried, secondary nerves 6-8,
arcuate ascending, obvious only beneath, the veins obscure; inflores-
cence on defoliate branchlets, 6-8 cm. long, thyrsoid-paniculate,
densely brown puberulent, bracts and bractlets soon caducous;
pedicels about 1 mm. long; flowers 2-2.5 mm. wide, sepals and petals
membranous, entire, the latter yellowish, oblong-elliptic, about 1 mm.
long; disk erect, entire, slightly thicker at tip than at base; anthers
dehiscing by confluent horizontal apical clefts; style short, conical,
truncate; ovules 2 per cell. Similar but leaves oblong or obovate-
oblong, abruptly short acuminate, inflorescence dichotomous and
glabrous is S. glabra (A. C. Sm.) Macbr., comb. nov. (T. glabra
A. C. Sm. I.e. 500), of adjacent Bolivia; ovary terete, petals spreading
but disk thicker at apex than at base and in this latter character
suggesting some Hippocratea species; probably variations of this
sort have occurred independently in each of these genera, so that
the character cannot be taken phylogenetically as significant. It
seems to me that Smith has overemphasized the importance of disk
development, which pretty obviously nearly exhausts the imagina-
tion in conceivable forms.
Loreto: Mishuyacu near Iquitos, Klug 712, type.
Salacia opacifolia (Macbr.) A. C. Sm. Brittonia 3: 434. 1940.
Hippocratea opacifolia Macbr. Field Mus. Bot. 8: 123. 1930. S.
gracilis A. C. Sm. Bull. Torrey Club 66: 249. 1939, fide Smith.
FLORA OF PERU 215
Branchlets cinereous, often prominently lenticellate; petioles
2-7 mm. long; leaves obovate-elliptic or narrowly elliptic, attenuate
to narrowly rounded at base, abruptly acuminate (acumen usually
rounded, 5-15 mm. long), entire or slightly undulate, 10-17 cm. long,
3.5-7.5 cm. wide, chartaceous or thin-coriaceous, brown or above
olivaceous when dried, secondary nerves 6-10, rather conspicuous
both sides as the reticulate veins; inflorescence axillary, 2-5 cm.
long, 3-5-times dichotomous, often from base, the papyraceous
bracts 1-1.5 mm. long; flowers sometimes paired, 5-6 mm. wide on
pedicels 8-12 mm. long; sepals sometimes punctate, erose, the obo-
vate petals faintly if at all so, 1.9-2.6 mm. long, often punctate;
disk flattened, uniformly thickened; anther dehiscence by horizontal
apical confluent clefts; style minute; ovules 2. Slender tree; attains
7 meters.
Junin: Chanchamayo Valley, 1,500 meters, Schunke 305, type.
Yapas on Pichis Trail, Killip & Smith 25498. Cahuapanas on Rio
Pichis, Killip & Smith 26733. Loreto: Yurimaguas, Killip & Smith
28024. Brazil.
Salacia spectabilis A. C. Sm. Brittonia 3: 436. 1940.
Slender tree with stout closely lenticellate branches; petioles 4-5
mm. thick, strongly rugose, flattened above, 2.5 cm. long or longer;
leaves elliptic-oblong, obtuse or acute at base (apex unknown),
25-37 cm. long, 10-14 cm. wide, drying olivaceous, chartaceous,
secondary nerves 11-13, ascending, straight, the veins horizontally
reticulate; inflorescence on defoliate branchlets, 1.5-2.5 cm. long,
compactly paniculate, the peduncle short, the clustered flowers
3-3.5 mm. wide; pedicels about 1 cm. long; sepals membranous,
cuspidate or subacute, subentire or erose as the pink orange-tinted
elliptic-oblong petals, these irregularly tuberculate within; disk
conic-pulvinate at margins, produced into 3 deltoid teeth alternate
with stamens, these reflexed and with anthers dehiscing by hori-
zontal apical confluent clefts; ovary connate, the style filiform-subu-
late, truncate or capitate, ovules 2 in each cell. S. acreana A. C.
Sm., with similar disk-teeth, has thicker petioles, still larger leaves,
the veins immersed and inflorescence 7-12 cm. long.
Loreto: In dense forest near San Antonio, Rio Itaya, Killip &
Smith 29416, type.
Salacia Weberbaueri (A. C. Sm.) Macbr., comb. nov. Tontelea
Weberbaueri A. C. Sm. Brittonia 3: 489. 1940.
216 FIELD MUSEUM OP NATURAL HISTORY BOTANY, VOL. XIII
Liana, with stout conspicuously lenticellate branches; petioles
11-13 mm. long; leaves obovate-elliptic, cuneate at base, rounded
or obtusely cuspidate at apex, 8-14 cm. long, 4.5-7.5 cm. wide,
coriaceous, olivaceous above and brown beneath (dried), secondary
nerves 8-11, spreading, prominent both sides, veins obscurely
reticulate; inflorescence 1-2.5 cm. long, peduncle stout; branchlets
densely farinose-ceriferous; flowers subsessile, crowded, 3.5 mm.
wide, sepals rounded, erose or fimbriolate, petals submembranous,
about 2 mm. long, erose, sometimes puberulent; disk erect-spreading;
stigmas deeply bilobed, the segments of the stigmatic surface
essentially equal; ovules 2 per cell; fruiting inflorescence much
swollen, the oblong-ellipsoid fruits to 3.5 cm. long, 1.5 cm. thick,
blackish and rugose in herb. Stigmas apparently opposite the
stamens. T. cuspidata A. C. Sm., I.e. p. 490, of Bolivia has thinner
proportionately narrower leaves, conspicuously cuspidate at tip.
Illustrated, Smith, I.e. p. 466 (flower).
Libertad: Above Ongon, valley of the Mixiollo, 1,700 meters,
Weberbauer 7068, type.
4. HIPPOCRATEA L.
Pristimera Miers, Trans. Linn. Soc. 28: 360. 1872. Cuervea [Tr.
& PL] Miers, I.e. 370.
Lianas or at least the branches finally scandent. Inflorescence
paniculate-corymbose, axillary, congested at branchlet tips or on
short leafless branchlets, the flowers apically crowded, lateral or
lax, always pedicellate and with 5 imbricate rounded sepals, 5
spreading petals, 3 erect to reflexed stamens, the anthers dehiscing
by confluent horizontal clefts. Ovary acutely trigonous or early
obviously 3-lobed, style slender to rather stout, stigmas obscure to
rather prominent, alternate or opposite the stamens, the 3 cells
each with 4-6 (2-8) ovules. Fruiting inflorescence more or less
swollen, developing 1-several capsular fruits consisting of 3 separately
attached dehiscing flat or convex flabellate pods with 2-8 oblongish
seeds that are strikingly winged at base. The disk varies as in the
drupaceous fruited groups in development as to form, size and in-
dument (to absence) and is regarded by Smith as of taxonomic
importance.
Petals barbate, 2.5-4 mm. long; disk cylindric or truncate-conical.
H. volubilis.
Petals not barbate; disk various.
FLORA OF PERU 217
Stigmas opposite stamens; disk cylindric, prominent; petals 5-9
mm. long H. Kappleriana.
Stigmas alternate with stamens; disk minute, annulate; petals
1.2-2.2 mm. long.
Inflorescence nodes by contraction at base, pseudoarticulate;
disk 5-sided H. nervosa.
Inflorescence nodes not contracted; disk about circular.
Leaves obtuse or obtusely apiculate; ovules 6-8 per cell.
H. andina.
Leaves long-acuminate; ovules 4 per cell H. tenuiflora.
Hippocratea andina (Miers) Macbr., comb. nov. Pristimera
andina Miers, Trans. Linn. Soc. 28: 364. 1872; 376.
Liana, with often densely lenticellate branchlets; petioles 8-15
mm. long; leaves narrowly to broadly elliptic or ovate-elliptic, obtuse
to narrowed at base, obtuse or obtusely cuspidate at tip, 4-16 cm.
long, 1.5-7 cm. wide, undulate to obviously crenate, with 7-10
secondary nerves prominent beneath; inflorescence to 11 cm. long,
often much smaller, the bracts less than 1 mm. long; flowers 2.5-4
mm. wide, pedicels to 1.5 mm. long, sepals scariose, subentire or
erose, petals 1.3-2.2 mm. long, entire or nearly, sometimes glandular
or pulverulent; disk annular; filaments gradually wider at base;
style fleshy; ovules 6-8 per cell; capsules elliptic or obovate-elliptic,
to 5 cm. long, half as wide, the seeds with wings to 3 cm. long, half
as wide. Type locality San Luiz, "Peru, alt. 7,000 ft.," but locality
perhaps Bolivian. There are two earlier names very inadequately
published and I believe they should be ignored, since no specimens
are cited (Smith). Illustrated, Schnizl. Icon. pi. 237, figs. 10-15.
F.M. Negs. 13336; 13338 (synonyms).
Peru (?; see remark above). Southern Brazil and Bolivia to
Argentina.
Hippocratea Kappleriana Miq. Linnaea 26: 220. 1853; 399.
Cuervea Kappleriana (Miq.) A. C. Sm. Brittonia 3: 399. 1940.
Liana, the branchlets often rugose; petioles 8-13 mm. long; leaves
elliptic-oblong or ovate-oblong, more or less narrowed to rounded at
base, gradually acuminate (acumen 5-10 mm. long), entire or ob-
scurely crenulate, 8-19 cm. long, 4-10 cm. wide, the 6-9 secondary
nerves as the fine reticulation prominent beneath; inflorescence 5-12
cm. long, branching from near base; flowers white, fragrant, 10-17
218 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
mm. wide, pedicels 1.5-3 mm. long, sepals subentire, petals obovate-
elliptic, 5.5-9 mm. long, entire or a little crenulate; disk mem-
branous, short-cylindric; filaments much widened at base; style
stout, the stigmas usually conspicuous but not always; ovules 4-6
per cell; capsules broadly obovate or suborbicular, 5-9 cm. long,
5.5-10 cm. wide, abruptly stiped, 1-1.5 cm. thick, the seed wings
about 2 cm. long, 5 mm. wide.
Loreto: Rio Napo near Mazan, Mexia 6446a. To Central
America and the Guianas; West Indies.
Hippocratea nervosa (Miers) Macbr., comb. nov. Sicyomorpha
nervosa Miers, Trans. Linn. Soc. 28: 412. 1872. H. Ulei Loes. Verb.
Bot. Ver. Brandenb. 48: 177. 1907. Pristimera nervosa (Miers)
A. C. Sm. Brittonia 3: 370. 1940.
Branchlets elongate, sometimes a little quadrate; petioles 8-15
mm. long; leaves elliptic or broadly ovate-elliptic, 7-20 cm. long,
3-9 cm. wide, obtuse or acute at base, obtusely acuminate, undulate
or crenate, thin, the 7-9 secondary nerves prominent beneath;
inflorescence 5-8 times branched, to 7 cm. long, often much smaller,
the bracts and bractlets conspicuous; flowers 1.7-3.5 mm. wide,
pedicels less than 1 mm. long, sepals erose or ciliate, petals more or
less erect, 1.2-1.8 mm. long, entire; disk pentagonous, annular;
filaments broadened at base; style fleshy, ovules 4-8 per cell. F.M.
Neg. 13344 (H. Ulei).
San Martin: Near Tarapoto, Spruce 4486; Ule 6354 (type, H.
Uki). Loreto: Yurimaguas, Killip & Smith 28897. Cuzco: Cos-
nipata, 700 meters, Weberbauer 6948. To Panama, Guiana and
Bolivia.
Hippocratea tenuiflora Mart, ex Peyr. in Mart. Fl. Bras. 11,
pt. 1: 132. 1878; 382. Pristimera tenuiflora (Mart.) A. C. Sm.
Brittonia 3: 382. 1940.
Branchlets subterete or slightly angled; petioles 3-8 mm. long;
leaves narrowly elliptic, acute or obtuse at base, long-acuminate,
the slender obtuse acumen 1-2 cm. long, obsoletely crenulate or
undulate, secondary nerves 6-8, prominent beneath; inflorescence
1-2.5 cm. long, 3-5 times branched; bracts about 1 mm. long;
flowers 2-2.7 mm. wide, subsessile, sepals minutely erose, petals
also or subentire or glandular-punctate, 1.5-2 mm. long; disk slightly
elevated; filaments only little wider at base; ovules 4 per cell (5
according to Peyritsch). Illustrated, Mart. I.e. pi. 42.
FLORA OF PERU 219
Loreto: La Victoria, Williams 2638. Caballo-Cocha, Williams
2412. Brazil and adjacent Colombia.
Hippocratea volubilis L. Sp. PL 1191. 1753; 359. H. ovata
Lam. Encycl. 1: 100. 1791. H. viridis R. & P. Fl. Peruv. 1: 44.
1798. H. copiosiflora Miers, Trans. Linn. Soc. 28: 341. 1872. H.
perspicua Miers, I.e. 342.
Liana, the younger parts including the 2-6 times well-peduncled
inflorescences densely brown puberulent; petioles slender, some-
times margined, 4-18 mm. long; leaves ovate or elliptic or oblong-
elliptic, usually 6-14 cm. long, 3-7 cm. wide, rounded to attenuate
at base, often decurrent, rounded to short acuminate at apex, the
acumen sometimes 12 mm. long but obtuse; margins obviously
crenate or serrate, the teeth often callose-apiculate, or again merely
undulate or subentire; secondary nerves 5-10, prominent, especially
beneath; flowers 4-8 mm. wide, pedicels 1-3 mm. long; sepals erose
or ciliolate; petals thin, 2.5-4 mm. long, ciliolate and often puberulent
both sides as well as transversely barbellate within near tip, this
subacute; disk sometimes expanded at base as the ligulate filaments;
capsules obovate- or oblong-elliptic, 4-8 cm. long, 1.5-4 (5) cm.
wide, slightly swollen at base, rounded or emarginate at tip; seeds
sometimes apiculate, the obovate wing finally 2-4 cm. long, 10-25
mm. wide with prominent funicle to 4 mm. long. Illustrated, Smith,
I.e. p. 358. F.M. Negs. 35934 (H. copiosiflora); 29389 (H. viridis).
San Martin: Juan Guerra, Williams 6923. Tarapoto, Spruce
(type, H. copiosiflora); Williams 6651. Zepelacio near Moyobamba,
Klug 3676. Amazonas: Vitoc, Ruiz & Pavdn. Chachapoyas,
Mathews 3033 (type, H. perspicua). Huanuco: Pozuzo, Ruiz &
Pavdn (type, H. viridis). Loreto: Tessmann 4421. Rio Nanay,
Williams 980; 1206; 1218; 1219. Caballo-Cocha, Williams 2440.
To Mexico, Florida and Argentina. "Yaco yaje" (Williams).
5. ANTHODON R. & P.
Lianas vegetatively similar to Hippocrateas but the petals con-
spicuously and irregularly serrulate, the anthers broadly reniform,
the ovules 8-14 per cell and imbricate in a mass instead of biseriate
and the flattened capsules firmly connate by the emarginate or
scalloped wings.
Anthodon decussatum R. & P. Fl. Peruv. 1: 45. pi. 74. 1798;
420. Hippocratea Anthodon Pers. Syn. PI. 1: 41. 1805. Tonsella
220 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
decussata (R. & P.) Vahl, Enum. 2: 30. 1805. Tontelea decussata
(R. & P.) Poir. in Lam. Encycl. Suppl. 5: 316. 1817. Salacia decus-
sata (R. & P.) G. Don, Gen. Syst. 1: 627. 1831. H. decussata (R. &
P.) Peyr. in Mart. Fl. Bras. 11, pt. 1: 138. 1878.
Branchlets terete; petioles 5-12 mm. long; leaves elliptic or ovate-
elliptic, 4-11 cm. long, 1.5-5 cm. wide, obtuse or acute at base,
gradually acuminate (acumen to 1 cm. long), membranous, lustrous,
crenulate or serrulate, secondary nerves 7-9, prominent as reticulate
veins both sides; inflorescence 1.5-5 cm. long, a many-flowered
rather open cluster to 6 cm. broad, the slender peduncle 5-15 mm.
long, bracts 1-2 mm. long; flowers 5-8 mm. wide, pedicels slender,
2-5 mm. long, sepals ciliate or erose, petals 3-4 mm. long, to half as
wide, minutely serrulate (serrations 7-11 per mm.); disk short-
cylindric; filaments broader, base and apex; style stout, stigmas
minute, opposite stamens, sometimes a little plicate and appearing
alternate; capsule connate into a fruit to 14 cm. across with 8-14
seeds in each capsule, the wings to 3.5 cm. long, 1.5-2 cm. wide.
Illustrated, Smith, I.e. p. 419 (fruit); Mart. Fl. Bras. 11, pt. 1.
pi 44; HBK. Nov. Gen. & Sp. 5: pi. 443 (floral details inaccurate,
Smith). F.M. Neg. 13335.
San Martin: Near Tarapoto, Ule 6620 (type, H. decussata, var.
parviflora}. Huanuco: Near Pozuzo, Ruiz & Pavdn, type. Bolivia
to Venezuela and Colombia.
BUXACEAE. Boxwood Family
Usually ligneous plants, with evergreen coriaceous simple estipu-
late leaves, either alternate or opposite, the small monoecious or
dioecious apetalous flowers in bracteate spikes or dense racemes.
Sepals if present imbricate and usually 4 with as many stamens
opposite them or these 6 with 2 pairs opposite the inner sepals.
Anther cells bivalved or opening longitudinally. Female flowers
often larger than male and fewer or solitary. Ovary superior, 3-celled
with entire approximate or widely separated styles, the 1 or 2 ovules
pendulous. Fruit capsular or drupaceous with black lustrous seeds,
the endosperm fleshy, embryo straight with flat or thick cotyledons.
STYLOCERAS A. Juss.
Glabrous trees, with alternate entire coriaceous leaves, the veins
very prominent beneath, and sessile short dense male spikes in the
axils, the female flowers often solitary and pedicellate or sometimes
FLORA OF PERU 221
at the base of the male spike. Disk none. Calyx lacking in male
flowers, stamens 6-30, filaments very short, anthers oblong, erect,
receptacle adnate to bracts, rudimentary ovary none. Female
flowers with 5 sepals similar to the several bracts, ovary 2 (-4) -celled,
styles 2, long, entire, ovules solitary. Drupe fleshy but drying hard,
the styles persisting.
Styloceras laurifolia (Willd.) HBK. Nov. Gen. & Sp. 7: 173.
1825. Trophis laurifolia Willd. Sp. PI. 4: 733. 1806. S. Kunthianum
A. Juss. Euphorb. Tent. 117. pi 17, jig. 56? 1824.
Branchlets angled; petioles 1.0-2.5 cm. long; leaves oblongish,
obtuse or barely acute, 1-2 dm. long, usually 4-4.5 (6.5) cm. wide,
lustrous above, paler beneath; spikes about 2.5 cm. long, flowers
sessile, unibracteate, the ovate acute bracts ciliate; anthers often
about 12, sessile, stout, tetragonous, lightly curved; female flowers
axillary, solitary, peduncled, 8 mm. long, with many small bracts
and cupulate 4-lobed calyx; ovary usually 2-celled, the long styles
spreading-recurving. Ovary of S. Kunthianum 4-celled, capsule
globose, 2-3-horned, pleasantly scented. Illustrated, HBK. I.e.
pi 638.
Yields, according to Weberbauer, a good wood.
Libertad: Usquil, Weberbauer 7000; 6999. Huanuco: Rio Pozuzo,
Weberbauer 6788; 6733a. Junin: Huancayo, Pariahuano, 3,100
meters, Weberbauer 6995. Utcuyacu, Woitkowski 497 (det. Morton).
To Colombia. "Sapanque," "limoncita de Caches," "curape."
ICACINACEAE
Reference: Richard A. Howard, Journ. Arnold Arb. 21: 461-
489. 1940; 23: 55-78, 479-494. 1942; Contr. Gray Herb. 142: 1-92.
1942.
Ligneous plants, often trees, with exstipulate mostly alternate
leaves and bisexual or functionally unisexual 4-5-merous bracted
flowers, axillary or terminal, extra-axillary or supra-axillary but
always with small inferior imbricately lobed calyx (usually articulated
below) and valvate free or united, rarely absent, petals (usually 5,
rarely 4-6), the stamens alternate with them, the fleshy or filiform
filaments, if present, commonly pubescent, the anthers with 4,
rarely 2 cells, often deeply lobed. Disk often lacking, sometimes
fleshy. Ovary 1(3-5) -celled; ovules usually 2, pendulous from near
tip of ovary, collateral or rarely superposed. Style usually short,
222 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
stigma large or punctiform. Fruit drupaceous, l(-3)-celled, l(-3)-
seeded, symmetrical or flattened, the funicle in a special tubular
canal of the putamen or free in the sarcocarp; embryo minute, small
or large, the endosperm copious.
It is fortunate for the usefulness of this compilation that the work
of Dr. Howard appeared before its publication; his thoughtful
studies, which have resulted in obvious clarification of the taxonomy,
have been followed, it is hoped accurately, and my indebtedness is
acknowledged with pleasure.
Flowers unisexual or rarely a few perfect; fruits crested or appen-
daged.
Flowers 4-parted, the male spicate without rudimentary ovary,
the cymose female pubescent; fruit glabrous within, reticulate
crested without 1. Calatola.
Flowers 5-parted, cymose; ovary glabrous, rudiment present in
male flowers, fleshy pulvinus in female persisting as an ap-
pendage on fruit 2. Discophora.
Flowers perfect; fruits (known) neither crested nor appendaged.
Ovary as drupe 2-3-celled; axillary inflorescences fascicled-panicu-
late; petals reddish pubescent within (Peru). . .3. Emmotum.
Ovary 1-celled (rarely 2-celled, Citronella); petal pubescence, if
present, white or pale.
Petals glabrous within; fruit (known) partly divided, small, the
seed hippocrepiform.
Lianas; flowers not articulated, inflorescence axes often
flattened 4. Pleurisanthes.
Erect or only branches clambering; flowers articulated, axes
terete 5. Citronella.
Petals pubescent within; fruit without partial partition, 3.5-7
cm. long (known).
Ovary, as cell within, glabrous; trees 6. Poraqueiba.
Ovary, as cell within, pubescent; shrubs or lianas.
Inflorescence axillary; connective linear 7. Leretia.
Inflorescence terminal; connective broadly triangular.
8. Humirianthera.
1. CALATOLA Standley
Reference: Richard A. Howard, Contr. Gray Herb. 142: 13-5
1942.
FLORA OF PERU 223
Trees with revolute margined leaves (blackening in herbarium)
md axillary minute dioecious 4-merous flowers, the male valvate
md in dense pseudo-spikes, gamopetalous, the lobes with obvious
nidrib villous within, the female solitary or in racemes, the petals
^conspicuous. Anthers oblong, basifixed, longitudinally dehiscent.
Dvary cylindric, pubescent, the style obsolete. Drupes typically
bicrestate and usually reticulate dentate-crested; seed solitary, the
embryo minute.
Wood white and desirable because immune to insect-attack; the
fruits of some species are said to be edible at least when roasted and
contain, as the bark and leaves, a pigment used as a blue dye.
Gala tola venezuelana Pittier, Bol. Soc. Venez. Cienc. Nat. 4:
360. 1938; 18.
Branches terete, minutely appressed puberulent; petioles 1.5 cm.
long, almost winged; leaves broadly oval to oblong, acute at the
usually oblique base, abruptly acuminate to rounded at tip, 17-20
cm. long, 8-12 cm. wide, glabrous above, sparsely villous beneath
becoming glabrate except on the 10-12 pairs of veins; flowers not
known; drupes ovoid to subglobose, 4.5-6.5 cm. long, 3.5-4.5 cm.
in diameter, the putamen bivalved, sutures prominent, valves rugose,
the crests conspicuous and more or less dentate-reticulate. Known
to 17 meters tall with a trunk diameter of 4 dm. According to Pittier
the fruits finally dehiscent, recorded for no other species. Nearly
C. costaricensis Standley but there appear to be reliable differences
in the ornamentation of the putamen (Howard). Other species to
be expected include the Ecuadorean C. pastazana Sleumer and C.
colombiana Sleumer, both fewer leaf veins and fruits constricted or
conical at base, those of the former subglobose, 4 cm. long, of the
latter, ellipsoid-oblong, 5 cm. long. Illustrated, Notizbl. Bot. Gart.
Berlin 15: 249. fig. 4a and 6.
Fruits of this species are inedible unless roasted.
Huanuco: Shapajilla, Woytkowski 19. Colombia; Venezuela.
"Orosul," "venenito."
2. DISCOPHORA Miers
Reference: Richard A. Howard, Contr. Gray Herb. 142: 21-30.
1942. Kummeria Mart, ex Engler in Mart. Fl. Bras. 12, pt. 2: 52.
1872.
Trees or shrubs with short-strigose or essentially glabrous
branches, the bark smooth, and small unisexual or polygamous
224 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
flowers in axillary diffuse panicles with strong basal branches or
rarely strict, elongate and divaricate in fruit, the very short strigose
pedicels articulated above the bract. Calyx broadly and often
unevenly 5-lobed. Petals free, glabrous or nearly, the inflexed tip
mucronulate, midrib prominent. Filaments medially enlarged or
appendaged and there pubescent, abruptly narrowed below the
versatile anthers, their ovate cells introrsely longitudinally dehiscent.
Ovary in male flowers reduced, either immersed in a fleshy disk or
eccentric in orifice of hippocrepiform disk, in the female 1-celled
and with a lateral basal fleshy pulviniform avascular appendage,
the style obsolete, the stigma fleshy, rugose; ovules 2, nearly col-
lateral. Drupes compressed, with large fleshy appendages on con-
cave side, putamen with two pairs of equal primary ribs on the
lateral edges and one median ridge on both sides, intermediate
ridges more or less developed on convex side; seed solitary, embryo
minute (after Howard).
Discophora guianensis Miers, Ann. Mag. Nat. Hist. ser. 2.
10: 118. 1852; Contr. Bot. 1: 105. 1851-61; 27. Kummeria brasilien-
sis Mart, ex Engler in Mart. Fl. Bras. 12, pt. 2: 52. 1872. Lasian-
thera amazonica Barbosa Rodrigues, Vellosia 1: 12. 1891. ed. 2,
fide Howard.
Branches, young leaves and panicles except the flowers more or
less sericeous-strigose with silvery or golden trichomes; petioles
1-2.5 cm. long, stout, canaliculate; leaves oblong-lanceolate to ellip-
tic-oblong, acute or rarely rounded at base, 1.5-3 dm. long, 7-13 cm.
wide, acute or acuminate, with apically rounded acumen 2 (-4) cm.
long, coriaceous, revolute-margined, midnerve prominent, strigose
beneath, conspicuously reticulate on both sides, the veins 7-10 pairs;
calyx campanulate, glabrous except for tips of teeth; petals obovate
to elliptic, 2-3 mm. long, glabrous except within inflexed acumen;
stamens 3.5-4 mm. long; basal pulvinus in female flowers 1 mm. high,
surrounding about half of ovary, stigma strongly rugose; rudiment
in male flowers usually completely surrounded by fleshy disk; drupes
2 cm. long, 1 cm. wide, 9 mm. thick, strongly 5-ribbed on the convex
surface. A shrub or tree to 13 meters tall, trunk 1 dm. in diameter.
Illustrated, Rodrigues, I.e. pi. 7; Engler, I.e. pi. 12 (inaccurate
except for habit, Howard). F.M. Neg. 35916 (K. brasiliensis).
Loreto: Balsapuerto, Klug 3017. Mishuyacu near Iquitos, King
452; 478; 942. To British Guiana and Bahia, Brazil.
FLORA OF PERU 225
3. EMMOTUM Desvaux
Reference: Richard A. Howard, Journ. Arnold Arb. 23: 479-
493. 1942.
Shrubs or trees with pubescent branches, alternate coriaceous
entire leaves and axillary fascicled paniculate bracteate inflores-
cences, the perfect 5-parted flowers articulated. Calyx campanulate,
persisting; petals fleshy, rarely glabrate without but the minute
inflexed apex glabrous. Anther cells 2, the fleshy connective bilobed
at base, the glabrous broad and flattened filament attached there
or dorsally near there. Ovary glabrous or hirsute, often with a
fleshy sterile ring of tissue at base, 2-3-celled. Style terminal or
slightly eccentric, glabrous. Drupe rarely with 3 seeds, the coty-
ledons orbicular, the radicle elongate.
Ovary glabrous; mature leaves tawny pubescent beneath.
E. floribundum.
Ovary hirsute; leaves glabrous at maturity E. glabrum.
Emmotum floribundum Howard, Journ. Arnold Arb. 23: 487.
1942.
Subterete branches cinereous, pubescent at tips as the narrowly
sulcate petioles, these 1.5-2 cm. long; leaves ovate or oblong, rounded
at base, acute, 9-12 cm. long, 4.5-5 cm. wide, castaneous, lustrous
and glabrous above, shortly appressed castaneous-hirsute beneath,
the 10 or 11 subprominent lateral nerves arcuate to the free tips;
panicles 3-4-fasciculate, shortly fuscous sericeous; calyx campanulate,
2 mm. broad, 1.3 mm. high, the triangular lobes 0.6 mm. long, slightly
wider; petals lanceolate to ovate, about 6 mm. long, 1.5 mm. wide,
densely ashy-sericeous without, castaneous or reddish lanate within;
stamens to 6 mm. long, incurved, the ovate-oblong anthers 1.2 mm.
long, the filaments dilated toward the base; ovary globose, glabrous,
3-celled, the glabrous eccentric style 4 mm. long. Nearest the
Amazonian E. acuminatum (Benth.) Miers, 486, with shorter petioles,
smaller white flowers. The type was a tree about 15 meters tall
with cream-colored flowers (Klug).
Loreto: Mishuyacu near Iquitos, Klug 1212, type. "Ingaina."
Emmotum glabrum Benth. ex Miers, Ann. Mag. Nat. Hist,
ser. 3. 4:366. 1859; 488.
Branches slender, terete, shortly cinereous pubescent; petioles
7-9 mm. long, with flaring almost winged sparsely crispose or hirsute
margins, canaliculate above; leaves ovate to elliptic, 6-9 cm. long,
226 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
3.5-4.5 cm. wide, rounded at base, narrowly attenuate to mucronate
acumen 1-1.5 cm. long, glabrous and usually dull above, early short
hirsute beneath but soon glabrate, veins almost inconspicuous both
sides, irregularly arcuate, free at tips; panicles slender, slightly
exceeding the petioles or to 1.5 cm. long, sericeous; calyx campanu-
late, 2 mm. wide, 1 mm. high, the rounded or acute lobes about 0.6
mm. long, glabrous except tips; petals ovate-lanceolate, 3-3.6 mm.
long, to 1 mm. wide, glabrous without but for medial row of pilose
trichomes, densely red-brown lanate on mid-nerve within; stamens
to 3.6 mm. long, incurved, anther sacs folded inward, the anther
attached basally to filament; ovary globose, densely silky hirsute
and with differentiated sterile more or less pubescent base; style
eccentric, glabrous, to 2 mm. long, 3-toothed or rounded at tip.
The Peruvian specimen was given an herbarium name by Standley;
it differs from type in inflorescence shorter than petioles, sterile
base of ovary essentially glabrous or with few trichomes, cells
abutting on the narrow ends instead of lying parallel, differences
I do not consider worthy of specific distinction (Howard). Tree
8 meters tall with cream-colored flowers (Klug). Illustrated, Miers,
Contr. Bot. 1. pi. 22 (as to habit only, Howard). F.M. Neg. 23274.
San Martin: Chazuta, 200 meters, Klug 3990. Amazonas, Brazil.
4. PLEURISANTHES Baillon
Reference: Richard A. Howard, Contr. Gray Herb. 142: 41-49.
1942.
Lianas, the terete young stems and branches tendril-like in
appearance, the entire or spinose-denticulate leaves pinnately veined
with prominent reticulate secondary veins. Inflorescence axillary,
supra-axillary or terminal, paniculate or spiciform-racemose, the
rachis commonly flattened, the sessile or pedicellate perfect flowers
nonarticulated and in bracteate glomerulate usually secund clusters.
Calyx with 5 triangular acute lobes, corolla polypetalous, the fleshy
petals strigose or tomentulose without, glabrous within with often
conspicuous midrib. Stamens 5, the free filaments filiform, the
oblong anthers longitudinally dehiscent. Ovary conical, hirsute,
1-celled with two collateral ovules; style minute or developed, the
stigma frequently papillose; fruit unknown.
Pleurisanthes flava Sandw. Kew Bull. 467. 1931; 48.
Younger stems, petioles, these flexuose, to 18 mm. long, leaf-
nerves beneath and axillary racemes, including the flowers without,
FLORA OF PERU 227
more or less tomentose with often yellowish trichomes; leaves ovate
to elliptic-oblong, rounded to subcordate at base, acute or acuminate
but acumen broadly obtuse to abruptly mucronate, 6-14 cm. long,
3-7 cm. wide, membranous, lustrous above, pubescent and paler
beneath, entire, the veins 7-9 pairs; racemes to 5.5 cm. long, the
flowers two-ranked in clusters of 3-5; calyx cupulate, 1.75 mm. wide,
teeth acute; petals elliptic, 2.5 mm. long, the stamens somewhat
shorter; ovary ovoid, densely pilose; style 0.5 mm. long, glabrous
stigma discoid. P. simplicifolia Sleumer, 48, has broadly elliptic
leaves, 8-14 cm. long, 5-9 cm. wide, rounded to obtuse or sub-
acuminate; since both species have been found at Sao Paulo de
Olivenca, they probably occur, in common with many others, within
Peru. Illustrated, Howard, I.e. pi. 2B, opposite p. 17.
Peru (probably). To British Guiana.
5. CITRONELLA D. Don
Reference: Richard A. Howard, Contr. Gray Herb. 142: 60-89.
1942.
Villaresia R. & P. Fl. Peruv. 3: 8. pi. 231. 1802, not Prodr. 35.
1794. Briquetina Macbr. Field Mus. Bot. 11: 26. 1931, fide Howard.
Villaresiopsis Sleumer, Notizbl. Bot. Gart. Berlin 15 : 232. 1940, fide
Howard.
Striate terete branches occasionally scandent, the leaves entire
or spinose dentate. Inflorescence paniculate or thyrsoid, the axes
terete, the perfect or polygamous 5-parted flowers with free apically
inflexed but not clavate petals, glabrous within, the midrib soon
prominent. Ovary subgibbous, the usually solitary cell with a
conspicuous parietal ridge, the 2 ovules pendent from near the apex.
Drupe scarcely fleshy, unappendaged, seed hippocrepiform, the
embryo small. Funicle ridge-like along cell-wall, the 2 ovules sub-
apical on each side at maturity, this funicle forming a vertical
radial partition, only one ovule developing and the seed finally
curved around the partial partition; this character, while found else-
where, is distinctive for this genus within the family. Leaves of
some species have small cavities in vein-axils, often called glands,
which may contain pubescence; corolla aestivation peculiar, the fleshy
petals with protruding midrib overlap at usually membranous
margins and thus have been interpreted as imbricate but actually
a modified valvate condition (these observations taken from Howard,
Journ. Arnold Arb. 21: 471 et seq. 1940, and also I.e. 62-66.
228 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Leaves with large pores in axils of primary veins, entire, glabrous.
C. melliodora.
Leaves without large pores as above but with many small ones in
secondary axils.
Leaves entire C. incarum.
Leaves spinose-dentate C. ilicifolia.
Citronella ilicifolia (Sleumer) Howard, Contr. Gray Herb. 142:
79. 1942. C. peruviana Howard, Journ. Arnold Arb. 21: 474. pi. 4-
1940, fide Howard. Villaresiopsis ilicifolia Sleumer, I.e.
Tree or shrub similar to C. incarum; branchlets fulvous hirsute,
the young leaves sparsely so both sides; petioles 5-6 mm. long;
mature leaves broadly lanceolate or elliptic, 12-18 cm. long, 5-8 cm.
wide, rounded and more or less oblique at base, narrowly acute,
spinose-dentate with teeth 2 mm. long; inflorescences axillary or
extra-axillary, composed of racemose cymes, branchlets 5-10-
flowered, recurving at maturity; petals 5 mm. long, 2 mm. wide;
stamens to 2.8 mm. long; style 0.8 mm. long, stigma oblique; fruit
unknown. As noted by the author, this species differs from C. in-
carum in the shape of leaves and their spinose margins, those of the
younger leaves, however, being only sinuate; since this character
is known to be variable in some other species, the exact status of
C. ilicifolia remains to be established.
Junln: Rio de Comas, 2,400 meters, Weberbauer 6617, type.
Citronella incarum (Macbr.) Howard, Journ. Arnold Arb.
21: 472. 1940; 77. Briquetina incarum Macbr. Field Mus. Bot. 11:
26. 1931. B. mollis Sleumer, Notizbl. Bot. Gart. Berlin 15: 231.
1940, fide Howard.
Large tree, the terete glabrous rambling branches with pale
brown lustrous bark; branchlets and peduncles of the racemosely
disposed short spikes (2 cm. long) glabrate or sparsely puberulent;
petioles stout, 7-15 mm. long; leaves nearly elliptic or broadly ovate-
elliptic, more or less oblique at base, shortly and obtusely acuminate,
mostly about 1.5 dm. long, 6-8 cm. wide, coriaceous, barely lustrous
above, opaque and early hirsute-tomentose or sometimes with a few
short crisp trichomes beneath, entire, undulately and very narrowly
cartilaginous-margined; nerves scarcely obvious above but strongly
prominent beneath with the transverse veins; pores present in axils
of secondary veins but absent from the primary; inflorescences 5-13
FLORA OF PERU 229
cm. long, lax, rarely 2 cm. broad even in fruit, the short (5-10-20
mm. long) branchlets strongly incurved-recurved and densely flow-
ered; flowers sessile, yellowish green, rather fulvous pilose, the
calyx shortly 4-5-lobed, 1.2 mm. high; petals 5, valvate, free, glabrous
but minutely papillose, about 1.25-2 cm. long, suboblong, definitely
narrowed at middle and inflexed, with elevated medial nerve but
unappendaged; stamens 5, little shorter than petals and alternate
with them, the glabrous filaments much dilated, thus cuneate;
anthers erect, cells ovate-oblong, connective not at all enlarged; disk
none; ovary free, 1-celled, ovoid; ovules 2 or apparently only one
commonly developing, pendulous; drupes ellipsoid, ligneous, obtuse,
about 1 cm. in diameter. Howard has shown that this is properly
a part of this group considered as including both old and new world
species, which seems to be the correct interpretation; cf. Howard,
l.c. 21: 476-477.
Huanuco: Muna, about 2,200 meters, 4050, type. Yanano, 2,000
meters, 3748, San Martin: Juanjui, King 3807. Libertad: Valley
of Mishiollo below Ongon, Weberbauer 7056.
Citronella melliodora (Sleumer) Howard, Contr. Gray Herb.
142: 77. 1942. Briquetina melliodora Sleumer, Notizbl. Bot. Gart.
Berlin 15: 230. 1940.
Branches glabrous; petioles 12-17 mm. long; leaves oblong,
rarely ovate-oblong, scarcely inequilateral at the broadly cuneate or
rounded base, acuminate with subacute curved acumen 1.5-2 cm.
long, entire, 18-25 cm. long, 7-19 cm. wide, subcoriaceous, lustrous
both sides, the 4-5 pairs of primary veins with pores in their axils;
inflorescence 6-10 (20) cm. long, rachis laxly pilose, the cymes
obviously peduncled ; flowers fragrant, subsessile, the scarious bracts
ciliate, the pubescent ovate sepals 1 mm. long; petals oblong, 1.5 mm.
long, glabrous, yellow; stamens half as long, with subulate filaments
and ovate-oblong anthers; ovary ovoid, glabrous, the short style
with subcapitate stigma; fruit unknown. Not seen by Howard, who
notes that ex char, it differs from C. incarum only in the possession
of pores in the vein axils, and has a sweet odor. A tree 12-15 meters
tall with trunk diameter of 2-3 dm.
Loreto: Mouth of the Santiago, (Tessmann Wk, type).
6. PORAQUEIBA Aublet
Reference: Richard A. Howard, Contr. Gray Herb. 142: 49-58.
1942.
230 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Trees, the veins of the coriaceous entire leaves weakly if at all
anastomosing near the margin, the paniculate axillary inflorescence
strongly branched from base, the articulated flowers subtended by 3
imbricate pubescent bracts, the buds globose. Calyx lobes 5, fleshy,
lightly imbricate; petal edges incurved, midrib prominent, furrowed
and more or less white silky pubescent. Stamens with fleshy flattened
glabrous filaments, erect anthers with broad connective extended to
inflexed tip, dehiscence lateral. Disk none. Ovary globose, glabrous,
1-celled, the shorter style terminal, capitate, the stigma minute.
Drupes somewhat oblique, fleshy, the woody and smooth endocarp
with a longitudinal ridge around the fruit, the cotyledons foliaceous.
Cultivated in Brazil for the abundant starch and oil in the fruits,
which are edible; the pulp according to Le Cointe contains 12 per
cent of its weight in oil.
Poraqueiba sericea Tul. Ann. Sci. Nat. ser. 3. 11: 172. 1849; 56.
Terete often striate branches sericeous to glabrate as the younger
leaves beneath and inflorescence including the flowers without;
petioles 2-4 cm. long, deeply canaliculate, usually 4-angulate; leaves
broadly elliptic-ovate, rounded at base, 17-24 cm. long, 8-15 cm.
wide, acuminate, the acumen obtuse, rounded or emarginate, 10-22
mm. long, the adult glabrous above, glabrate beneath, coriaceous,
midrib strongly sulcate above, veins 7-9 pairs; inflorescence 5-10
cm. long; calyx 2.3 mm. wide, ovate lobes 1 mm. long and wide;
petals ovate-lanceolate, 3-4 mm. long, densely villous on ridges in-
side, transverse ridge lacking; style stout; filaments 1 mm. wide;
drupes ovate-oblong, slightly compressed, 6-8 cm. long, 4.5-6 cm.
in diameter, in Peru yellowish-green when mature and fresh
(Williams). Attains 30 meters (Krukoff and Ducke), the straight
trunk with a diameter of 6 dm., the crown spreading, bark purplish
or dark brown, the wood compact and used for carpentry, according
to Williams, who has also given the anatomy, Field Mus. Bot. 15:
291. 1936.
Loreto: Iquitos, Killip & Smith 29837; Williams 8078; 8080.
Caballo-Cocha, Williams 2099. Amazonian Brazil. "Umari,"
"umari amarillo," "umari negro" (all Williams); "umari roco,"
"mary negro," "mari preto."
7. LERETIA Vellozo
Reference: Richard A. Howard, Journ. Arnold Arb. 23: 58-60.
1942.
FLORA OF PERU 231
Lianas, or at least the branches scandent, lenticels inconspicuous,
leaves entire, alternate, short-petioled, inflorescences axillary much-
branched peduncled cymes or panicles, the flowers articulated
to short bracteate pedicels. Calyx lobes 5, subacute to rounded.
Petals and stamens 5 (4), the former valvate with inflexed tips, the
latter with arcuate filaments arcuately affixed dorsally, the linear
connective usually prolonged into an obscure tip. Two styles
usually abortive, the functional glabrous. Ovary with columniform
base, pubescent at least within, ovules 2, pendent, disk lacking. Fruit
drupaceous, large, with thin and dry mesocarp, pubescent within;
funicle in a tubular canal in the putamen; cotyledons folded, not
displaced laterally, their margins superimposed. The genus Mappia
Jacq., not yet known from Peru, has ebracteate pedicels, terminal
style, filaments affixed to base of connective, funicle in the mesocarp
(all after Howard).
Leretia cordata Veil. Fl. Flum. 99. 1825; 58. Mappia Poeppig-
iana Baillon, Adansonia 11: 175. 1874, fide Howard. L. nitida
Miers, Ann. Mag. Nat. Hist. ser. 3. 4: 365. 1859, fide Baehni. Ica-
cma(?) Poeppigiana (Baillon) Valeton, Grit. Overz. Olac. 187.
1886. L. Poeppigiana (Baillon) Sleumer, Notizbl. Bot. Gart. Berlin
15: 245. 1940.
Branches red-brown, early appressed short-strigose becoming
glabrate and with darker corky bark; petioles stout, 8-15 mm.
long, commonly twisted, glabrate and corky in age; leaves ovate-
lanceolate to oblong or elliptic, acute or rounded at base, acute and
apiculate to rounded or obtuse, 10-19 (30) cm. long, 4-12 cm. wide,
dark green and often lustrous above where early sparsely white-
pilose, usually whitish or yellowish beneath with malpighiaceous
trichomes or rarely with weak stellate clusters of pilose trichomes,
the stout midrib strigose, laterals 6-8, veins many; inflorescences to
18 cm. long, 10 cm. wide, strigose, the peduncles, pedicels (these
2-4 mm. long), and teeth of patelliform calyces densely so, the
trichomes golden; petals at maturity 4.8-5.5 mm. long, about 2 mm.
wide, densely hirsute without, long-pilose or tomentose within except
for the inflexed tip; ovary densely pubescent within, rudimentary
styles minute; fruit acute both ends or narrowed to base, finally
glabrate, scarcely woody, 3.5-4.5 cm. long, 2-2.5 cm. wide, 1.5-2
cm. thick, the seed to 2.5 cm. long, the raphe extended obliquely to
a circular chalaza near base. Illustrated, Mart. Fl. Bras. 12, pt. 2.
pi. 11 (as M. nitida); Baehni, Candollea 7. pi. 4 (as M. cordata and
232 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
M. Poeppigiana). F.M. Negs. 23278 (L. nitida); 35915 (M. Poep-
pigiana).
San Martin: Tarapoto, Williams 6842; Spruce 44,92. Juanjui,
King 3892. Loreto: Yurimaguas, Poeppig 2329 (type, L. Poep-
pigiana}. Puerto Arturo, Killip & Smith 27869. Mishuyacu near
Iquitos, King 345; 641; 666; 1215. Caballo-Cocha, Williams 2067;
2055; 2084- Iquitos, Williams 3541. Peru-Colombian boundary,
King 1633. Brazil to British Guiana. "Puca-yacu" (Williams).
8. HUMIRIANTHERA Huber
Reference: Richard A. Howard, Journ. Arnold Arb. 23: 73. 1942.
More or less scandent rhizomatous shrubs, the large rhizomes
sometimes tuberiform, the leaves alternate, usually membranous,
entire, the flowers perfect, in terminal or pseudoaxillary cymes
articulated at base of deeply lobed calyx to bracteate pedicels. Petals
ovate to lanceolate, valvate, appressed pubescent without, villous
or crispose pubescent within, inflexed at tip. Stamens free, the
filaments thickened below, inflexed toward attenuate tip, the con-
nective triangular, broadest at base and extending beyond the
globose or oblong anther sacs; style incurved, glabrous; otherwise
much like Leretia.
Howard, Contr. Gray Herb. 142: 58. 1942, has suggested that
Casimirella Hassler, Repert. Sp. Nov. 12: 249. 1913, may prove to
be an earlier name but he notes several technical points which from
character remain vague.
The starchy material of the tubers, these in at least one species
said to attain as much as 4 dm. in diameter, to weigh nearly 100
kilos and to yield 16 kilos of starch, may be used as food when
powdered and repeatedly boiled to remove a poisonous substance;
cf. Howard, who draws upon Ducke, Arch. Jard. Bot. Rio 4: 118.
1925, and Le Cointe, Arvores e Plantes Uteis 457. 1934.
Humirianthera ampla (Miers) Baehni, Candollea 7: 182. 1936;
76. Leretia ampla Miers, Ann. Mag. Nat. Hist. ser. 3. 4: 364. 1859.
Mappia ampla (Miers) Engler in Mart. Fl. Bras. 12, pt. 2: 51. 1872.
H. Duckei Huber, Bull. Soc. Bot. Geneve se>. 2. 6: 184. 1914, fide
Baehni.
Young branches glabrous, lenticels conspicuous; petioles 8-10
mm. long, glabrate as the inflorescence and leaves in age, the pubes-
cence usually simple, the trichomes rarely clustered; leaves broadly
FLORA OF PERU 233
lanceolate to elliptic, subacute to rounded at base, obtuse to acumi-
nate at tip, 8-20 cm. long, 3-10 cm. wide; veins arcuate and strongly
anastomosing; calyx lobes densely hirsute; petals ovate above,
3.5-4.3 mm. long, strigose without, villose to rarely slightly crispose
within except inflexed tip; mature anther sacs at most one-third
length of anther; connective extension 0.6-0.8 mm. long; ovary
hirsute; style glabrous. Illustrated, Valeton, Grit. Overz. Olac.
pi. 5, fig. 31. 1886. F.M. Negs. 23270 (H. Duckei)', 23277.
Baehni was the first to designate this as the type of Huber's
genus, which consists to date of two other species: H. rupestris
Ducke, 75, and H. crispula Howard, 77. The former, of Amazonian
Brazil, has reddish-brown stellate pubescence, veins weakly anas-
tomosing at margins, petals glabrous below; the latter, Colombian,
resembles H. ampla but is yellow-brown stellate, petals with
trichomes within, curly-crispose to base, broader leaves slightly
plicate or cordate and anther sacs one-half to two-thirds length
of anthers; all these are to be expected within Amazonian Peru.
Peru (probably). Amazonian Venezuela and Brazil. "Mandioca
ossu" (Le Cointe), "maria," "apolo" (Ducke).
STAPHYLEACEAE
Reference: Joh. Krause in Pflanzenfamilien ed. 2. 20b: 255-321.
1942.
Shrubs or sometimes trees with various 1-3-foliate or odd-
pinnate (rarely simple) leaves usually with binate stipules and
stipels and small perfect or polygamous flowers in axillary or terminal
panicles. Sepals and petals 5, imbricate, the latter inserted on or
below the hypogynous crenate or lobate disk and accompanied
alternately by the 5 free stamens. Anthers 2-celled, longitudinally
dehiscent. Ovary 2-3-celled and lobed, the styles at least finally
free; ovules few to many in 1-2 series on the ventral suture. Fruit
a membranous inflated capsule opening apically or sometimes inde-
hiscent and fleshy, with 1-few seeds, these truncate at base with
scanty endosperm, straight embryo, the cotyledons plano-convex.
Leaves opposite; sepals free 1. Turpinia.
Leaves alternate; sepals united about one third 2. Huertea.
1. TURPINIA Ventenat
Shrubs with odd pinnate or rarely simple leaves, the leaflets
opposite and small white flowers in terminal panicles. Calyces
234 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
persisting, the sessile petals suborbicular. Filaments complanate.
Ovules few, anatropous. Fruit subglobose, fleshy at least slightly,
3-celled, the seeds compressed.
Turpinia heterophylla (R. & P.) Tul. Ann. Sci. Nat. ser. 3.
6: 363. 1846. Staphylea heterophylla R. & P. Fl. Peruv. 3: 29. pi 253.
1802.
Branches slender, terete, spreading, subvaginate at nodes,
smooth; petioles elongate, the short petiolules canaliculate, the
pinnate leaves with often 5 (3-7) oblong-lanceolate to ovate-oblong
glabrous acutely acuminate serrate leaflets often 8-12 cm. long, 3-5
cm. wide, lustrous, at maturity more or less pubescent and nervose
beneath but green both sides; panicles usually little longer than
broad; bractlets subulate, deciduous; pedicels 3-5 mm. long; calyx
white or purplish, the oval sepals unequal, at least one persisting;
petals white, suboblong, concave, nearly equaled by the filaments;
styles 3; capsules obscurely 3-lobed, seeds two in each cell. Trunk
stout with leafy crown (Ruiz & Pavon). Harms, Notizbl. Bot.
Gart. Berlin 14: 335-336. 1939, called attention to the publication
of Tulasne overlooked by him, by Loesener, Bot. Jahrb. 37: 575.
1906, and even by the careful Briquet, Ann. Cons. Jard. Bot. Geneve
20: 423. 1919! T. megaphylla Tul. I.e., 361, of Ecuador, perhaps to
be expected within Peru, has 1-2 pairs of larger leaflets (often 10-15
cm. long, half as wide) narrowed at base, densely white pubescent
beneath, ample panicles 3 dm. long, flowers subsessile. F.M. Neg.
13380.
Cajamarca: Cutervo, Raimondi (det. Harms). Tambillo, Jelski
414- San Martin: San Roque, WiUiams 7099. Huanuco: Chinchao,
Sawada 54- Mesaporta and Mufia, Ruiz & Pavdn, type. Huancayo:
Rio Mantaro, 2,500 meters, Weberbauer 6564. Puno: Near Puno,
Soukup 27. To Colombia. "Yaua-nullaca," "inquinca."
2. HUERTEAR. & P.
Tall trees with reddish wood, terete branches and alternate leaves,
these with 8-10 pairs of more or less cordate and acute petiolulate
leaflets that are glandular dentate and deciduously stipellate glandu-
lar. Flowers small, many on delicate, apparently not articulated
pedicels, the lax inflorescences rather yellowish pubescent above
the bracts becoming strongly reduced. Sepals and petals 5 or 6,
the outer sepals persisting, scarcely if at all exceeded by the imbricate
FLORA OF PERU 235
pubescent petals, the 5 or 6 stamens alternate to these, somewhat
longer and themselves alternate to the small disk lobes. Ovary
2-celled only at base, style short with 2 spreading stigmas, the
ovules solitary on inner side of each cell. Fruit drupaceous, obovoid,
1-seeded. Named for the Spanish scholar (1579-1649) Hieronymus
de Huerta, who translated Pliny with comments.
Huertea glandulosa R. & P. Fl. Peruv. 3: 4. pi. 227, fig. a.
1802.
Tall thick-stemmed tree; leaves to 3 dm. long; leaflets lustrous
above, somewhat pubescent beneath; petioles and petiolules with
two dark glands at base which however are soon caducous; flowers
2 mm. in diameter; sepals oval, acutish, united at most only one
third. The position of this still imperfectly known tree has long
been open to question and at one time it was confused with Crepido-
spermum Goudotianum (Tul.) Tr. & Planch, of the Burseraceae,
and suggested as possibly belonging to various other botanical
families.
Huanuco: Mufia, Ruiz & Pavon, type. "Cedro macho."
CALLITRICHACEAE. Water Starwort Family l
By Norman C. Fassett
Fragile herbs in shallow water, often maturing emersed, in
drying pools or on muddy banks, or in Peru between Distichia and
other polsters of the wet highlands. Leaves opposite, estipulate,
entire, sometimes forming a rosette at the ends of the branches,
usually pitted, the floating 3-, the submersed 1-nerved. Flowers
monoecious, almost minute, axillary, solitary or 2-3 together, with-
out perianth but often with 2 membranous sometimes inflated bracts.
Stamen 1, the cordate laterally dehiscent anther on filiform filament;
female flowers with 4-celled ovary, ovules 1 in each cell, styles 2,
filiform, papillose. Fruit compressed, emarginate or 4-lobed and
separating finally into 4 1-seeded carpels.
1 Herbert L. Mason, as reported by Jepson, Fl. Calif. 2: 433. 1936, observed
that C. marginata Torr. and at least some other species may have 1-3 male flowers
in an axil, sometimes accompanied by 1 female flower, or the flower, when solitary,
either sex. Dr. Fassett has kindly supplied the key and species descriptions,
including however C. turfosa and C. deflexa at my suggestion. The generic descrip-
tion has been omitted as it would only repeat the family characters. J.F.M.
236 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
CALLITRIGHE L.
Plants usually aquatic, with opposite linear submersed leaves or a
rosette of rotund leaves or both; fruit 1.2-1.4 mm. wide, the
width equaled or exceeded by the height; margins of carpels
wingless C. heteropoda.
Plants terrestrial or subterrestrial with essentially uniform spathu-
late leaves; fruit 0.9 mm. or less wide, the width exceeding the
height; carpels narrowly winged.
Height of fruit 0.6-0.8 mm., 0.1 mm. less than the width; face of
mericarps plump C. turfosa.
Height of fruit 0.3-0.5 (-0.6) mm., 0.2 mm. less than width; face
of mericarp nearly flat C. deflexa.
Callitriche deflexa A. Br. ex Hegelm. Monogr. Gatt. Callitriche
58. pL 3. 1864. C. deflexa A. Br. var. subsessilis Fassett, Rhodora,
53: 150. 1951. C. deflexa A. Br. var. Austini Hegelm. as to South
American plants but excluding North American cotypes.
Fruits 0.5-0.7(-0.8) mm. wide, 0.3-0.5 (-0.6) mm. high on pedicels
0.1-0.3 mm. long; wings of carpels 0.03-0.05 mm. wide, a little
narrower toward the base of the fruits; leaves 2-4 mm. long, 0.8-
2.3 mm. wide, the midrib and often the two lateral veins evident.
A specimen by Weberbauer from near Poto, Puno, determined at
Dahlem as C. marginata Torr., could possibly be this.
Peru (possibly). Mexico to Argentina.
Callitriche heteropoda Engelm. ex Hegelm. Verh. Bot. Ver.
Brandenb. 9: 40. 1867.
Fruit 1.2-1.4 mm. wide, 1.2-1.5 mm. high, the height equaling
or slightly exceeding the width; edges of carpels wingless, rounded
or very obtusely angled with a broad V-shaped commissural groove
between them; fruit sessile or on pedicels up to 15 mm. long on the
same plant; leaves linear and 1-nerved below and broadly ovate
above or all of the ovate-rotund type, or, when plants are stranded
on shore, oblong and of firmer texture. The Pennell specimen seen
has leaves 4-10 mm. long, 2-8 mm. wide. The specimen listed by
Weberbauer, 219, from Poto, Puno, above 4,000 meters between
Distichia polsters, may belong here but compare C. deflexa.
Cuzco: Pools in Distichia moor, 4,200-4,500 meters, La Raya,
Pennell 13493 (det. Fassett). Northwestern Bolivia.
FLORA OF PERU 237
Callitriche turfosa Bert, emend. Hegelm. Monogr. Gatt. Cal-
litriche 59. 1864, in part; Hegelm. Verb. Bot. Ver. Brandenb. 9: 13.
1867. C. turfosa Bert. Amer. Journ. Sci. 19: 308. 1831, nomen nudum.
Fruits 0.7-0.9 mm. wide, 0.6-0.8 mm. high, very narrowly and
evenly winged all around, sessile or on peduncles up to 0.4 mm. long;
mericarps plump, thinner toward the margins; leaves essentially
uniform, 2-4 mm. long, 0.75-1.5 mm. wide, 3-nerved. In damp
soil and shallow water. C. platycarpa Kuetz. of Ecuador has fruits
twice as large with longer wings. Since Fassett has no record of the
following specimen the determination may be my error. Illustrated,
Hegelmaier, I.e. pi. 3, jig. 4 (fruit).
Junin: Carapata above Huacapistana, Killip & Smith 21$86 (det.
Fassett). Ecuador to northern Chile and eastern Argentina.
CORIARIACEAE. Coriaria Family
Shrub (Peru) with opposite branches and leaves (or the former
ternate below) and perfect or subpolygamous small flowers borne in
slender racemes. Sepals 5, imbricate in bud, persisting and spreading,
the shorter hypogynous petals fleshy and carinate within, becoming
thicker and intruded between the fruit carpels, the 10 stamens free
or 5 adherent to petals. Carpels 5-10, adnate in a whorl about the
fleshy conic torus, 1-celled, the styles same number, free, long and
stigmatose all around; ovules solitary, pendulous. Fruit of 5 or
more carpels included in the accrescent petals, compressed and
carinate laterally and dorsally.
CORIARIA L.
A single genus of few species, the Mediterranean C. myrtifolia L.,
like the Peruvian, rich in tannin and also furnishing a black dye.
Coriaria thymifolia Humb. & Bonpl. ex Willd. Sp. PL 4: 819.
1806.
Slender-stemmed, the usually spreading-recurving branches
suggesting fern fronds because of the many small approximate leaves
disposed in one plane; glabrate or lightly puberulent except the
rachis of the racemes densely so; leaves sessile or subsessile, oblongish,
acute, a cm. or 2 long, 5-12 (20) mm. wide, prominently nerved;
flowers 2 mm. long, reddish and green; fruits globose, 3-4 mm. thick,
juicy, dark purple. Often forms hedge-like thickets sometimes 1-3
238 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
meters tall, especially along brooks between 2,000 and 3,200 meters
(Weberbauer). Illustrated, HBK. Nov. Gen. & Sp. 7: pi. 636.
The plant is poisonous to species of llama (Herrera); and with
the racemes the Indians dye their cotton and wool (Ruiz & Pavon).
Cajamarca: Valley of the Llaucan by Hualgayoc, Weberbauer
4043; 261. Cutervo, Raimondi. Huanuco: San Rafael, Soukup
2245. Mito, 1525. Pampayacu, Kanehira 230. Junin: Near
Tarma, Ruiz & Pavdn. Vitoc, Isern 2290. Huacapistana, Weber-
bauer 2008; 246. Uspachaca, 1320. Huancavelica: Near Surcu-
bamba, Stork & Norton 10343. Ayacucho: Aina, Killip & Smith
23119. Apurimac: Raimondi. Huancarama to Cochacaya, West
3761. Marcapata, Vargas 9701; 1354Cuzco: Mollepata, Valley
Apurimac, Herrera 1185. Pillahuata, Pennell 14131; West 7085.
Achirani, Vargas 11112. Ollantaytambo, Cook & Gilbert 607. Valley
Paucartambo, Herrera 2969. Valley Urubamba, Weberbauer, 243.
Puno: Valley Sandia, Weberbauer 554; 237. To Mexico. "Miyo-
miyo," "llama-miyo" (Herrera), "saca-saca" (Raimondi), "mio,"
"mio-mio," "chanche" (Bonpland).
JULIANIACEAE. Juliania Family
For this family, which might be sought here since some regard
it as related to Anacardiaceae, see this work part 2, no. 2: 266-267.
1937. Record has shown that the wood structure suggests that it
may actually be near Burseraceae or here.
ANACARDIACEAE. Cashew Family
Reference: Engler in DC. Monogr. Phan. 4: 171-500. 1883.
Shrubs, sometimes scandent, or trees with alternate (Peru)
estipulate simple 1-3-foliate or (rarely odd-) pinnate leaves, normally
epunctate, and small perfect or polygamous usually regular flowers
in axillary or terminal panicles. Calyx 3-7-parted, more or less
deeply, sometimes completely, rarely spathaceous or irregularly
ruptured; petals ordinarily same number, infrequently wanting or
persisting; stamens commonly twice as many, rarely fewer or more
numerous inserted at base of the usually annular disk. Ovary 1 (2-5)-
celled, rarely with distinct carpels; styles 1-3; ovules solitary,
pendulous or ascending. Fruit superior, infrequently semi-inferior,
sometimes inserted upon the accrescent and fleshy calyx-base and
pedicel (hypocarp), but usually a drupe (seldom dehiscing). Endo-
sperm none or scant. The family is further characterized by the
FLORA OF PERU 239
presence of an oil or resin and a more or less caustic sap that may
occur even in the fruit.
Cashew and pistachio nuts, mangos and mulli or molle trees
as well as species of mayco (Rhus and Mauria) that may irritate the
skin when touched, are among a few of the members of this varied
family. All these are described below except Pistacia (P. vera L.),
which strangely seems to be rarely if at all cultivated.
For convenience the following key applies only to the Peruvian
species; the genera are illustrated, Engler, I.e. pis. 4-15, flowers and
fruits.
Leaves simple.
Leaves sessile or subsessile, nearly linear; female flowers apetalous.
10. Haplorhus.
Leaves petioled, broader; petals present in both sexes.
Carpels solitary; style excentric and filiform or lateral, stigma
simple.
Stamens 8-10, several or all fertile; style excentric; fruit
receptacle enlarged, topped by nut-like drupe.
1. Anacardium.
Stamens 1 or 4-5, 1-2 fertile; style lateral; fruit fleshy.
2. Mangifera.
Carpels 3; style short with 3 stigmas; fertile stamens 10.
Leaves 7 cm. long or longer (one species) ; drupes compressed.
5. Mauria.
Leaves about 4 cm. long or shorter (two species); drupes
globose 6. Schinus.
Leaves pinnate, rarely 1-3-foliate.
Stamens (8) 10, that is, usually twice as many as petals.
Styles and ovary cells (3) 4-5; drupes ovoid or subglobose,
edible.
Leaflets 5-12 pairs; calyx deciduous; stigmas spathulate.
3. Spondias.
Leaflets 2-5 pairs; calyx persisting; stigmas rounded.
4. Tapirira.
Style 1, sometimes trifid; ovary 1-celled; drupes inedible.
Leaflets petiolulate; drupes compressed, endocarp chartaceous.
5. Mauria.
Leaflets sessile; drupes globose, endocarp osseous. 6. Schinus.
240 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Stamens only as many as petals or sepals.
Drupes subglobose; leaflets (Peru) many, petiolulate. .7. Rhus.
Drupes compressed, subalate or alate apically; leaflets, if many,
sessile.
Leaflets 2-4 pairs, petiolulate 8. Loxopterygium.
Leaflets many pairs, sessile 9. Schinopsis.
1. ANACARDIUM L.
Evergreen trees with petioled or sessile entire coriaceous leaves
and small bracteate polygamous flowers borne in terminal panicles.
Calyx divisions as narrow petals imbricate, 5, the latter recurving.
Torus spiciform, occupying the calyx base and adnate to the more
or less unequal connate filaments .of the often only partly fertile
stamens. Ovary free, sessile, 1-celled with slender excentric style,
the stigma punctiform; ovule lateral, ascending on a very short
funicle. Fruit reniform, laterally compressed, umbilicate by a lateral
sinus borne on a pyriform conspicuously enlarged fleshy hypocarp.
The peculiar fruits of the popular Maranon, when mature pendent
in clusters from the medium-sized tree, consist, at least colloquially,
of a lower rather spongy but juicy obovoid portion, reddish-yellow,
sweet-acid and astringent, pleasant fresh, preserved or as a flavor,
and an upper much smaller kidney-shaped part (the actual fruit)
which encloses the well-known cashew nut, edible only after roasting,
since its hull contains a resinous irritant (anacardic acid and cardol,
a caustic oil) that must be avoided with care when eating the fruit-
like base upon which it rests.
Anacardium occidentale L. Sp. PL 383. 1753; 219.
Smooth, glabrous, the branchlets densely leafy toward the tips;
petioles 1-1.5 cm. long; leaves broadly oblong-obovate, rounded,
truncate or slightly emarginate at tip, often 1-1.5 dm. long, 6-8 cm.
wide; panicles a dm. or two long with slightly pilose bracts 5-10 mm.
long; pedicels 2-3 mm. long; flowers yellowish green with roseate
stripes, calyx segments lanceolate, petals 7-8 mm. long, puberulent
without; stamens very unequal, connate only at base; hypocarp to
7 cm. long, 5 cm. thick, fruit 2-2.5 cm. long, 1.5 cm. broad, the
reniform seed 1.5-2 cm. long, 1 cm. broad. Related species or forms
native to the upper Amazonian areas include A. giganteum Hancock
with semiovate calyx segments much shorter than petals and A.
FLORA OF PERU 241
Spruceanum Benth. with longer petioles, filaments connate into a
tube. Illustrated, Chicago Nat. Hist. Mus. Pop. Ser. 26: 47.
San Martin: Tarapoto, Williams 5548. Junin: Rio Perene",
clearing, Kittip & Smith 25118. La Merced, 5479. Loreto: Cul-
tivated, Nanay, Williams 400; 483. Near Iquitos, Williams 1388.
Caballo-Cocha, Williams 2183. Brazil; generally planted and often
naturalized in tropical regions of both hemispheres. "Maranon,"
"cashew," "caju," "casu."
2. MANGIFERAL.
Popular shade and fruit tree with petioled entire deep green but
often reddish-tinged early and in age coriaceous leaves crowded at
the ends of the many branchlets, these early slightly puberulent
toward the tips, and polygamo-dioecious bracteate flowers in much-
branched terminal panicles. Sepals and petals imbricate, 4-5.
Stamens 1 or 4-5, connate below and with pulvinate or stipitiform
disk, 1-2 fertile. Ovary free, sessile, 1-celled, compressed with
lateral or subterminal curved style, the stigma simple; ovule 1,
ascending. Drupe ovoid or subreniform, fleshy, with a mass of
fibers (more or less reduced in cultivated forms) holding the some-
what flattened seed.
The greenish-yellow usually pink-tinged mango, very juicy and
with sweet aromatic flavor, often with a suggestion of turpentine
(this in better cultivated forms largely lacking), is probably the most
appreciated of the tropical fruits. Its cultivation in Peru is not as
frequent as would be expected but Williams found it planted exten-
sively in the northern departments. It has a curious tendency, as
the avocado, of often bearing flowers at one time only on part or
perhaps half of the tree, the contrast being great because the flowers
are usually abundant.
Mangifera indica L. Sp. PL 200. 1753; 198.
Becoming a tall tree, the densely leafy crown spreading, the trunk
sometimes a meter in diameter; leaves usually narrowly oblong-
lanceolate, 1-2 dm. long, 5-8 cm. wide, acute or acuminate both
ends; sepals to 2.5 mm. long, pilose, the narrow apically recurved
petals twice as long; fertile stamens 1-2; fruit often oblong-ovoid
and slightly lop-sided; varying from small to large in cultivated
trees. Illustrated, Chicago Nat. Hist. Mus. Pop. Ser. 26: 4; Engler,
I.e. pi. 4, figs. 10-12.
242 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Ancash: River flat, Yautan, 2567. Junin: Puerto Bermudez,
Kittip & Smith 26471. Lore to: Yurimaguas, Williams 3980; 4519.
Caballo-Cocha, Williams 2354. Iquitos, Klug 1477. Cuzco: Valle
de Santa Ana, Hacienda Huadquina, Hen era 3612. Southern Asia.
"Mango."
3. SPONDIASL.
Branches below the crowded odd-pinnate often ample leaves
roughened by petiolar scars and lenticels, glabrous. Leaflets many
pairs, petiolulate or sessile, more or less oblique at base, acuminate.
Flowers usually pedicellate in pyramidal terminal or lateral panicles,
polygamous, the subimbricate calyx lobes small, the 4-5 spreading
petals valvate in bud. Stamens 9-10 inserted below the cupulate
crenate disk. Ovary sessile, free, 4-5-celled with as many styles
connivent above; ovules solitary, pendulous. Fruit a fleshy drupe
with a large osseous stone. Embryo straight.
The yellow and red mombin ("ciruela agria") are smaller than
mangos, ovoid or plum-like with subacid aromatic thin flesh, fibrous,
especially the yellow species; according to Williams both are planted
commonly in lowlands of Loreto, where persisting in thickets.
Leaflets subsessile or shortly petiolulate, mostly 2-3 (5) cm. long,
rounded to acute S. purpurea.
Leaflets well-petiolulate, often 6-10 cm. long, acuminate. S. Mombin.
Spondias Mombin L. Sp. PI. 371. 1753; 244. S. lutea L. Sp.
PI. ed. 2: 613. 1762.
Becoming a tall tree, the small fragrant white flowers conspicuous
from the abundance of the often ample panicles; leaflets thin, 5-9
pairs, oblong or ovate-oblong, 2.5-4 cm. wide, usually pilose on the
8-10 subparallel spreading lateral nerves; petals 3 mm. long, reflexing,
equaled by the stamens; fruit yellow, ovoid, ordinarily 3-4 cm. long.
Illustrated, Chicago Nat. Hist. Mus. Pop. Ser. 26: 14.
Grown at Surco near Lima, according to Ruiz and Pavon. Only
a few of the many collections by Williams and others are cited.
Loreto: Frequent in thickets, Lower Itaya, Williams 217; 258;
259. Rio Mazan, Jose Schunke 250 (det. Standley). Lower Nanay.
Williams 408. Pebas, Williams 1761. Caballo-Cocha, Williams
2156. La Victoria, Williams 2651, and others. Upper Itaya,
Williams 3401 ; 3446. Lower Huallaga, Williams 4437; 4933. Near
Iquitos, Williams 8034. Rio Acre: Mouth of Rio Macauhan,
FLORA OF PERU 243
Krukoff 5767. To Brazil and Mexico. "Ciruelas agrias" (Ruiz &
Pavon), "tobo de montana," "shungu," "ubo," "uvo," "ushun" (all
Williams), "hubus" (Schunke).
Spondias purpurea L. Sp. PI. ed. 2. 613. 1762; 243.
Often a shrub-tree smaller than S. Mombin, the few branches
stout and the panicles narrow, only 2-4 cm. long and borne mostly
on the larger branches at defoliate nodes, the flowers reddish or
purplish; leaflets 5-12 pairs, variable in shape, mostly subsessile;
fruit commonly red or purple, sometimes yellow, plum-like or egg-
shaped, 3-5 cm. long or larger. Illustrated, Chicago Nat. Hist.
Mus. Pop. Ser. 26: 48; Engler, I.e. pi 6, figs. 19-24.
This species like the other is only in cultivation or naturalized;
Ruiz and Pavon noted it at Macora near Huanuco as S. Mombin,
"its fruit is red and has a bittersweet taste, quite agreeable," but
on account of the color this was probably S. purpurea. Williams
recorded it as fairly common in old clearings in lowlands.
An anomalous specimen by Lorenz and Lescamp from Pucallpa,
Huanuco, referred by Standley to S. Mombin with query, has leaflets
to 5.5 cm. long, 2.5 cm. wide, inflorescence to 1.5 dm. long but this
detached; material broken and inadequate, possibly mixed, but of
interest especially for the native names "tapiriba" and "mangana
decoro."
San Martin: Tarapoto, Spruce 4093. Pongo de Cainarachi,
Klug 2610. Huanuco: Near Huanuco, Ruiz & Pavdn. Loreto:
Lower Itaya, Williams 216. Near Yurimaguas, Williams 4207. To
Mexico and the West Indies. "Ciruela agria" (Ruiz & Pavon),
"ciruela," "ajuela" (Williams).
4. TAPIRIRA Aublet
Shrubs or trees with alternate odd-pinnate leaves and small green-
ish-yellow polygamo-dioecious flowers borne in axillary or terminal
panicles. Calyx segments and oval petals 5, imbricate, the former
persisting, the latter more or less spreading. Stamens 10, inserted
under 5-lobed disk, exserted in male flowers. Ovary ovoid, free,
partly immersed in disk, 1-celled, the short conic styles 4-5 with
simple stigmas; ovule laterally pendent. Drupe obliquely sub-
globose, fleshy, with rugulose-crustaceous stone, the oblong seed
with thin testa. Cotyledons large, piano-compressed, the radicle
superior; embryo curved.
244 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Branchlets gray-puberulent, glabrate or glabrous; leaflets obtusish
or emarginate; petals reflexing T. guianensis.
Branchlets typically reddish-tomentulose; leaflets cuspidate-acute;
petals suberect T. myriantha.
Tapirira guianensis Aublet, PL Guian. 1: 470. pi. 188. 1775;
277.
Branchlets and petioles early rusty appressed puberulent; leaves
large, often at least 1-2 dm. long, typically with 5-7 distant pairs of
variable but mostly about oblong-elliptic leaflets equally narrowed
both ends, 4-15 cm. long, often only about a third as wide, the
obtusish or emarginate acumen at most 5 mm. long, distinctly petiolu-
late, membranous, glabrous or beneath on the little prominent nerves
minutely puberulent; male panicles ample, much-branched, the larger
about equaling the leaves, the dense bracteolate ultimate branches
subspicate, the smaller female mostly in leaf -axils; flowers very many,
subsessile, slightly puberulent or glabrous, the calyx segments
scarcely 1 mm. long and wide, the equally narrow petals 2.5 mm.
long, yellow with pale greenish tinge (Williams), reflexed; filaments
in male flowers filiform, 3 mm. long; ovary pilose or glabrescent;
drupe about 8 mm. long, 6 mm. thick or larger. Williams, Field
Mus. Bot. 15: 288. 1936, has described the tree as 10-25 meters tall,
the straight columnar trunk sometimes buttressed and unbranched
to one-half the height. It is known in Brazil as "pao pombo" and
is said to furnish a wood (described by Williams) of good quality.
It seems probable that at least within Peru only one species is con-
cerned; the collections referred by Gross to T. Marckandii are
glabrous, the broadly elliptic leaflets about half as wide as long.
Loreto: Balsapuerto, King 3008; 3156 (det. Standley). Mishu-
yacu, King 1377 (det. Gross); 588 & 614 (both det. Gross, T.
Marchandii); 266 (det. Harms). Lower Nanay, Williams 652 (det.
Gross). Near Iquitos, Williams 3649 (det. Harms). Yurimaguas,
Williams 3828. Rio Acre: Mouth of Rio Macauhan, Krukoff 5325;
5565; 5684 (this distr. T. myriantha?). Brazil to the Guianas and
Colombia. "Isa-paritsi" (Williams); "joy-ey" (Klug); "jemeco"
(Klug).
Tapirira myriantha Tr. & PI. Ann. Sci. Nat. se>. 5. 14: 295.
1872; 279.
Branchlets and petioles appressed reddish pulverulent; leaves
typically with 2 pairs (but as determined often 3-4 pairs) of obovate-
FLORA OF PERU 245
oblong leaflets unequal at base, cuspidately acute, the lateral spread-
ing nerves arcuate toward margins, rather prominent beneath;
much-branched ample axillary panicles leafless, flowers shortly
pedicellate; calyx pilosulous, the segments ovate; petals narrowly
oblong, concave, contracted and erose-dentate at tip, not at all re-
flexed, 3-4 times longer than calyx, glabrous, nearly as long as
stamens. Differs especially from T. guianensis in the acute leaflets
and the more erect petals erose-dentate at tip (Engler) but in Peru
at least the characters seem intangible. Williams specimen from a
tree about 20 meters tall, trunk nearly 4 dm. in diameter, exuding
when incised some oily or resinous fluid. The following material
could apparently be included in T. guianensis. Determinations by
Standley except as indicated.
San Martin: Pongo de Cainarachi, Klug 2720. Zepelacio, King
8294. Huanuco: Cuchero, Poeppig 1333 (det. Engler). South of
Monzon, 1,600 meters, Weberbauer, 256. Junin: Chanchamayo,
Schunke 414- Loreto: Mishuyacu, Klug 592 (det. Gross). Florida,
Klug 2137; 2346. Yurimaguas, Mexia 6089. Lower Huallaga,
Williams 4712? Colombia. "Itil" (Mexia).
5. MAURIAKunth
Trees with odd-pinnate leaves or sometimes in part or all of them
simple, the leaflets petiolulate, the shortly pedicelled hermaphrodite
or polygamo-dioecious 5-merous flowers in terminal and axillary
usually very composite panicles. Calyx short-cupulate with short
broad teeth. Petal aestivation valvate. Stamens 10 in hermaphro-
dite or male flowers, the filaments subulate, anthers dorsifixed.
Ovary attenuate into short or obsolete style with broadly 3-lobed
stigmas. Drupe compressed, endocarp chartaceous; seed laterally
affixed.
Some species (like Rhus striata} are said to cause skin irritation,
often serious, merely from contact. The species as defined are
much alike and probably actually fewer.
Leaves all simple M. simplicifolia.
Leaves, at least some of them, pinnate.
Leaves obscurely reticulate above, densely and minutely beneath.
M. aurantiodora.
Leaves obviously reticulate and more laxly both sides.
Pubescence on the leaves beneath conspicuous, soft.
M. Killipii, M. sericea.
246 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Pubescence on the leaves beneath lacking or obscure.
Leaflets entire or nearly entire.
Leaflets 1-2 pairs, coriaceous, subrigid.
Inflorescence glabrate M. heterophylla.
Inflorescence densely pubescent M. trichothyrsa.
Leaflets 2-4 pairs, rather thin or flexible.
Leaflets typically smooth M. biringo.
Leaflets puberulent or sometimes with tufts of hairs in
the nerve axils M. suaveolens.
Leaflets, at least some, obviously denticulate.
Some of the leaflets entire M. subserrata.
All of the leaflets denticulate M. denticulata.
Mauria aurantiodora [Ruiz] Engl. in DC. Monogr. Phan. 4.
326. 1883. Sorindeia aurantiodora [Ruiz] Engl. in Mart. Fl. Bras.
12, pt. 2: 391. 1876.
Shrub with fulvous puberulent branchlets and panicles, these
terminal, few-branched, scarcely 1 dm. long, the short branchlets
1-2-flowered with rounded bractlets not more than 1 mm. long and
fruiting pedicels at most twice as long; petioles 4-7 cm. long; leaflets
petiolulate, ovate, obtuse both ends, unequal, the lateral 6-8 cm.
long, 4-5 cm. wide, the terminal larger, all with reticulate veins
faint even beneath, the lateral rather prominent; calyx lobes rounded,
1 mm. broad; drupes ovoid, little oblique, glabrous, 6-8 cm. long,
4-5 cm. thick. Ruiz' name was in herb, under Schinus. F.M. Neg.
13163.
This is one of several species that, as with Rhus striata, the Indians
believe cause an irritating skin eruption if one sleeps in their shade;
according to Ruiz and Pavon this could be cured in 8-10 days by
application of very hot bunches of Valeriana, "albergilla." See
remarks under Rhus striata.
Huanuco: Near Muria, Ruiz & Pavdn, type. Cueva Grande
near Pozuzo, 4810. Rio Pozuzo, Ruiz & Pavdn; Weberbauer 6745;
6750. "Mayco."
Mauria biringo Tul. Ann. Sci. Nat. se"r. 3. 6: 365. 1846; 328.
Densely leafy flowering branchlets erect, the subterete petioles
1.5-2 dm. long or longer, the leaves with 2-3 pairs of thin green
oblong leaflets shortly and obtusely acuminate, 8-12 cm. long, 4-6
FLORA OF PERU 247
cm. wide, the reticulate veins prominent both sides, paler beneath,
the petiolules 4-5 mm. long; panicles puberulent-pilose, to about
15 cm. long with many-flowered angulate branchlets, the bractlets
acute; calyx minutely puberulent with subtriangular ciliolate teeth;
petals ovate, acute; disk rather thick; stamens in hermaphrodite
flowers half as long as petals; ovary glabrous, contracted into short
style which persists on the obliquely ellipsoid drupe, this about 8 mm.
long. The var. Ruizii Engler (the typical form) has oblong-elliptic
leaves, the lateral nerves ascending, while var. Weberbaueri Loes.
has the leaflets pilose in the nerve axils. This may prove to be a
part of M. suaveolens Poepp. & Endl. and M. heterophylla HBK.,
at least almost certainly of the former. F.M. Negs. 13164; 13165
(var.).
Cajamarca: Tambillo, Jelski 295. Junin: Casapi, Mathews 2104.
Near La Merced, in open wood, Weberbauer 1935 (type, var. Weber-
baueri); 282. Without locality, Ruiz & Pavdn (type, var. Ruizii).
To Bolivia and Colombia. "Birringo" (Colombia).
Mauria denticulata Macbr. Field Mus. Bot. 8: 122. 1930.
Branches glabrous; petioles of the rarely unifoliate leaves 2-3
mm. long, the leaflets especially toward the tips irregularly mucro-
nate-dentate, ovate-lanceolate, subacute at base where somewhat
oblique, gradually and acutely acuminate, about 10 cm. long, 3.5 cm.
wide, subcoriaceous, glabrous above or obscurely pubescent on the
midnerve, densely fulvous hirsute in the nerve axils beneath, other-
wise glabrous, the reticulation prominent; panicles subsessile, lightly
hirsutulous, to 1.5 dm. long, the pedicels 2 mm. long; stamens 10,
about 1 mm. long, the free filaments and anthers equal and between
the lobes of the subannulate disk. A 3-meter shrub. One or two
other species have obscurely or remotely serrulate leaflets but they
are essentially entire, not sharply denticulate and there appear to
be none with such dense tufts of pubescence in the nerve-axils; of
course the range of variation of these apparent differences is unknown
and therefore may some day be shown to be without significance.
Ayacucho: Choimacota Valley, Weberbauer 7546, type.
Mauria heterophylla HBK. Nov. Gen. & Sp. 7: 13. pi. 606.
1824; 327.
Resembles in general M. simplicifolia but some of the leaves
usually with 1-2 pairs of somewhat less coriaceous ovate, ovate-
oblong or elliptic leaflets 8-10 cm. long, 3 cm. wide; panicles some-
248 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
times larger; upper bracts acute; petals 3.5 mm. long, oblong, acute;
filaments 2 mm. long, anthers ovate. The var. puberula (Tul.)
Engl. has leaves pilose in axils of nerves beneath, these obscure above,
while var. contracta Loes. has the denser inflorescence scarcely half
as long as leaves, flowers a little larger. This may prove to be the
earlier name for several forms as M. aurantiodora, M. biringo, M.
suaveolens.
Cajamarca(?). Near Celendin, 3,000 meters, Weberbauer 4247
(type, var. contracta). Piura: Toward Ayavaca, Bonpland, type.
Amazonas: Near Chachapoyas, (Mathews 1458). Huanuco: Muna,
Vitoc and Chinchao, 3967 (det. Johnston); Ruiz & Pavdn. Puno:
Valley of Sandia, in rocks, 2,300 meters, Weberbauer 564 (var.
puberula, det. Loesener) ; 237. To Colombia and Venezuela. "Caspi"
(Colombia).
Mauria Killipii Barkley, Bull. Torrey Club 74: 77. 1947.
Branches striate, conspicuously lenticellate, soon glabrate but
tips as stout petioles, these about 2 cm. long, ferruginous pilose;
leaves pinnately 3-foliate; petiolules pilose, the lateral 2-4 mm. long,
terminal, 6-12 mm. long; leaflets ovate or broadly lanceolate, acute
or subacute, broadly cuneate at base, entire, membranous, glabrous
above, ferruginous tomentose on nerves beneath, prominently
reticulate both sides; panicles axillary and terminal, equaling the
leaves; peduncles sparsely pilose as the deltoid ciliate subacute
bracts, these 1-1.5 mm. long, subpersisting; calyx about 1.5 mm.
wide, with broad pilose lobes 0.3 mm. long; pe&ls ovate, glabrous,
1.3 mm. long; stamens 0.45 mm. long, the rotund yellow anthers
0.3 mm. wide. Type from a 3-4 meter tree with few branches. In
character of inflorescence resembles M. ovatifolia Turcz. but ap-
parently is most closely allied to M. sericea Loes. with smaller inflores-
cence and 5-7 leaflets (Barkley); maybe these differences could be
expected within a reasonable range of variation. However, the
petals of Loesener's species, ex char, are 3 mm. long. Illustrated,
Barkley, I.e., opposite p. 78 (photo of type).
Junin: Huacapistana, Killip & Smith 24182, type.
Mauria sericea Loes. Bot. Jahrb. 37: 571. 1906.
Branchlets striate-sulcate, subfuscous pubescent, about 5 mm.
thick; petioles and petiolules densely pubescent or puberulent as
rachis, the former 3-5 cm. long, petiolules 2.5-5 mm. long or that of
terminal leaflet to 22 mm. long; leaflets imparipinnate, 2 or mostly
FLORA OP PERU 249
3 pairs, subchartaceous, entire or undulate, ovate-oblong to sub-
elliptic, the lateral more or less obliquely cuneate to base, acute or
shortly acuminate, 8-13 cm. long, 2.5-5.5 cm. wide, all puberulent
above, on the face sparsely, on the (10) 12-16 lateral nerves rather
densely, especially beneath where also slightly hirtellous in the upper
nerve axils, the reticulation beneath very dense and fine, the nerves
not prominent; panicles few, fasciculate in the axils or branched
from base into one composite densely hirtellous or pubescent inflores-
cence, 7 or 8 cm. long, branches spreading, ultimate pedicels about
1 mm. long, primary carinate deltoid bracts to 3 mm. long; calyx
about 3 mm. wide, lobes acuminate; petals narrowly ovate-deltoid,
acutish, puberulent without, about 3 mm. long, half as wide; stamens
all little shorter than petals; stigma obsoletely 3-lobed. Seems to
be much like M. ferruginea Tul. of Colombia, known to me only
from description, the leaflets with very prominent nerves (Loesener).
Fruits of my specimen immature, 1 cm. long, 7 mm. wide, lustrous,
and with a rather pungent aroma.
Junin: Huacapistana to Palca, in shrubs, 1,700 meters, Weber-
bauer 1981, type. Huanuco: Muna, 3937 (det. Johnston, ex char.).
Mauria simplicifolia HBK. Nov. Gen. & Sp. 7: 13. pi 605.
1824; 326.
Glabrous shrub or small tree without any compound leaves;
branchlets 3-4 mm. thick; petioles subterete, 6-7 mm. long; leaves
oblong, obtuse both ends, 7-9 cm. long, 3-4 cm. wide, green above,
the veins and lateral nerves prominent beneath; panicles terminal,
exceeding the leaves, to 1 dm. long, laxly many-flowered, the angulate
branchlets 1-3 cm. long, the pedicels longer than the globose buds,
puberulent as the ovate obtuse 1-2 mm. long bracts; flowers 5-6-
merous with short broad apiculate calyx lobes, ovate acute petals
3 mm. long, more than 1 mm. wide; stamens more than half as long,
anthers oblong; disk 10-crenate; ovary with 3-lobed stigma, in the
male flower elongate into a thick style; drupe oblong-ovoid, to 9 mm.
long, 5-6 mm. broad. Observed to 10 meters high.
Cajamarca: Cutervo, Jelski 296. Huanuco: Muna, Ruiz &
Pavdn. Chaglla, 3,000 meters, Weberbauer 6687; 6692. South and
southwest of Panao, 2211; 3582. Colombia.
Mauria suaveolens Poepp. in Poepp. & Endl. Nov. Gen. & Sp.
3: 77. 1845; 329.
Branchlets minutely puberulent as the leaf nerves (unless in age)
and petioles, these terete, 1.5-2 dm. long, petiolules 2-4 mm. long;
250 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
leaflets 2-3 pairs, membranous, ovate or ovate-oblong, shortly
acuminate, obtusish, 7-10 cm. long, 3-5 cm. wide, fulvous pilose in
the axils of the nerves beneath, these with the reticulate veins promi-
nent; panicles ample, with many-flowered spreading branches, the
lower 1-1.5 dm. long; bracts and bractlets acute, the latter 2-3 mm.
long or longer than the pedicels; calyx puberulent, cupulate, the short
teeth acute, 1 mm. wide; petals white, ovate, 2.5 mm. long, puberu-
lent, the stamens in female flowers half as long; ovary ovoid, attenuate
into short style with 3-lobed stigma; ovule pendulous below the
apex of the cell; drupe obliquely ovoid or subglobular, at least 8 mm.
long, 5 mm. broad. Williams noted it occasionally as tall as 20
meters, the erect columnar trunk 4 dm. or more in diameter and
found it limited to the upland. Perhaps a variant of M. heterophylla
HBK.
San Martin: Tarapoto, Spruce 4268. Lamas, Williams 6408.
San Roque, Williams 7098; 7227. Chazuta, King 4016 (det. Stand-
ley). Zepelacio, Klug 3473. Junin: Near La Merced, Weberbauer,
282. Huanuco: Cuchero, Poeppig 1390, 1391, type. Banks of Rio
Ysabel, Churubamba, 1,100 meters, Mexia 8161 (det. Standley,
M. heterophylla). Valley of the Monzon, Weberbauer, 284. Co-
lombia; Dutch Guiana? "Ingaina blanca," "itil," "itil bianco,"
"yurac ingaina" (all Williams).
Mauria subserrata Loes. Bot. Jahrb. 37: 572. 1906.
Similar to M. sericea; branchlet tips merely pulverulent as the
petioles, these (1.6) 2-3 cm. long; leaves sometimes 3-foliate, coria-
ceous, glabrous or subglabrous; leaflets entire or more or less serru-
late, the lateral obtuse or subrotund at base, the others acute or
subacute, all gradually narrowed to acute or obtuse acumen, (4.5)
6.5-10 cm. long, 2-4 cm. wide, the 9-14 lateral nerves and very dense
reticulation only prominent beneath, the latter less so; panicles
puberulent, 4-5 cm. long or in fruit to 15 cm. long, pedicels 1-2 mm.
long, bracts 1-1.5 mm. long; calyx about 2 mm. wide, the lobes
subacute; petals sparsely puberulent, scarcely 2 mm. long; staminodia
staminiform; drupes 7 mm. long with persisting calyx at base and
minutely capitate thickly 3-lobed stigma at tip. On account of the
acuminate and acute leaflets seems to be allied to M. sericea
(Loesener). F.M. Neg. 13170.
Cuzco: Valle del Urubamba, among shrubs, mostly evergreen,
2,000 meters, Intuhuatani to Torontoi, Weberbauer 5070 and 5077,
type.
FLORA OF PERU 251
Mauria trichothyrsa Loes. Bot. Jahrb. 37: 571. 1906.
Sulcate branchlets at tip, petioles, these 2- nearly 3 cm. long,
rachises and leaf nerves beneath densely puberulent; leaves tri-
foliate, rarely imparipinnate, coriaceous, the ovate-lanceolate or
oblong leaflets to subobovate, sessile or subsessile or terminal leaflet
with petiolule to 7 mm. long, all cuneate to subobtuse at base, obtuse
or rounded and shortly apiculate at tip, entire or undulate, usually
3.5-8 cm. long, rarely to 10 cm. long, 1.5-3.5 cm. wide, glabrous
above, the 11-15 lateral nerves rather prominent beneath with the
dense reticulation; panicles axillary, densely pubescent, 2.5-4.5 cm.
long, about 1.5 cm. wide, flowers hermaphrodite, closely glomerulate,
the deltoid bracts 1 mm. long; calyx 2.5-3 mm. wide; petals lanceo-
late-linguliform, about 3 mm. long, 1.5 mm. wide, scarcely acute;
stigma lobes bilobed. Seems to be near M. ferruginea Tul., which
has leaves puberulent above, densely ferruginous tomentose beneath
and much longer petioles (Loesener). F.M. Neg. 13172.
Cajamarca: Near Lajas, 2,500 meters, among shrubs, Weberbauer
4218, type.
6. SCHINUSL.
Reference: Fred Barkley, Brittonia 5: 160-198. 1944.
Trees or shrubby, with alternate simple or odd-pinnate (rarely
paripinnate) leaves with sessile leaflets, these often somewhat
crenate-dentate, especially toward the tips. Flowers polygamo-
dioecious, bracteate, in axillary or terminal panicles or pseudo-
racemes, the 5 calyx segments and oblong petals imbricate, the former
rounded, the latter with the 10 stamens inserted at base of thick
disk, finally reflexed; filaments subulate, anthers oval. Ovary free,
sessile, 1-celled, the ovule suspended from near the apex. Style
terminal, trifid or simple, with 3 minute capitellate stigmas or
simple and 3-lobate. Drupes globose, small, the putamen coriaceous
or osseous, usually oily.
The mulli, or more usually molle, tree is Schinus molle L., notable
as a native plant that not only is distributed in all the drier and
warm climatic areas to 3,000 meters in the south (about 2,000 meters
in the north; Weberbauer) but is the popular planted tree for shade
and ornament. A small example in a highly characteristic setting
may be seen in an excellent photograph by Weberbauer, opposite
p. 297.
Leaves simple, rarely more than 3 cm. long, usually much shorter.
Inflorescences 2-3 cm. long; pubescence sparse S. polygamus.
252 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Inflorescences 1-10 mm. long; pubescence at least on younger
parts dense S. microphyllus.
Leaves at least in part, more or less divided or pinnate, rarely simple,
always 4 cm. long or longer.
Leaves irregularly divided, if at all, 4-6 cm. long; branchlets erect
or spreading S. Pearcei.
Leaves multipinnate, ample; branchlets lax or pendent. . .S. molle.
Schinus microphyllus Johnst. Journ. Arnold Arb. 19: 258. 1938;
178. S. andinus (Engl.) Johnst. Journ. Arnold Arb. 19: 259. 1938.
S. dependens Ort. var. andinus Engl. in DC. Monogr. Phan. 4: 341.
1883.
Slender terminally spinose branchlets early softly puberulent
pilose as the petioles, these 1-3 mm. long, and the 7-10 mm. long,
few-flowered inflorescences; leaves obovate, cuneate at base, obtuse,
entire or slightly serrate, 4-18 (26) mm. long, 3-10 mm. wide,
typically firm-membranous; bracts broadly deltoid, puberulent and
ciliate; pedicels about 3 mm. long, sparsely pilose as the lanceolate-
deltoid truncate to subacute sepals; petals broadly ovate, emarginate
to truncate. Could be included in S. polygamus but apparently an
endemic of more northern distribution. S. andinus, however, seems
to be merely a less pubescent variant with somewhat more coriaceous
leaves, at least in age, and reduced inflorescences, these 1-3 or some-
times to 6 mm. long and may become S. microphyllus Johnst. var.
andinus (Engl.) Macbr., comb. nov. Type a 2-meter shrub.
Ayacucho: Tambo, Prov. de la Mar, 3,100 meters, Weberbauer
5552 (var.). Apurimac: Argama to Andahuaylas, 3,800 meters,
West 3747, type; at 2,500 meters, Weberbauer 5839. Pincos, Prov.
Andahuaylas, 2,700 meters, Stork & Horton 10679 (det. Barkley,
S. andinus, but intermediate). Cuzco: Chaspyoc, Huarocando
River, Edmund Heller 2173 (var.). Ollantaytambo, Cook & Gilbert
427 (var.). Tarapata, Vargas 11052 (var.). Urubamba Valley,
Herrera 2104 (var.); Weberbauer, 175. Bolivia. "Checche" (Cook
& Gilbert); "muluaca" (?, West).
Schinus molle L. Sp. PI. 388. 1753; 184.
Abundantly but loosely leafy and gracefully branched tall shrub
or tree, the slender upper branches and branchlets more or less
pendent and part of the year colorful with large panicles (commonly
8-15 cm. long) of small but numerous yellowish-white flowers or in
FLORA OF PERU 253
many specimens with abundantly developed rose-red berry-like fruits
that may long persist among the feathery light green foliage; leaves
usually with an unequal number of pairs (often 7-13) of linear-
lanceolate subentire or serrate acute or subobtuse leaflets ordinarily
several cm. long, the rachis margined; flowers 5-merous with oblong
petals; styles free, fruits inedible, globose, about 5 mm. in diameter.
The var. Rusbyi Barkley, I.e. 186, is more puberulent, leaflets 34-38,
the terminal lacking and panicles 2-6 cm. long. Illustrated, Bot.
Mag. pi. 3339.
In cultivation becomes a large picturesque tree with short but
huge trunk widely over-topped by the spreading rounded crown, the
loosely borne divided leaves resulting in lovely light effects, especially
with a breeze, in the sun. In some Spanish-speaking lands, as in
Mexico, it is called "Peru," after the country of its origin, or "pi-
miento (del Peru)," since when rubbed between the fingers the fruits
suggest black pepper because of a volatile oil, and thus the common
English name is "pepper tree." But in Peru, as noted by Weber-
bauer, the Quechua name "mulli" is commonly modified to "molle."
According to Herrera, in Calca the leaves are used for an "abono
catalitico" called R'ockescca in which are sunk the seeds of maize.
Ruiz and Pavon, noting that the Incas are said to have called it the
"tree of life," listed a number of uses, many medicinal, including a
"chicha" prepared by fermentation of the fruits with water, "sweet
and agreeable to them and excellent against dropsy" and the use of
the white fragrant resin as plasters for fractures and ulcers. But
they warn that the seeds cannot be used for pepper without "great
harm to health." The ashes, however, of the slow-burning wood
give "an excellent fixed alkali for the purification of sugar and for
dyes" while the trunk furnishes "excellent" lumber.
Yet in its favored natural environment among shrubs of brooks
and rivers, it rarely attains timber size or, quite possibly, it is
destroyed before maturity.
Cajamarca: Near Huaraz, 2,200 meters, Weberbauer, 172; 173.
Puccha Valley, Weberbauer, 174. Below San Miguel, Weberbauer,
188. Below Santa Cruz, Weberbauer, 189. Lima: Hacienda de
San Jadeo and at Huacho (fide Engler). Rio de Supe, Weberbauer,
162; 166. Native in dried-out gulches, 243. Huanuco: Near
Huanuco, Ruiz & Pavdn; Kanehira 60. Junin: Near Tarma, Killip
& Smith 21775; Weberbauer, 111. Valley of the Utcubamba, Weber-
bauer, 191. Arequipa: Yura, Schmidt. At 2,500 meters, (Williams
2555 in part, type, var. Rusbyi). Tiabaya, rocky river bank, Pennell
254 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
13083. Quequena, Eyerdam & Beetle 22181; 22178. Cuzco: Uru-
bamba, Weberbauer, 175; 245. Quebrado de Oropeza, Herrera 922.
Near Ollantaytambo, Weberbauer, 243. Lower valley slopes and
along river, Calca, West 8076. Tacna: Near Tacna, Weddell.
Bolivia; Chile; Ecuador and cultivated or perhaps native over a
much wider area. "Molle," "mulli."
Schinus Pearcei Engl. Bot. Jahrb. 1: 423. 1881; 184.
Glabrous shrub or small tree with (1) 2-5 pairs, sometimes reduced
to 1 leaflet, of very unequal subcoriaceous oblong obtuse but apicu-
late leaflets, the upper 2 or 1 and connate with the terminal, this
2-3 cm. long, the lateral 1.5-2 cm. long, 5-6 mm. wide, the leaves
4-6 cm. long, the petioles winged; panicles or racemes axillary,
4-6 cm. long, minutely puberulent as the deltoid bractlets, the
pedicels 2-3 mm. long; calyx lobes scarcely 0.5 mm. long, obtuse
as the longer ovate petals; anthers ovate; fruits lavender-red, about
5 mm. in diameter (Barkley). To 6 meters tall, associated with
S. molle to which it has great resemblance but quite distinct in habit,
the branches erect (West). F.M. Neg. 35972.
Apurimac: At 2,900 meters, Weberbauer 5853. Cuzco: Near
Calca, 3,000 meters, West 8075; Herrera 2080; 2089; Weberbauer,
175. Rio Chaupimayo, Soukup 587. Ollantaytambo, Cook &
Gilbert 773. Puno: Gay, type. Locality unknown, Orubomba,
(Pearce in herb. Kew). Bolivia. "Molle" or "mulli" (West),
"orcco-mulli" (Cook & Gilbert), "china-mulli."
Schinus polygamus (Cav.) Cabrera in Frenguelli, Inst. Mus.
Univ. Nac. La Plata Obra Cinquent. 2: 269. 1937; 174. Amyris
polygama Cav. Icon. 3: 20. pi. 239. 1794. S. dependens Ortega, Hort.
Matr. Dec. 8: 102. 1798.
Shrub or small tree often gnarled or in any case the very slender
glabrous or puberulent branchlets spinose; petioles only 1-3 mm.
long, pilose, merging into the cuneate base of the oblanceolate to
oblong or obovate obtuse or acute glabrous or rarely sparsely pilose
subcoriaceous obscurely nerved leaves, these entire or more or less
crenate-dentate, ordinarily 1.5-3.5 cm. long, 3-7 mm. wide; flowering
branchlets little longer than foliage, puberulent or glabrescent,
pseudo-racemose (2-3 cm. long), 2-3 flowers in axils of ovate minutely
ciliolate bracts 1 mm. long, the almost filiform pedicels 6-8 mm.
long, much longer than the globose buds; obovate-oblong petals
about 3 mm. long, half as wide, nearly three times as long as the
FLORA OF PERU 255
ovate ciliolate calyx lobes and little shorter than the filiform stamens
(male flowers); styles connate; drupes globose, about 4.5 mm. in
diameter, with thin-chartaceous lilac lustrous exocarp, the mesocarp
very resinous. A number of variants have been designated, often
as distinct species, to take care of differences largely or entirely
due, probably as Engler suggests, to habitat; the Peruvian material
consists only of scraps and is probably mislabeled as to origin or
collectors or both. So, in a strict sense it is highly doubtful if the
species occurs in Peru. Illustrated, Engler, I.e. pi. 12, fig. 21.
Peru(?): Without data, Dombey; Pavdn; Ruiz. Chile; Bolivia.
7. RHUSL.
Reference: Barkley, Ann. Mo. Bot. Card. 24: 265-498. pis. 10-
26. 1937.
Shrub with odd-pinnate leaves (Peru) and small polygamous
bracteate flowers borne in axillary or terminal, sometimes large,
panicles. Calyx segments and petals 4-6, imbricate (in bud),
the former persisting, the latter longer, spreading. Stamens 4-6 or
10, inserted at base of annular disk, the filaments subulate. Ovary
sessile, styles 3, sometimes very short, free or connate at base, the
stigma obtuse or capitate; ovule pendent from a basal funicle.
Drupe slightly if at all fleshy, sometimes compressed, the seed ovoid
or reniform.
Genus notable for the beneficence and infamy of its members;
there is, for example, R. vernicifera DC. and R. succedanea L., both
of eastern Asia, which supply lac from which lacquer, perhaps the
most durable varnish, is made; and there is R. striata R. & P. of
Peru, among others, counterpart of the poison sumac, poison oak
or ivy of North America in its poisonous attributes as noted below
but similar and related to the Asian species, especially to R. suc-
cedanea.
Rhus striata R. & P. Fl. Peruv. 3: 29. pi 252. 1802; 436. R.
juglandifolia Willd. ex Roem. & Schult. Syst. 6: 649. 1820. Toxico-
dendron striatum (R. & P.) Kuntze, Rev. Gen. 1: 153. 1891. Rhus
samo Tul. Ann. Sci. Nat. ser. 3. 6: 367. 1846?
Tall shrub or becoming a slender trunked densely branched
shapely tree 10-15 meters tall, usually nearly glabrous except for
the puberulent branchlet tips and the lateral panicles that are shorter
than the large leaves, these with 5-6 pairs of oblong abruptly acute
256 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
to acuminate membranous entire slender-petiolulate leaflets, often
6-12 cm. long, 4-5 cm. wide, very oblique at base, a little pilose on
the nerves and midnerve beneath, the veins obscure; sepals rounded,
1 mm. wide, glabrous; petals white, 2.5 mm. long; drupes whitish,
about 8 mm. in diameter, the seed 5 mm. long, 7 mm. broad.
R. juglandifolia var. samo (Tul.) Engl., branchlets as leaves beneath
ferruginous pubescent, the fruit ex icone a little larger, seems as to
Peru to be the type. F.M. Neg. 13190 (R. juglandifolia Willd.).
There is a well-grounded belief among the country people that
one taking even a siesta in the shade of this small tree will suffer
the often agonizing torment of the skin poisoning "mayco" to which
many are susceptible; actually, as shown by McNair (at least for
northern species) and others, contact with the sap is necessary but
may occur indirectly from touching clothing already brushed against
the tender foliage or flowers or from fallen parts of the plant. Ruiz
and Pavon in their Journal, English ed., Field Mus. Bot. 21: 80
and 95. 1940, recorded their own experience and the native supersti-
tion which I found persisting a century and a quarter later. A
modern and effective preventative and treatment for relief according
to James B. McNair, Am. Biol. Teacher, 2: 100-101. 1940, or Field
Mus. Leaflet 12: 10-12. 1926, both from the same author's "Rhus
dermatitis," Univ. of Chicago Press, Illinois, is application of 5 per
cent iron chloride in a half and half mixture of alcohol and water,
applied freely. In the United States immunity for some time is
often acquired from poisoning (by similar species) by the use of a
diluted and purified rhus toxin (.0130 gm. in 13 cc. ethyl alcohol)
as prepared by chemical laboratories.
Amazonas: Chachapoyas, Mathews. San Martin: Near Moyo-
bamba, Weberbauer, 291. Huanuco: Chinchao to Macora, Ruiz &
Pavon. Yanano, 3787. Junin: Chanchamayo Valley, Weberbauer.
Below Huacapistana, Sandeman 4571. Cuzco: Valle de Santa Ana,
Hacienda Idma, Weberbauer, 281. Valle de Santa Ana, 1,000
meters, Herrera 3483. Rio Apurimac, 1,100 meters, Weberbauer
5889. Rio Chaupinmayo, Soukup 805. To southern Mexico.
"Mayco," "incati" (Cuzco), "itil" (San Martin).
8. LOXOPTERYGIUM Hook. f.
Trees with 2-4 pairs of large odd-pinnate leaves, the opposite
petiolulate leaflets usually crenate, the small polygamo-dioecious
flowers crowded in branched panicles. Calyx lobes obtuse, 5,
FLORA OF PERU 257
imbricate as the incurved petals. Stamens 5, inserted outside the
disk base, the distinctly dorsifixed anthers on filiform-subulate
filaments. Ovary 1-celled with solitary compressed ovule pendulous
from ascending basal funicle. Styles connate or short or obsolete,
the often more or less medially lateral stigmas sessile on the lightly
falcate samara, the oblique obtuse wing-portion membranous,
nervose. Seed testa membranous, embryo curved.
Loxopterygium huasango Spruce ex Engl. in DC. Monogr.
Phan. 4:461. 1883.
Younger branches, petioles and panicles pilose with short and
long trichomes intermixed, the 4-5 pairs of oblong subacute sub-
sessile leaflets white-hirsute on the midnerve and nerves beneath,
otherwise glabrous, the lower 6 cm. long, half as wide, the inter-
mediate 6-8 cm. long; panicles about 1 dm. long, the secondary
branchlets 3-5 cm. long, pedicels 1-3 mm. long; calyx segments ovate,
shortly pilose, the oblong petals about four times longer or 2 mm.
long; filaments nearly 3 mm. long; samara 1.5 cm. long, 5-6 mm.
broad, the lower part including the seed 6 mm. long. Tree 8-10
meters tall, the wood hard. Illustrated, Weberbauer, 151; Svenson,
Amer. Journ. Bot. 33: 400 (branch). F.M. Neg. 13184.
The bark and leaves have a resin which affects the skin much
as in Rhus (Svenson).
Piura: Quebrada Ancha near La Brea, ( Haught & Svenson 11558}.
Above Payta, Weberbauer, 153. Ecuador. "Huasango."
9. SCHINOPSIS Engler
Younger branchlets puberulent, glabrous and blackish in age,
the leaves either odd- or even-pinnate, the common petiole narrowly
winged, the leaflets many pairs, small, entire. Flowers polygamous,
the ovate sepals definitely imbricate, the 5 oblong-elliptic petals
with prominent midnerve within, concave, spreading-reflexed. Disk
thickish, the short filaments inserted in each sinus of the 5 lobes.
Ovary rudimentary in male flowers, ovoid and strongly compressed
in female, the sessile stigma lateral, the pendulous ovule conforming
to the single cell. Calyx persisting at base of cultriform samara,
the endocarp osseous. Embryo curved as in the similar Loxop-
terygium.
Schinopsis peruviana Engl. in Mart. Fl. Bras. 12, pt. 2: 405.
pi 86. 1876; 462.
258 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Densely leafy branchlet tips, petioles and leaves beneath more
or less conspicuously pilose, the leaves 1-1.5 dm. long with as many
as 15 pairs of spreading opposite oblong sessile obtusish inequilateral
but subequal leaflets, the median longer, about 1.5-2 cm. long, 4-5
mm. wide; panicles 1.5 dm. long, the secondary branches spreading,
the subsessile flowers crowded apically, with semiovate bracts
scarcely 1 mm. long; sepals ovate, obtuse, pilose at base, marginally
ciliolate, 0.7 mm. long; petals oblong-ovate, 2 mm. long, half as wide,
subequaled by the stamens; thick wings of fruit thin-edged, medially
1 cm. wide, 2 cm. long, 5 mm. wide at obtuse tip. Cf. also S.
Haenkeana Engl. I.e. Illustrated, Engl. I.e. pi. 15, figs. 6-7.
San Martin: Near Tarapoto, Spruce 4549, type; Williams 5727.
"Baloquiro," "cocobolo."
10. HAPLORHUS Engler
Simple-leaved shrub or tree with slender branchlets, the short
axillary floriferous composite-paniculate with secondary or tertiary
flexuose racemose divisions. Flowers (male unknown) sessile in
axils of obovate bracts subtending the 5 imbricate sepals. Ovary
obovoid, laterally compressed, 1-celled, the solitary ovule pendent
from a basal funicle. Stigmas 3, sessile, minute, triangularly placed
laterally below the apex. Drupes obliquely obovoid, compressed
laterally, exocarp and mesocarp thin, endocarp coriaceous, the seed
with thin testa.
Haplorhus peruviana Engl. Bot. Jahrb. 1: 419. 1881; 283.
Glabrous (unless panicles) the sparsely leafy slender glaucescent
branches with internodes about 1 cm. long; leaves linear-lanceolate,
narrowed to the scarcely petioled base, coriaceous, the lateral nerves
immersed, 5-9 cm. long, 5-10 mm. wide; panicles obscurely puberu-
lent, 2-3 cm. long; pedicels 0.5 mm. long; bracts ciliate, nearly 1 mm.
long; drupes 3.5 mm. long, 3 mm. thick. Wide crowned-tree to
15 meters tall with short irregular trunk, dark brown fissured bark,
with Schinus molle dominant in Mantaro Valley for some 50 kilom-
eters below La Mejorada, most plentiful on canyon floor but extend-
ing up the slopes singly or in groups to about 500 meters; collected
by Dr. 0. Usandivaruas of Andahuaylas (West). Illustrated,
Engler, I.e. pi. 9.
Huancavelica: Rocky canyon slopes, La Mejorada to Mayoc,
3,000 meters, West 8354 (det. Johnston). Puno: Gay, type.
FLORA OF PERU 259
CELASTRACEAE. Bittersweet Family
Reference: Loesener, Pflanzenfamilien ed. 2. 20b: 87-197. 1942.
Trees or shrubs, sometimes scandent, the simple leaves alternate
or opposite, the stipules, if present, small and caducous. Flowers
small, cymose or fasciculate, often perfect, the 4-5 calyx lobes as
the 5 petals imbricate or rarely valvate, the latter alternate with
the 4-5 stamens (these exceptionally more) and inserted on the
ordinarily flat and fleshy disk or this rarely obsolete. Anthers
2-celled, longitudinally dehiscent. Ovary superior, free or adherent,
1-5-celled, the short style more or less 3-lobed; ovules mostly 2
from the inner angle of each cell. Fruit various but seeds mostly
with abundant endosperm and rather large straight embryo, the
flat cotyledons thin. Often said to be distinguishable from Aqui-
foliaceae by the presence of the disk and the usually capsular
dehiscent fruit but these characters are not in themselves definitive
for two Peruvian genera, as noted below. The genus Alzatea R. & P.
formerly included here will be placed for convenience in this work
in the Rhamnaceae. Cf . Loesener, I.e. 196. The genus Cheiloclinium
Miers, transferred by Loesener, I.e. 17, here from the similar Hippo-
crateaceae, is retained for expediency in that family.
Fruit dehiscent; seeds arillate; ovules usually 2; disk present, un-
dulate; leaves entire unless above middle; flowers axillary.
1. Maytenus.
Fruit drupaceous, sometimes coriaceous; ovules solitary; disk obso-
lete or 4-5-lobed; leaves serrate if flowers axillary (Peru).
Inflorescences fasciculate, forked; leaves entire or nearly.
2. Rhacoma.
Inflorescences simple (known); leaves serrate (Peru).
3. Schaefferia.
1. MAYTENUS Feuill^e
Usually glabrous or essentially glabrous shrubs or small trees
with persisting coriaceous often distichous alternate petioled leaves,
the minute stipules deciduous, and small polygamous axillary white
or yellowish flowers. Calyx 5-parted, the petals spreading. Stamens
5 (filaments subulate, anthers ovate-cordate), inserted below the
orbicular undulate-margined disk in which the 2-4-celled ovary is
immersed and confluent; ovules solitary or geminate, erect. Style
none or short, the stigma 2-4-lobate. Fruit capsular, coriaceous,
1-3-celled, loculicidally 2-3-valvate, the seed arillate.
260 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
It has not been practical for me to attempt evaluation of the
many species proposed in this group; the key is only suggestive and
some of the characters indicated for reasons of expediency will
prove, it seems to me, to be less significant when a comprehensive
study is possible. Cf. Rhacoma for M. magnifolia Loes. Verh. Bot.
Ver. Brandenb. 48: 176. 1907.
Leaves all or many distinctly shorter than 3 cm.
Leaves more or less oblanceolate, cuneate to base.
Leaf -nerves obsolete above M. andicola.
Leaf -nerves and usually the veins obvious above.
Leaves a cm. wide or narrower M. cuzcoina.
Leaves, at least some of them, wider M. apurimacensis.
Leaves more or less obovate or suborbicular, at least in Peru,
rounded to acute at base.
Leaves about half as wide as long.
Inflorescences simple M. conferta.
Inflorescences peduncled M . boarioides.
Leaves more than half as wide as long M. octogona.
Leaves all or many 3 cm. long or longer.
Leaves obviously if minutely serrate, at least toward the tip.
Leaves more or less oblong-lanceolate, often about half as wide
as long or even narrower.
Nerves obsolete (Peru) ; leaves coriaceous . . . . M. alaternoides.
Nerves more or less obvious.
Leaves coriaceous; veins obscure.
M. pseudoboaria, M. Pavonii, M. verticillata.
Leaves membranous with reticulate venation both sides.
M. Jelskii.
Leaves more or less elliptic or more than half as wide as long.
Leaves conspicuously acuminate M. Krukovii.
Leaves shortly if at all acuminate, often obtuse or emarginate.
M. durifolia, M. retusa.
Leaves quite or essentially entire, even at tip.
M. tarapotensis, M. macrocarpa.
Maytenus alaternoides Reiss. in Mart. Fl. Bras. 11, pt. 1: 15.
1861.
Glabrous; branchlets terete, many; petioles 4 mm. long, sub-
margined by the decurrent blade, this 3.5-7 or 8 cm. long, 2-3 cm.
FLORA OF PERU 261
wide, oblong or narrowly lanceolate and acuminate or elliptic and
cuspidate, crenate-dentate nearly to base, the 8-15 nerves both sides
or beneath prominent or obsolete, often drying brownish above,
early glaucous, finally coriaceous; cymes 2-3 times longer than
petioles, fasciculate or branched; basal bractlets of filiform pedicels
deltoid, stipuliform; flowers 3-4 mm. wide; calyx segments rounded,
half as long as the oval or obovate petals; disk fleshy, subpentagonous;
filaments subulate, anthers subcordate; fruiting pedicels 4 mm. long;
style very short, stigma entire; capsules obovoid or subglobose, sub-
ligneous, 16-18 mm. long, the solitary ellipsoid compressed brown
lustrous seed 8-10 mm. long, the aril drying crustaceous (after
Reissek). Var. peruana Loes., Bot. Jahrb. 37: 574. 1906, has the
leaf nerves fainter above. Illustrated, Reissek, I.e. pi. 2, fig. 12
(leaves). F.M. Neg. 13270.
Cuzco: Prov. de la Convention, Valle del Urubamba, Weberbauer
5066 (type, var.); 245. Central Brazil.
Maytenus andicola Loes. Bot. Jahrb. 50: Beibl. Ill: 9. 1913.
Branchlets spreading, the older subterete, at the third year about
2 mm. thick, the young shoots pulverulent-papillose, sometimes
obsoletely; leaves crowded, the internodes 4-9 mm. long; petioles
1-3 mm. long; leaves lanceolate or elliptic-lanceolate to obovate-
elliptic, acute or narrowly cuneate at base, rounded or obtuse to
acute at tip, this mostly very minutely apiculate, 1-2.6 cm. long,
4-8 mm. wide, entire or rarely obsoletely and remotely appressed
serrulate, rigid-coriaceous, the 3-4 pairs of lateral nerves more or
less arcuate to tips, obsolete or faint above, sometimes rather
marked beneath, not reticulate; pedicels under anthesis scarcely
1 mm. long and flowers then hardly 2 mm. wide, the former in fruit
barely 2 mm. long, glabrous; calyx glabrous, except the fimbriolate-
ciliolate suborbicular lobes; petals irregularly orbicular, margins
glandular (?)-fimbriolate, about 1 mm. broad; anthers cordiform;
ovary 2-celled, stigma bilobed; capsules ellipsoid, 7-8 mm. long,
4-6 mm. wide, by abortion 1-celled, tardily dehiscent, 1-seeded, the
seed invested by the flat aril. Affine M. viscifolia Griseb. of Argen-
tina, with larger thicker leaves, larger flowers, ovary 3-celled
(Loesener). Aspect of M. cuzcoina and with it near probably M.
conferta.
Ancash: Closely branched shrub, Huaraz, 2525 (det. in herb.,
M. conferta). Ayacucho: Near Ayacucho in rocky shrubby out-
crops, 3,000 meters, Weberbauer 5507, type.
262 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
May terms apurimacensis Loes. Bot. Jahrb. 50: Beibl. Ill: 9.
1913.
Branchlets erect-spreading, finally terete and densely lenticellate,
by the second or third year to 5 mm. thick, the new shoots striate-
angulate, closely pulverulent-papillose, scarcely to 2 mm. thick; leaf
internodes 5-8 mm. long; petioles hardly 1 or to 2 mm. long; leaves
obovate- or rarely ovate-elliptic, acute or sometimes cuneate at base,
obtuse or mostly rounded at apex where often minutely apiculate,
1-3.2 cm. long, 4-19 mm. wide, the margin densely but frequently
obsoletely serrulate (under a lens), rigid-coriaceous, nitidulous and
very minutely closely punctulate both sides, lateral nerves 4-6 pairs,
equally prominent above and below, reticulation faint or obsolete;
flowers few, densely fascicled, under anthesis scarcely 2 mm. wide,
the glabrous pedicels 1.5 (-2) mm. long; calyx glabrous except the
ciliate obtuse lobes, these 0.75 mm. long; petals suborbicular, about
1.5 mm. wide; anthers subreniform, broader than long; stigma
capitellate. Said to attain 2 meters. Affine M. cuzcoina Loes.,
perhaps a variant^ but larger leaves broader, rounded at tip and
nerves more manifest (Loesener). The var. Trollii Loes., Notizbl.
Bot. Gart. Berlin 13: 219. 1936, has slightly narrower or narrowly
elliptic and thinner leaves. F.M. Neg. 13268.
Apurimac: Between Rio Pachachaca and Rio Pampas, 2,400
meters, Weberbauer 5873, type. Cuzco: Yucay, Valle del Urubamba,
Herrera 2556 (var. Trollii). Bolivia. "Paltai-paltai" (Herrera).
Maytenus boarioides Loes. Bot. Jahrb. 36: 381. 1905.
Glabrous, the lightly striate shoots 1-1.5 mm. thick; petioles
2-4 mm. long; leaves typically oblong or lanceolate-oblong, cuneate
(in Peru merely acute) at base, acute or obtuse and minutely apicu-
late, 2-2.5 cm. long, 9-12 mm. wide, coriaceous, above slightly
lustrous, the acute midnerve prominent, the 4-7 slender lateral
nerves obsolete, these rather prominent beneath and reticulate
especially toward the serrulate margins; inflorescences solitary in
axils, only about 1 mm. long, peduncled, bifurcate, many-flowered,
pedicels 2-3 mm. long, bracts deltoid, sparsely fimbriolate, scarcely
1 mm. long; flowers under anthesis about 3 mm. wide; sepals rotund,
fimbriolate, less than half as long as the subrhombic-ovate petals;
stamens inserted without and within the subpulvinate obsoletely
lobed disk; ovary costate, scarcely semi-immersed, 2-celled, ovules
solitary, stigma capitate, 4-sulcate; capsule bivalved, the valves
6-7 mm. long, arillate base nearly to middle. Much like M. boaria
FLORA OF PERU 263
Molin. but that species with less densely serrulate, more obtuse leaves
(Loesener). The peduncled inflorescences separate it from M.
cuzcoina and M. apurimacensis. F.M. Neg. 13273.
Huanuco: Yanahuara Valley, Huanta, Weberbauer 7600; 7599.
Ecuador.
Maytenus conferta (R. & P.) Reiss. & Loes. in Pflanzenfam. 3,
Abt. 5: 206. 1892. Celastrus confertus R. & P. Fl. Peruv. 3: 7. 1802.
Much-branched, the crowded verticillate branchlets as the short
petioles lightly pubescent; leaves black-serrulate above the middle,
to mucronate tip, oval or subrotund or often ovate-oblong (at least
about 10 mm. long, half as wide), coriaceous, lustrous above, venose;
pedicels many, short, the solitary whitish flowers minute; capsules
2-3-celled and with as many valves and seeds, the latter obovoid
with fleshy yellowish aril. Pedicels according to Briquet scarcely
as long as flowers. M. Jamesonii Briq. Ann. Cons. Jard. Bot.
Geneve 20: 353. 1919, of Ecuador is similar with nerves prominently
reticulate beneath but the branchlets are glabrous, flowers larger,
the ovate-triangular petals about 1 mm. long (Briquet). F.M.
Neg. 13280.
Huanuco: In cold woods toward Muna and Pillao, Ruiz & Pavdn,
type. Monzon, Weberbauer 3385 (det. Loesener); 253. San Martin:
Vitoc, Ruiz & Pavdn.
Maytenus cuzcoina Loes. Bot. Jahrb. 37: 574. 1906.
Branchlets terete, the grayish bark densely lenticellate and longi-
tudinally rimulate, the slender tips under a lens most minutely
pulverulent-papillose; petioles about 1 mm. long; leaves about
oblanceolate, above the middle broader, subentire or obsoletely and
appressed serrulate, narrowly acute to base, rounded, obtuse or
subacute at tip, only under a lens apiculate, 12-25 mm. long, 3.5-
7 mm. wide, coriaceous, concolorous, lateral nerves few, obvious
above, prominent or obsolete beneath, the reticulation inconspicuous;
flowers 5-merous, fasciculate in the leaf -axils, greenish, the glabrous
pedicels about 1 mm. long; sepals rotund, under lens denticulate-
subciliate, scarcely 0.5 mm. long and wide; petals broadly ovate-
suborbicular, little oblique, about or hardly 1.5 mm. long, 1 mm.
wide; stamens inserted in fleshy pentagonally angled disk, subequal-
ing sepals, subreniform anthers scarcely as long as subulate filaments;
ovary conical (stigma capitate, obsolete or slightly bilobulate),
264 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
2-celled, cells biovulate, ovules erect. M. boaria Molin. has the
leaves more narrowed to the tip (Loesener). F.M. Neg. 23248.
Ancash: Openly branched canyon shrub, Llata, 2290 (det. in
herb. M. conferta). Ayacucho: Tambo, Weberbauer 5553 (det.
Loesener). Cuzco: Valle del Urubamba, 3,200 meters, in open
thicket along brook, Weberbauer 4915, type. Rio Chaupimayo,
Soukup 595 (det. in herb. M. andicola). Ollantaytambo, Cook &
Gilbert 418 (det. in herb. M. conferta}. "Paltay-paltay" (Cook &
Gilbert).
May terms durifolia Briq. Ann. Cons. Jard. Bot. Geneve 20:
351. 1919.
Glabrous except the ciliate bracts; branchlets subterete; petioles
to 5 mm. long; leaves nearly elliptic, acute at base, contracted to
obtusish tip, 3-4 cm. long, 2-2.7 cm. wide, coriaceous, slightly
lustrous and obsoletely venose above, paler beneath, the 6-8 lateral
nerves there very fine; flowers fasciculate from a pulvinus of crowded
apiculate bracts; pedicels to 3 mm. long; sepals ovate, fimbriate, to
0.6 mm. long; petals oblong-ovate, rounded at tip, subfimbriate, to
1.2 mm. long, 1 mm. wide; disk about 1.2 mm. wide. Allied to
M. retusa (Poir.) Briq. as determined in herbaria but easily dis-
tinguished by leaves obtuse both ends, fewer nerves, shorter pedicels
(Briquet). F.M. Neg. 23249.
Peru(?): Without locality, Ruiz & Pavdn, type.
Maytenus Jelskii Zahlbr. in Szyszyl. Diag. PI. Nov. 1: 8. 1895
(Dissert. Classis Math. Phys. Acad. Litt. Cracov. 29: 222. 1895).
M. Mathewsii Briq. Ann. Cons. Jard. Bot. Geneve 20: 348. 1919.
Branchlets slender, subangled, minutely puberulent; petioles a
little puberulent, 2-3 mm. long; leaves oblong-lanceolate, acute both
ends or the lower obtuse, broadest above the middle, 2.5-4.5 cm. long,
1-18 mm. wide, green, paler beneath, glabrous, finely dentate toward
the apex, lateral nerves 10-12, prominently reticulate both sides;
inflorescence solitary, axillary, peduncles 5-7 mm. long, densely
spreading-puberulent, the 3-4 mm. long pedicels glabrescent; bracts
ciliolate; flowers 4-5 mm. wide; sepals oblong, rounded, about 0.6
mm. long; petals to 1.8 mm. long; disk nearly 1 mm. broad (after
Briquet). Marked by the membranous leaves with fine delicate
venation both sides and by the puberulence (Briquet). F.M.
Neg. 23257 (M. Mathewsii).
FLORA OP PERU 265
Cajamarca: Cutervo, Raimondi (det. Loesener); Jelski, type.
Amazonas: Chachapoyas, Mathews (type, M. Mathewsii). Cuzco:
Pumahuanca, 3,500 meters, Vargas 7692?
Maytenus Krukovii A. C. Smith, Journ. Arnold Arb. 20:
294. 1939. M. chuchuhuasha Raymond-Hamet & Colas, Compt.
Rend. Acad. Sci. Paris, 1886. 1937; (Dissert. Faculte" de Pharmacie,
Paris, 1937)?
Glabrous tree to 28 meters tall, the slender branchlets terete;
petioles nearly alate or conspicuously canaliculate, 6-9 mm. long;
leaves oblong-elliptic or obovate-elliptic, 9-14 (18) cm. long, 3.5-5 (7)
cm. wide, attenuate to base, acuminate (acumen about 1 cm. long),
obscurely crenulate-serrulate, coriaceous, the 7-9 lateral nerves im-
mersed above and obscure, lightly prominent beneath; inflorescences
axillary, glomerulate, 4-5 mm. across; flowers sessile or pedicels 0.5
mm. long, 50 or more to an inflorescence; sepals deltoid, 0.8-1 mm.
long, minutely glandular fimbriolate, subacute; petals oblong-deltoid,
0.8-1.2 mm. long, obtuse or rounded; disk fleshy, undulate; style
0.4 mm. long; capsules coriaceous, 2 cm. long, 14 mm. wide, bivalved.
M. ebeniifolia Reiss. of Brazil and M. Karstenii Reiss. of Colombia
apparently resemble this in foliage but both have pedicellate flowers,
those of the latter in fascicles. M. micrantha A. C. Smith, I.e., 295,
has smaller thinner leaves with spreading lateral nerves obvious on
both sides.
Probably this is M . chuchuhuasha as suggested by the collector
but that was described in rather running comment and in French;
while it would be appropriate for the scientific and native names to
coincide, and convenient, it is better in the interest of uniformity to
accept the equally well-chosen name used when the plant was
described, in conformity with modern custom, in Latin and with
formal typification.
According to P. Le Cointe, Arvores e Plantes Uteis 120. 1934,
the bark yields an alkaloid which is a powerful stimulant similar
to cathin from the leaves of Catha edulis Forsk.
Peru: Apparently the type of M. chuchuhuasha; see note above.
Brazil. "Chuchuhuasca," "chuchuhuasha."
Maytenus macrocarpa (R. & P.) Briq. Ann. Cons. Jard. Bot.
Geneve 20: 361. 1919. M. multiflora (R. & P.) Loes. Bot. Jahrb.
36: 380. 1905, not Reiss. Haenkea multiflora R. & P. Syst. Veg. 65.
1798. Celastrus macrocarpus R. & P. Fl. Peruv. 3: 8. pi. 230. 1802.
266 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Haenkea macrocarpa (R. & P.) Steud. Nomencl. ed. 1: 170 and 387.
1821.
Glabrous with verticillate branches, the leafy branchlets angulate;
petioles to 4 mm. long; leaves oblong-lanceolate or elliptic, entire,
acuminate but emarginate, coriaceous, lustrous above, 1-2 dm. long;
flowers 5-merous, minute, many in the axils; calyx colored, the teeth
deciduous; petals obovate, whitish, spreading; styles mostly simple;
capsules obovoid, often 1-seeded, 2-3-valved, the oblong seeds with
white aril. F.M. Neg. 13298.
Huanuco: Muna and Chacahuasi, Ruiz & Pavdn, type. San
Martin: Zepelacio, King 3721. Ecuador.
Maytenus octogona (L'He>.) DC. Prodr. 2: 9. 1825. Celastrus
octogonus L'HeY. Sert. Angl. 7. 1788. Ziziphus peruviana Lam.
Encycl. 3: 320. 1789. Maytenus obovata Hook. f. Trans. Linn. Soc.
20: 230. 1847, fide Svenson. Celastrus orbicularis Willd. ex Roem.
& Schult. Syst. 5: 423. 1819. Maytenus uliginosa HBK. Nov. Gen.
6 Sp. 7: 65. 1824. M. orbicularis (Willd.) Loes. Hot. Jahrb. 50:
Beibl. Ill: 10. 1913.
Sprawling to erect shrubs often 3 meters tall with glabrous or
minutely pubescent branchlets and with fleshy leaves varying from
ovate to suborbicular or obovate, these commonly 1-2 cm. long,
7-12 mm. wide, the margins entire to repand dentate; petioles about
2 mm. long; pedicels 2-3, axillary, fasciculate, usually bracteate,
2-3.5 mm. long, glabrous or pubescent as the calyces; capsules ovoid,
obtusely trigonous, usually 3 (2-4) -celled, thick- or thin-walled, about
7 mm. wide and long. In part after Svenson, Amer. Journ. Bot. 33:
461. 1946. Illustrated, Svenson, I.e. p. 462. F.M. Neg. 35918.
Tumbez: Cancas, Weberbauer 7754. Piura: Talara, Haught 58;
76 (det. Svenson). Paita, Haught 97; Gaudichaud. Rio de Lomas,
700 meters, Weberbauer 5740. La Brea, ( Haught & Svenson 11573).
lea: Above Pisco, near Santa Ines Mine, 4,600 meters, Weberbauer
5381 (det. Loesener, leaves narrower and thicker). Ancash: River
valley gravel, 2565. Tacna: Near Tacna, Rusby 2618. Chile;
Ecuador; Galapagos. "Realingo" (Weberbauer).
Maytenus Pavonii Briq. Ann. Cons. Jard. Bot. Geneve 20:
349. 1919.
Glabrous or nearly, the rather stout branchlets angled, the petioles
1-2 mm. long; leaves narrowly lanceolate-oblong, minutely dentate
above the cuneate base, acute or subacute, 2-4 cm. long, 7-10 mm.
FLORA OF PERU 267
wide, coriaceous, lustrous above, the few indistinctly pinnate lateral
nerves sub-immersed; peduncles rarely to 5 mm. long, pedicels to
3 mm. long; bracts obscurely ciliolate; sepals to 0.6 mm. long; petals
obovate, fimbriolate, 1 mm. long; disk nearly 1 mm. wide. Easily
distinguished from M. verticillata by the smaller indistinctly crenate-
dentate leaves, less apparent venation, few-flowered inflorescence,
smaller flowers and fimbriolate petals (Briquet). F.M. Neg. 23259.
Peru(?): Without data, Pavdn, type.
Maytenus pseudoboaria Loes. Notizbl. Bot. Gart. Berlin 12:
29. 1934.
Glabrous shrub with spreading or erect branchlets, the older
densely lenticellate, the younger sulcate-striate; petioles 2-5 mm.
long; leaves lanceolate or oblanceolate, rarely suboblong or obovate-
elliptic, cuneate or acute at entire base, densely serrulate to tip,
obtuse or subacute, 2.5-5 cm. long, 8-17 mm. wide, coriaceous or
chartaceous, the 5-7 lateral nerves rather prominent both sides, the
reticulation only so beneath; inflorescences solitary or lateral or in
the axils and 1- or 1-3-flowered, the peduncles of the latter 3-6 mm.
long, the pedicels of the 1-flowered 4-7 mm. long, of the 2-3-flowered
to 4 mm. long and minutely bracteolate; flowers in bud (type),
sepals rounded; capsules 3-celled, 2-3-valved, the valves 5 mm. long
and wide; seed ellipsoid, 4 mm. long, more or less arillate. The
description includes the var. triantha Loes. and the var. monantha
Loes., the species according to the author much like both M. boaria
Molin. of Chile, with more coarsely serrate and acute leaves, flowers
mostly fascicled in the axils, and M. boarioides Loes., with more
oblong shorter leaves, 2-celled ovary and capsule; also M. Pavonii
Briq. ex char, seems to be approximate but the smaller leaves are
said to be denticulate-crenate, the lateral nerves sub-immersed
above. The name was originally spelled as above and without
hyphen. F.M. Neg. 29280 (var. triantha).
Huanuco: Macora, Ruiz & Pavdn (type, var. triantha). Bolivia.
Maytenus retusa (Poir.) Briq. Ann. Cons. Jard. Bot. Geneve
20: 351. 1919. Celastrus retusus Poir. Encyl. Suppl. 2: 146. 1811.
C. emarginatus R. & P. Fl. Peruv. 3: 6. pi. 229. 1802, not Willd.
Glabrous unarmed shrub, the erect little leafy stems with a few
slender angulate branchlets; petioles 4 mm. long, canaliculate above;
leaves ovate-oblong-elliptic, acuminate but emarginate at tip,
slightly crenate, coriaceous, venose both sides, lustrous above;
268 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
peduncles congested in the axils, 1-flowered, unequally elongate;
petals yellow, spreading; stigma often entire or bifid or trifid;
capsules obovoid, yellow, 1-3-celled (Ruiz & Pavon). Allied to
M. conferta (R. & P.) Briq. but leaves much larger, obtuse or emargi-
nate, not apiculate, rounded at base, lateral nerves more numerous,
petioles to 5 mm. long, flowers larger on elongate pedicels (Briquet).
F.M. Negs. 13286; 23267.
Huanuco: Near Mufia, Ruiz & Pavdn, type.
Maytenus tarapotensis Briq. Ann. Cons. Jard. Bot. Geneve
20: 264. 1919.
Glabrous, except the ciliate bracts of the pulvinus of the axillary
congested flowers; petioles 5-8 mm. long, sulcate above; leaves
oblong or lanceolate, cuneate at base and somewhat narrowed to
obtuse tip, entire, to 8 cm. long, 2.8-3.7 cm. wide, coriaceous, the
8 lateral nerves immersed on upper lustrous surface, subimmersed
on opaque not at all papillose under side; pedicels to 3 mm. long;
sepals ovate, scarcely longer than 0.5 mm.; petals obovate, rounded,
about 1 mm. long, 0.8 mm. wide; disk about 0.7 mm. wide. Allied
to M. ligustrina Reiss. of Brazil but remarkable by the conspicuously
immersed nerves (Briquet); but the type was referred by Loesener
to M. macrocarpa. F.M. Neg. 23022.
San Martin: Campana near Tarapoto, Spruce 4329, type.
Maytenus verticillata (R. & P.) DC. Prodr. 2: 10. 1825.
Celastrus verticillatus R. & P. Fl. Peruv. 3: 6. pi. 229. 1802.
Stems terete, nigrescent; branches aggregate-verticillate, virgate,
erect, the younger angled, leafy, often purplish as the short petioles;
leaves crowded, often glaucous, oblong-lanceolate or oblong-obovate,
serrulate, lustrous both sides, venose beneath, coriaceous, to 1 dm.
long; peduncles solitary and aggregate, 1-5-flowered, the pedicels
short; calyx segments rounded, half as long as the ovate spreading
petals; disk plane; style obsolete, stigmas 3; capsules 3-celled, 3-
valved, the solitary seed with aril (Ruiz & Pavon). Flowers greenish-
white; seeds orange-yellow. To 8 meters tall. The var. latifolia
Loes., Bot. Jahrb. 36: 381. 1905, of Ecuador has leaves to 8.5 cm.
long, 3-5 cm. wide, capsules to 6 mm. in diameter; var. Lehmannii
Loes., Rep. Nov. Sp. 1: 161. 1905, from near Cuenca recedes from
the type in its less densely serrulate leaves, obviously long-ciliate
sepals and smaller 2-merous ovary. F.M. Neg. 13326.
FLORA OF PERU 269
The strong wood is very suitable for handles of axes and other
tools (Ruiz & Pavon).
Huanuco: Mito, 1478; 1947; 3418. Cani, 3456; 3479. Muna,
4059; Macora and Muna, Ruiz & Pavon, type. Yanano, compact
3-meter tree, 3762 (?leaves thinner). Junin: Huasahuasi, Ruiz &
Pav6n, type (part). Near Palca, Weberbauer 1768; 2181; 246.
Ayacucho: Aina, 8-meter tree, Killip & Smith 22517; 23139
Apurimac: Rio Pinco, 3,200 meters, Weberbauer 5912; 591 4-
Cuzco: Valle de Santa Ana, about 2,000 meters, Weberbauer 5914.
Valle del Paucartambo, 3,400 meters, Herrera 2961. Bolivia?
Chile? "Pilpus," "duraznillo" (Herrera), "rurama" and "picma"
or "pigna" (Ruiz & Pavon).
2. RHACOMA L.
Glabrous or pubescent with mostly opposite (in Peru alternate)
sometimes crenate leaves, the stipules small, and inconspicuous
flowers cymose or subsolitary at the ends of short or long peduncles.
Calyx urceolate-patelliform with 4-5 rounded lobes, the 4-5 petals
reflexed. Stamens 5, inserted between the lobes of the nearly plane
disk, the filaments subulate, the anthers subglobose. Ovary con-
fluent with disk, usually 3-4-celled, the very short style with 3-4-
lobate or punctiform stigmas; ovules erect, solitary in each cell.
Fruit dry or fleshy, coriaceous or drupaceous, most often 1-celled,
the erect seeds without an aril. The alternate-leaved species are
probably better referred to Maytenus (Bentham & Hooker). In-
deed M. magnifolia Loes. Verh. Bot. Ver. Brandenb. 48: 176. 1907,
in fruit and not placed by me, might be sought here.
Rhacoma Urbaniana Loes. Repert. Nov. Sp. 1: 162. 1905.
Glabrous, the subterete shoots 1-2 mm. thick, the older branch-
lets twice as stout and plicate-striate; petioles 5-10 mm. long; leaves
alternate, oblong-lanceolate, cuneate at base, mostly long-acuminate
or acute, entire or the younger minutely and sparsely callous-
denticulate (under lens), 13-21 cm. long, 3.5-5.5 cm. wide, charta-
ceous, with 7-9 lateral nerves arcuate at tip, rather prominent both
sides, the reticulation lax and obsolete; inflorescences fasciculate in
the axils, usually to 4-furcate, the peduncles 2-10 mm. long; bracts
obtuse, 0.5 mm. long; pedicels 2.5-5 mm. long; flowers 4-merous;
calyx patelliform, the rounded lobes scarcely 0.5 mm. long, about
0.75 mm. wide; petals imbricate, nearly 2 mm. long, 0.75 mm. wide,
the stamens half as long; disk inconspicuous; ovary 4-celled, sub-
270 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
lageniform, stigma punctiform. About 4 meters tall; nearest the
smaller-leaved R. spathulifolia Urban of Cuba according to the
author. F.M. Neg. 13332.
Junin: In open woods near La Merced, 1,000 meters, Weberbauer
1875, type; 282.
3. SGHAEFFERIA Jacq.
Rigid shrubs with alternate coriaceous usually entire (in Peru
serrate) leaves with small pedicellate or sessile dioecious 4-merous
flowers in their axils. Sepals orbicular. Petals oblong-elliptic,
obtuse. Disk small or none. Ovary sessile, the erect ovules solitary
in the 2 cells. Drupes with 2 pyrenes, the erect seed without aril.
Near Rhacoma but unique because of the reduced disk; in the male
flowers it approaches Aquifoliaceae (Loesener). The name com-
memorates Jakob Christian Schaeffer (1718-1790), author of a
notable work on the fungi of Bayern.
Schaefferia serrata Loes. Repert. Nov. Sp. 1: 162. 1905.
About 2 meters tall with spreading terete minutely pulverulent
branchlets 1-2 mm. thick, elongate or short, the latter often leafless
and bearing the fasciculately congested dioecious flowers; petioles
about 1.5 mm. long; leaves spathulate-obovate, cuneate at base,
obtuse or rounded at tip, denticulate-serrate, 6-11 mm. long, 3-7 mm.
wide, obsoletely nerved; male flowers 4-merous on older wood in
the axils of fallen leaves or sometimes in those of new leaves, about
3 mm. wide, pedicels slender, 3-5 mm. long; sepals free, rounded,
scarcely 0.5 mm. long and wide; petals free, elliptic, about 3.5 mm.
long (stamens much shorter), 1.75 mm. wide; disk obscure or none;
pistil little rugose, bilobed at tip. Differs from other species in
the serrate leaves. F.M. Neg. 13331.
Cajamarca: Chepe"n to Cajamarca, Prov. Pacasmayo, 1,000
meters, Weberbauer 4811, type; 188.
AQUIFOLIACEAE. Holly Family
Reference: Loesener, Monogr. Aquifol., Nov. Act. Acad. Leop.-
Carol. 78: 1-598. 1901; 89: 1-314. 1908.
Mostly evergreen, usually glabrous trees or shrubs with alternate
simple estipulate leaves and axillary perfect or polygamous 4-9-
merous flowers, sometimes solitary, often in cymes, racemes or
FLORA OP PERU 271
fascicles. Corolla imbricate in bud. Stamens as many as corolla
parts or petals and alternate with them, usually affixed to base.
Anthers 2-celled, oblong to subglobose, opening longitudinally.
Disk usually lacking. Ovary (3)4-9-celled; style terminal or obso-
lete, the pendulous ovules 1 or 2 in each cell. Fruit a drupe, often
berry-like, of (3)4-9 pyrenes, the seeds with abundant endosperm
and small straight embryo. Leaves opposite in one Philippine
species.
ILEX L.
Paltoria R. & P. Prodr. 13. 1794. Izquierdia R. & P. I.e. 140.
pi. 30.
Flowers often perfect, with persisting 4-5-parted calyx and rotate
4- rarely 5-6-parted corolla, the petals or lobes imbricate in bud,
ovate to suborbicular or elliptic, rounded or very obtuse. Anthers
ovoid or ellipsoid. Ovary 2-4-6(7-22)-celled. Drupes globose or
ellipsoid with 3-4 (or rarely many) pyrenes. Otherwise character
of the family which it shares with only two other genera.
The monograph of Loesener will doubtless become a classic for
its period; nevertheless modern research methods in systematic
botany may result in considerable change in his taxonomy of the
group. My compilation of course is directly from Loesener except
the key, which has been devised primarily on the basis of conveni-
ence and is only suggestive, since it accounts for little possible varia-
tion.
In South America the genus is of particular interest as the source
of the tea-like beverage "mate," which contains caffein, however,
considerably less than tea or coffee (cf. Loesener, I.e. 2: 254). A
principal source is I. paraguayensis St. Hilaire but a number of
other species serve apparently as well, including the Peruvian I.
tarapotina Loes.; cf. Sparn, Rep. Argent. Acad. Nac. Cienc. Miscel.
no. 22. 1937, and Sprecher von Bernegg, Trop. Weltwirtschafts-
pflanzen 3. 1936.
Leaves emarginate, rounded or obtuse (rarely mucronate or barely
acute), less than twice as long as wide (unless /. microsticta
and I. loretoica, imperfectly known).
Leaves small, mostly or all 3 (4) cm. long or shorter.
Leaves epunctate beneath.
Leaves entire.
Midnerve villous, at least of mature leaves.
272 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Leaves 4-7 mm. wide I. ericoides.
Leaves 15 mm. wide or wider I. villosula.
Midnerve of medium-sized leaves glabrous 7. quitensis.
Leaves serrulate 7. caniensis, I. elliptica.
Leaves punctate beneath, glabrous or puberulent.
Branchlets pulverulent; male flowers solitary. 7. Weberbaueri.
Branchlets pilose-puberulent; male flowers 3.
Leaves 1-3 cm. long.
Nerves of mature leaves obsolete above.
7. Lechleri, I. ovalis.
Nerves of mature leaves rather prominent both sides.
7. Matthewsii.
Leaves 7 to rarely 11 mm. long 7. minutifolia.
Leaves medium size, usually or mostly 4 cm. long or much longer.
Leaves pubescent beneath, at least midnerve, even at maturity,
more or less ovate.
Leaves 5-9 cm. long, asperous above 7. trachyphylla.
Leaves 3-5.2 cm. long, glabrous or pubescent above.
7. sessiliflora.
Leaves glabrous or essentially even beneath.
Leaves mostly or all notably narrower than 5.5 cm.; female
calyx lobes acuminate 7. microsticta.
Leaves mostly or all notably wider than 5.5 cm.; female
calyx lobes rounded to acute.
Leaves (9) 11-18 cm. long.
Petioles 2-6 mm. long 7. loretoica.
Petioles 10-25 mm. long.
Leaves subentire 7. anonoides.
Leaves serrulate 7. teratopis.
Leaves 5-10.5 cm. long 7. crassifolioides, I. crassifolia.
Leaves obviously but sometimes shortly acute or acuminate.
Petioles 3-6 mm. long; leaves 3.5-6 cm. long, serrulate.
Leaves punctate beneath 7. cuzcoana, 7. andicola.
Leaves epunctate beneath 7. hippocrateoides.
Petioles 8-17 mm. long; leaves mostly or all longer than 6 cm.,
entire or subentire.
Leaves 2-6 cm. wide, ovate-lanceolate or elliptic.
FLORA OF PERU 273
Leaves glabrous.
Leaves acutely subcaudate-acuminate I. tarapotina.
Leaves shortly and subobtusely acute or acuminate.
Leaves all about 1.5 dm. long I. aggregata.
Leaves mostly 5-10 cm. long, rarely some larger.
Leaves punctate beneath I. andarensis.
Leaves epunctate beneath 7. inundata.
Leaves puberulent, especially beneath 7. Jelskii.
Leaves 9-18 mm. wide, oblong-lanceolate 7. amygdalina.
Ilex aggregata (R. & P.) Loes. Notizbl. Bot. Gart. Berlin 11: 95.
1931. Izquierdia aggregata R. & P. Syst. 278. 1798. 7. amygdalifolia
Rusby, Mem. Torrey Bot. Club 3, no. 3: 15. 1893; 128, fide Loesener.
Glabrous unless for the umbelliform solitary or usually fascicled
often axillary inflorescences; branchlets lenticellate, about 3 mm.
thick, the slenderer shoots densely striate; stipules 1 mm. long;
petioles 5-10(17) mm. long, 1.5-2.5 mm. thick; leaves ovate-lanceo-
late or elliptic, rounded or obtuse at often slightly oblique base, acute
or acuminate, 8-19 cm. long, 2.5- nearly 7 cm. wide, entire margin
obscurely or not recurving, submembranous or thin-coriaceous,
slightly lustrous above, paler beneath, the 10-17 lateral nerves very
slender above, prominent beneath with the conspicuous reticulate
venation; flowers 3-many, inflorescence 2-3-dichotomous, bracts
scarcely 0.5 mm. long and wide, peduncles 9-25 mm. long, ultimate
pedicels 3-5 mm. long; male flowers 4(5)-merous, calyx 1-1.25 mm.
wide, lobes lacinulate-sinuate, 0.5 mm. long, acute; petals oval,
about 2.2 mm. long, the subequal stamens with ovoid anthers;
drupes globose, scarcely 5 mm. thick, stigma 4-lobed, pyrenes 4 or 5,
smooth but very finely striate-rugulose. A 4-meter shrub or a IO-
meter tree (Weberbauer) ; determinations by Loesener.
Huanuco: Macora, Ruiz & Pavdn, type. Pozuzo, Weberbauer
6768; 6746; 6778. Cueva Grande near Pozuzo, 4764 Junin: La
Merced, 5537. Bolivia.
Ilex amygdalina Reiss. ex Loes. Monogr. Aquifol. 1: 299. 1901.
Branchlets ascending, the older to 5 mm. thick, the puberulent
shoots scarcely 1 mm. thick with minute subulate-deltoid stipules;
petioles 9-11 mm. long, sulcate, 1.5 or at base nearly 2 mm. thick;
leaves lanceolate, acute or cuneate-acute at base, more or less ob-
tusely acuminate, 6- nearly 9 cm. long, 9-18 mm. wide, subentire
274 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
or remotely and obsoletely crenulate, the margin recurving, coria-
ceous, glabrous, slightly lustrous above, paler beneath; lateral nerves
straight or little arcuate, marginally anastomosing, not or scarcely
reticulate, obsolete above, rather prominent beneath; female in-
florescence few-fascicled, puberulent-hirtellous, 1-flowered, the
pedicel 3-4 mm. long; calyx under drupe about 3.5 mm. across, the
subciliolate lobes rounded-obtuse; drupes globose, 4-5 mm. in
diameter, pyrenes 4 or 5, dorsally striate. Flowers according to
author 4-5-merous, otherwise unknown. F.M. Neg. 19556.
Peru(?): Without locality, Haenke, type.
Ilex andarensis Loes. Monogr. Aquifol. 1: 394. 1901.
Strongly affine 7. tarapotina according to the author but the
fleshier leaves typically less acuminate, nerves less prominent
beneath, inflorescences peduncled, calyx glabrous or nearly; young
branchlets as flowers in type somewhat pubescent; petioles 7-11
mm. long or sometimes apparently shorter, sulcate-margined; leaves
ovate to elliptic, entire margin narrowly recurved, obtuse or rounded
at base, acute or obtusely acuminate (acumen sometimes 7-12 mm.
long, in Peru short), 5-8 (13) cm. long, 2.5-5.5 cm. wide, coriaceous,
glabrous, scarcely lustrous, minutely brown-punctate beneath, the
7-10 lateral nerves either obvious or obsolete above, slightly promi-
nent beneath, sometimes reticulate; inflorescences typically pulveru-
lent, peduncles 2-5, pedicels 1-4 mm. long; flowers 4(5)-merous,
more or less hirtellous-puberulent, the calyx 1.5- barely 2 mm. wide,
the corolla 2.5-3 mm. long; anthers ovoid; ovary 4-5-celled; stigma
4-5-lobed. The forma psila Loes., I.e. 2: 289. 1908, has essentially
or quite glabrous branchlets and inflorescences, globose drupes 4-4.5
mm. thick, estriate and esulcate, the 4 pyrenes obsoletely 1-striolate,
hardly 3 mm. long. No material seen recently and the determination
of the San Martin specimens may be open to question. Illustrated,
Loes. I.e. pi 11, fig. 2. F.M. Neg. 13194.
San Martin: Tarapoto, Williams 5999. Zepelacio, King 3287;
3361. Loreto: Cerro de Escaler, Ule 6353. Canella Ucsha, Ule
6718. Cerro de Usco, Ule. Ecuador.
Ilex andicola Loes. Monogr. Aquifol. 1: 161. 1901 and 2: 277.
1908.
Similar to /. crassifolia and /. crassifolioides; leaves 6-7.5 cm.
long, 2.5-3.5 cm. wide, rounded to acute, lateral nerves 8-10, im-
pressed or plane and obsolete; inflorescence of male flowers with
FLORA OF PERU 275
peduncles 10-18 mm. long, ultimate pedicels 1-2 mm. long, calyx
less than 2 mm. wide, glabrous except ciliolate obtusish lobes;
petals to 2.3 mm. long; female flowers about 7, peduncles 8-20 mm.
long, calyx 4-5-lobed, 1.5-2.25 mm. wide, the deltoid acute lobes
callous-apiculate; ovary depressed, stigma 4-lobed, 4-celled, cells
1-ovuled; drupes globose, 4.5 mm. thick, immature. Type from
Huaycani by Pearce. The Cuzco material could be referred to
7. cuzcoana if that is distinct. F.M. Neg. 13195.
Cuzco: Slender, 3-4 meters, rain-forest, Pillahuata, Vargas 3016.
Cedrobamba, 22 meters, Herrera 1564- Puno: Azalaya to Ichu-
bamba, Prov. Sandia, 1,600 meters, Weberbauer 1114 (det. Loesener);
278. Bolivia?
Ilex anonoides Loes. Monogr. Aquifol. 1: 164. 1901.
With many of the characters of the closely allied 7. crassifolia
and 7. crassifolioides but the much larger leaves prominently reticu-
late, especially beneath where very densely punctate and with 13-19
lateral nerves; petioles 14-25 mm. long; leaves broadly elliptic to
obovate, rounded both ends, 11.5-18 cm. long, 7.5-10.5 cm. wide,
more or less remotely and broadly serrulate, opaque and granulate
above, especially on the nerves; inflorescences glabrous, 4-7-flowered,
compressed peduncles 6-17 mm. long, pedicels none or to 1 mm.
long, the 4-merous flowers fasciculate apically; calyx to nearly 3 mm.
wide, the subrotund lobes to 2 mm. wide; female corolla scarcely
rotate, the firm ovate petals about 2 mm. long, the staminodia
shorter, anthers broadly cordiform; ovary 4-celled; ovary globose,
crowned with 4-lobed stigma, 5 mm. in diameter, with 4 smooth
pyrenes. Illustrated, Loes. pi. 2.
Puno: Tatanara, Lechler 2624, type.
Ilex caniensis Macbr. Field Mus. Bot. 8: 122. 1930.
Branches stout, short, the subangled branchlets, petioles (2-3
mm. long), and pedicels more or less densely hirtellous; leaves
crowded, elliptic or elliptic-oblong, sometimes obovate, rounded or
little narrowed at base, rounded-obtuse or mucronulately acute,
3-4.5 cm. long, 2-2.5 cm. wide, fleshy rigid-coriaceous, glabrous,
lustrous above, opaque and epunctate beneath, margin scarcely
revolute, serrulate to apex, the 5-6 lateral nerves little marked except
beneath where prominent with the reticulate venation; peduncles
1-3-flowered, to 2 mm. long; pedicels 1-2 mm. long; flowers 5-6-
merous, calyx 3 mm. wide, lobes rotund, obscurely ciliolate; petals
276 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
elliptic, 3-3.5 mm. long. Compared with I. uniflora Benth. and
I. scopulorum HBK. it differs from the former in pubescence and
in the leaves, which are longer in proportion to width, and also
from the latter, which is merely puberulent and has subrotund foliage.
The type was from a shrub 1.5-2 meters tall.
Huanuco: Near Cani, 3455, type.
Ilex crassifolia Hook. Icon. 2: pi. 149. 1837; 159.
Stout erect early sparsely pulverulent branchlets finally subterete
and 2.5-4.5 mm. thick; stipules deltoid, acute, to 1.5 mm. long;
petioles 8-11 mm. long, to 3.5 mm. wide, plane and striate above,
sulcate and decurrent-margined beneath; leaves broadly elliptic to
ovate-oblong, cuneate to obtuse or rounded at base, mostly rounded
and minutely apiculate at tip, usually 5-7.5 cm. long, 3-4.5 cm. wide,
adult glabrous, nearly opaque above, paler and punctate beneath,
nigrescent-apiculate denticulate margin recurved; lateral nerves
9-11, obsolete or little prominent above, spreading, mostly straight,
rather obvious beneath and often reticulate only at margin; inflores-
cences (including calyces) pulverulent, the male stoutly pedunculate
(peduncles to 7 mm. long), the 3-7 subsessile flowers fasciculate,
bracts ovate-lanceolate, the larger lower to 4 mm. long; flowers
4-5-merous, the calyx 4-4.5 mm. wide, the subequal ciliolate rounded
lobes to 3.5 mm. wide; petals connate below, oval, 4- nearly 5 mm.
long, 2-3 mm. wide, fleshy, papillose; anthers ovoid; pistil of male
flowers ovoid, 1.75 mm. long, about 1.5 mm. wide. Apparently a
variant of I. scopulorum HBK., 171, of Ecuador, with even heavier
leaves very narrowly recurved at margin if at all, the male inflores-
cences often many-flowered with pedicels finally 2-6 mm. long, female
flowers solitary. The species name may be untenable but there
perhaps are synonyms; a number of Andean shrubs of this group
appear to be very similar indeed.
Amazonas: Chachapoyas, Mathews, type.
Ilex crassifolioides Loes. Monogr. Aquifol. 1: 160. 1901.
In general like I. crassifolia but the leaf nerves obviously reticu-
late beneath and the female inflorescence dichotomously forked,
many-flowered, subumbelliform; stipules scarcely 0.5 mm. long;
petioles more slender, 7-14 mm. long; leaves obovate to broadly
oval, 7-10.5 cm. long, 3-6.5 cm. wide; calyx of female flowers 2.5
mm. wide; petals about 2 mm. long; ovary subovoid, crowned by
FLORA OF PERU 277
capitate stigma, mostly 4-celled. Type a small tree 6 meters tall.
F.M. Neg. 32565.
San Martin: Monte Campana near Tarapoto, 1,700 meters,
Spruce 4461, type.
Ilex cuzcoana Loes. Monogr. Aquifol. 2: 273. 1908.
Branchlets spreading or ascending, the older about 4 mm. thick,
the much slenderer tips sub velvety puberulent; stipules subulate-
filiform, scarcely 1 mm. long; petioles 3-6 mm. long, narrowly mar-
gined, only about 1 mm. thick; leaves ovate or oval to oblongish,
obtuse to subcuneate at base, shortly but obviously acuminate, 3.5-5
(6) cm. long, 1.5-2.4 cm. wide, serrulate margin obscurely or not at
all recurving, fleshy coriaceous, glabrous, lustrous above, opaque
beneath where densely and minutely brown punctulate; lateral nerves
7-10 clearly impressed above, reticulate marginally, little marked
or obsoletely beneath; inflorescence axillary or lateral, solitary,
usually furcate, peduncles 5 mm. long, intermediary axes 3 mm.
long, bracts barely 1 mm. long, unidenticulate, bractlets at base of
the 1.5-2 mm. long pedicels; flowers 4-merous, calyx about 2 mm.
wide, lobes deltoid, obtuse or rounded, ciliolate; petals ovate, 3 mm.
long, 2 mm. wide, subauriculately dilated at base, stamens little
shorter, the anthers ovoid-subcordiform; fruit and female flowers
unknown. Belongs to same group as I. crassifolia and rather near
to I. andicola Loes., I.e. 1: 161, with slightly larger leaves, somewhat
smaller flowers, probably not specifically distinct. Cf. also I.
boliviano, Britton, 154, the reticulation of the leaves typically more
prominent beneath. F.M. Neg. 13206.
Cuzco: Intihuantani to Torontoi, Prov. Convention, 2,100
meters, Weberbauer 5067, type; 245.
Ilex elliptica HBK. Nov. Gen. & Sp. 7: 70. 1824; 182.
Branchlets hirtellous, the older tardily glabrate; petioles 1-2 mm.
long; leaves elliptic or suboval, rounded both ends, 2-3.5 cm. long,
11-21 mm. wide, finely serrulate margin narrowly recurving, pul-
verulent and epunctate beneath, fleshy coriaceous, glabrous or
nearly above, the 5-6 lateral nerves there obsolete, rather prominent
beneath, not reticulate; male flowers unknown; female solitary in
the axils or on stems, 5-6-merous, pedicels 4 mm. long, medial
bractlets about 0.5 mm. long, hirtellous as the calyx, this 3.5- scarcely
5 mm. wide, the acutish lobes to 1.5 mm. long; petals oval, 4.5-5 mm.
long; ovary 5-celled, cells 1-ovuled; drupes subglobose, about 8 mm.
278 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
in diameter, esulcate and estriate, pyrenes 4, obsoletely trigonous.
Probably to be joined to I. Kunthiana Triana of Colombia with
smaller mostly 4-merous flowers (Loesener), calyx lobes obtuse,
petals about 2.5 mm. long. Maybe the Stork and Horton specimen
should go to /. caniensis Macbr., if that is distinct.
Cajamarca: Contumasa, Bonpland, type. Huancavelica: Near
Surcubamba, 3,000 meters, Stork & Horton 10380 (det. Standley,
I. caniensis?).
Ilex ericoides Loes. Monogr. Aquifol. 1: 176. 1901.
Resembles I. Lechleri and thus I. ovalis but leaves epunctate
beneath, strongly revolute, minutely serrulate, nerves impressed
above; villous pubescence of young branchlets subglandular; stipules
persisting, about 1 mm. long; petioles 1-2, rarely 2.5 mm. long;
leaves oval, doubtfully ovate, cuneate-rounded or cuneate-acute at
base, obtuse and mostly apiculate, 5-13 mm. long, 4-7 mm. wide,
even adult villosulous on midnerve beneath; petals of female flowers
oval, 2 mm. long; drupes broadly ellipsoid, 4.5 mm. long, 4 mm.
thick, persisting stigma obsoletely 4-lobed, pyrenes 4, finely striate
but esulcate. F.M. Neg. 13210.
Puno: Sachapata, Lechler 2591, type.
Ilex hippocrateoides HBK. Nov. Gen. & Sp. 7: 72. 1824; 155.
Glabrous tree, the new branchlets at most 2 mm. thick; petioles
about 6 mm. long, finally sulcate above; leaves broadly oblong to
oblong-lanceolate, acute both ends or obtusely acuminate at tip,
4-6 cm. long, 1.5-2.3 cm. wide, plane or nearly to the serrulate
margin, coriaceous, sublustrous above, the 8-10 larger lateral nerves
densely reticulate, the reticulation particularly conspicuous beneath
in the smaller; female flowers and fruit unknown or at least not recog-
nized; male inflorescences solitary at base of new wood or in leaf
axils, 3-7-flowered, glabrous, the peduncles 7-13 mm. long, pedicels
at tip 1.5-2.5 mm. long, bracts acute, the larger about 0.5 mm. long;
flowers 4-merous, calyx 2-2.5 mm. wide, the subsemiorbicular acute
or acuminate ciliolate lobes sometimes five; otherwise unknown.
F.M. Neg. 13217.
Peru?: Type from Peru according to Loesener but no locality
given by HBK.
Ilex inundata Poeppig ex Reiss. in Mart. Fl. Bras. 11, pt. 1:
43. pi. 11, fig. 5. 1861; 403.
FLORA OF PERU 279
Glabrous, the striate branchlets sparsely and minutely lenticel-
late, finally 3-5 mm. thick; petioles 6-12 mm. long, sulcate, little
margined by the decurrent leaf -base, this acute or obtuse; leaves
about oval or rarely subobovate, entire, mostly obtusely acuminate
(acumen to 12 mm. long), rarely obtuse, 7-15 cm. long, 3-6.5 cm.
wide, coriaceous or subchartaceous, not punctate; lateral nerves
7-10, straight or mostly arcuate toward the margin, laxly reticulate,
moderately prominent only beneath; inflorescences about 5-10-
fasciculate, the male often dichotomous, peduncles and pedicels
short (1 mm. long), the female 1-flowered, peduncles and pedicels
5-7 mm. long; flowers 4-merous, female calyx 2-2.25 mm. wide, lobes
ciliolate, petals to 3 mm. long, 1-2 mm. wide; anthers cordate, sub-
sagittate; stigma capitate, scarcely lobed; ovary 4-celled. F.M.
Neg. 13219.
Loreto: Near Iquitos, Ducke 1915. Florida, Klug 231 4- Mishu-
yacu, Klug 426; 427; 456. Amazonian Brazil.
Ilex Jelskii Zahlbr. in Szyszyl. Diagn. PI. Nov. 1: 9. 1894;
Dissert. Classis Math. Phys. Acad. Litt. Cracov. 29: 223. 1895;
413.
Branchlets straight, 4 mm. thick, apically pubescent; stipules
subulate, about 0.5 mm. long; petioles 8-13 mm. long, tip nearly
2 mm. thick; leaves oval or elliptic to ovate-elliptic, cuneate-obtuse
to subrounded at base, shortly and acutely acuminate or acute,
5.5-8 cm. long, 2-3.5 cm. wide, recurving margin entire, rigid-cori-
aceous, densely puberulent, especially beneath, notably lustrous
above; lateral nerves 9-10, little prominent, very slender, anastomos-
ing obsoletely near margin; male flowers unknown, the female 4-5-
merous, 5-10 in leaf axils, peduncles 6-10 mm. long, pedicels 2-4 mm.
long; calyx 2-2.5 mm. broad, pubescent lobes deltoid; petals oval,
slightly pilose without, to 2.5 mm. long; ovary 4-5-celled, the cells
1-ovuled; drupes ovoid, gray sericeous, pyrenes 4. F.M. Neg. 13220.
Cajamarca: Tambillo, Jelski 37, type.
Ilex Lechleri Loes. Monogr. Aquifol. 1: 177. 1901.
Similar to I. ovalis but the new branchlets villosulous-hirtellous,
the older to 4 mm. thick; stipules nearly 1 mm. long; petioles more
slender; leaves oval, or ovate or oblong-oval, crenulate-serration less
prominent, mostly 1.5(-2) cm. long, 6-11, mostly 9 mm. wide,
peduncles and pedicels slenderer, more pilose or villous, flowers
smaller. In view of the other similar plants described as distinct
280 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
species these differences can scarcely be valid; however, female
flowers unknown. F.M. Neg. 23192.
Puno: Sachapata, Lechler 2607, type.
Ilex loretoica Loes. Repert. Nov. Sp. 1: 166. 1905; Monogr.
Aquifol. 2: 288. 1908.
Glabrous, the young shoots 2-3 mm. thick, the year's 3.5-6 mm.
and the older branchlets to 9 mm. thick, early lenticellate and finally
sulcate; stipules soon absent; internodes 5-35 mm. long; petioles
2-6 mm. long, stout; leaves elliptic to obovate or oval, apparently
obtuse both ends or rounded at tip (mostly imperfect in type),
12-18.5 cm. long, 5-9 cm. wide, finally rigid-coriaceous, paler beneath
where minutely brownish puncticulate, the 9-12 lateral nerves
beneath rather prominent as the reticulation, this obsolete above;
inflorescences many in axils or often some extra-axillary, the male
mostly 3-flowered, the slender peduncles 9-16 mm. long; bracts
broadly deltoid, 3-dentate, about 0.25 mm. long, bractlets at base
of the 1-2 mm. long pedicels; flowers 4-merous, calyx explanate,
2.5-3 mm. in diameter, lobes hardly 2 mm. long and barely to 2 mm.
wide at base, obscurely repand-denticulate; corolla subrotate with
ellipsoid sublinguliform petals, connate or subfree, about 3.5 mm.
long, the stamens nearly as long; pistil depressed, lobulate; female
flowers and fruit unknown. Next to I. nervosa Triana of Colombia,
which differs especially in indument (Loesener). It also closely
resembles I. laurina HBK., 360, country unknown, probably Ecuador,
Colombia or Venezuela (Loesener) but that species has rather lanceo-
late acuminate leaves 4-5.5 cm. wide.
San Martin: East of Moyobamba, 1,500 meters, Weberbauer 4749,
type; 267.
Ilex Matthewsii Loes. Monogr. Aquifol. 1: 176. 1901.
Erect, densely leafy branchlets pulverulent-hirtellous to 3.5 mm.
thick; stipules subulate-filiform to nearly 1 mm. long; petioles 3-5
mm. long, decurrently margined to 1.75 mm. thick; leaves broadly
oval to subobovate, obtuse or sub-rounded at base, rounded and
minutely black apiculate at tip, 1.5 to 3 cm. long, 8-16 mm. wide,
the narrowly recurved margin densely serrulate, fleshy-coriaceous,
glabrous except sparsely pilosulous on midnerve beneath and above
at margin, especially the younger, paler beneath and densely brown
punctulate; lateral nerves 4-7, marginally anastomosing and reticu-
late, rather prominent on both sides; inflorescence solitary in leaf
FLORA OF PERU 281
axils or rarely at base of new shoots, pulverulent-hirtellous, the male
3-7-flowered, peduncles 5-10 mm. long, pedicels 1-2 mm. long,
the basal bracts and bractlets subulate-deltoid; male flowers 4-
merous, the calyx 2.5-3 mm. wide, the broadly deltoid rounded
or obtuse ciliolate lobes scarcely 1 mm. long; petals suborbicular,
about 2.5 mm. long and wide; anthers broadly ellipsoid; otherwise
unknown. Near I. ovalis with viscid branchlets, shorter slender
pedicels and more slender lateral nerves obsolete above in the adult
leaves. F.M. Neg. 23193.
Amazonas: Chachapoyas, Matthews, type.
Ilex microsticta Loes. Repert. Nov. Sp. 1: 164. 1905.
Glabrous, unless the bracts, the soon densely lenticellate branch-
lets to 6 mm. thick, the leaf internodes 1-2 cm. long; petioles 8-13
mm. long, subplane above, striate- and subplicate-angulate beneath;
leaves obovate to elliptic, rarely subovate, cuneate at entire base,
serrulate to rounded tip, (5) 7-10.5 (13) cm. long, (2.4) 3.6-5.5 cm.
wide, rigid-coriaceous, densely and minutely punctate beneath, the
7-10 lateral nerves and reticulation there rather prominent, the
latter obsolete above; female inflorescences solitary in the axils or
often at base of lateral shoots before the leaves, mostly about
4-dichotomous and on peduncles 1-2 cm. long, these applanate and
bicarinate, the spreading somewhat carinate bracts about 1.5 mm.
long, ciliolate or sparsely pilose; pedicels 2-3 mm. long; flowers
4-merous; calyx quadrangular-subcupuliform, 2.5-3 mm. wide, tube
subequaling the acuminate lobes, these about 1 mm. long; male
flowers unknown; female rotate, petals little oblique, about 3.25 mm.
long, stamens much shorter; ovary 4-celled with fleshy capitate
4-sulcate stigma. Would be given number 62a in monograph;
suggests /. teratopis Loes. but has smaller flowers and mostly smaller
leaves, especially, 1-ovuled cells; I. crassifolia Hook, pulverulent
inflorescence, sepals acutish (author). F.M. Neg. 13229.
Hudnuco: Monzon, 2,000 meters, Weberbauer 3532, type; 255.
Ilex minutifolia Macbr., nom. nov. I. microphylla Hook. Icon.
PI. 2: pi. 139. 1837; 174, not Spreng. ex DC., 1825.
Low much-branched shrub, with many mostly spreading straight
branchlets, the striately angled younger densely puberulent, finally
glabrate, subterete, to 4 mm. thick; stipules subulate, scarcely
0.5 mm. long; petioles 0.75-1.5 mm. long; leaves suborbicular-
282 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
ovate or oval, obtuse, rounded or rarely acute, often minutely
apiculate, 7-11 mm. long, 5-7 mm. wide, coriaceous and ridged,
the margin nearly plane, slightly lustrous, scarcely paler beneath
where minutely puberulent on the midnerve and black punctate;
lateral nerves 4-5, inconspicuous above, often less beneath, anasto-
mosing at the margin; inflorescence solitary in leaf axils, the male
3- the female 1-flowered, minutely hirtellous; bracts 3-dentate,
linear, at base of 3-flowered pedicel, at middle of 1-flowered pedicel;
male peduncles about 3 mm. long, pedicels scarcely 3 mm. long,
the female to 3 or in fruit to 4 mm. long; flowers 4-merous; calyx to
2 mm. wide, puberulent, the obtuse or acute triangular lobes ciliate;
corolla about 2.5 mm. long and broad or in the female flower narrower;
anthers broadly ellipsoid; ovary 4-celled, the cells 1-ovuled; drupes
subglobose, the 4-lobed stigma prominent, about 4.5 mm. thick, the
4 pyrenes scarcely 3 mm. long, dorsally convex, striate-sulcate.
F.M. Neg. 23194.
Ayacucho: Guancas, (Fielding 1636). Amazonas: Chachapoyas,
Matthews 1636, type?
Ilex ovalis (R. & P.) Loes. in Engl. & Prantl, Pflanzenfam.
Nachtr. 1: 219. 1897; 179. Paltoria ovalis R. & P. Fl. Peruv. 1: 54.
1798. I. Paltoria Pers. Syn. 1: 152. 1805.
Much-branched shrub, the many straight erect branchlets to
6 mm. thick, subviscid, the shoots also pilose-puberulent; stipules
subulate, about 0.5 mm. long, petioles 2-2.5 (3) mm. long, decur-
rently margined, to scarcely 1 mm. thick; leaves oval, suborbicular
or rarely ovate, obtuse or acute at base, rounded and obscurely
black apiculate at apex, 1-2.6 cm. long, 7-16 mm. wide, rigid-
coriaceous, lustrous above, paler and black punctulate beneath, the
plane or nearly plane margin crenulate-serrulate; midnerve prominent
beneath, the 3-6 lateral slender and anastomosing at margin, obsolete
above, more or less conspicuous beneath; inflorescence solitary in
leaf axils, hirtellous or nearly glabrous, the male 3-flowered, peduncles
3-4 mm. long, pedicels scarcely 1 mm. long, bracts ovate-acuminate,
1-2-dentate, about 0.5 mm. long, the smaller bractlets at base
acuminate, in female in middle of pedicels; flowers 4-merous or calyx
sometimes 5-lobed, in female flower 2-2.5 mm. broad, the lobes
subrotund; corolla (female) about 2.5 mm. long to 3 mm. broad;
stigma obsoletely 4-lobed, the ovary 4-celled, the cells 1-ovuled.
The male flowers are undeveloped in type and the female, according
to the monographer, may not belong to the species. Probably should
be interpreted to include as variants the several similar plants.
FLORA OF PERU 283
Junin: Churupallana and Siusa, Ruiz & Pavdn, type. Chancha-
mayo, Isern 2300. Huanuco: Chacahuassi and Huacahuassi, Ruiz
& Pavon. "Quinti," "quhioi."
Ilex quitensis (Willd.) Loes. Monogr. Aquifol. 1: 221. 1901.
Rhamnus quitensis Willd. ex Roem. & Schult. Syst. 5: 295. 1819.
I. bumelioides HBK. Nov. Gen. & Sp. 7: 71. 1824.
Branchlets ascending, early striate sulcate, the gray-barked
older 2.5-3.5 mm. thick, the usually pulverulent densely leafy
shoots about 1.5 mm. thick; stipules subulate; petioles 4-6 mm.
long, margined; leaves erect, obovate to elliptic or oval-oblong,
entire or some closely serrulate, acute or cuneate at base, rounded
or subobtuse, minutely mucronulate, mostly 2.5 (1.6-4) cm. long,
8-14 mm. wide, coriaceous, lustrous above; midnerve slender above,
prominent beneath as the 4-5 (8) lateral nerves (these obsolete
above) nearly straight, marginally minutely reticulate; inflorescences
fasciculate in the leaf axils or the female rarely solitary on new
branchlets, all glabrous or pulverulent, the male dichotomously
3-flowered or rarely 4-5-flowered, peduncles 1-4 mm. long, bracts
minute, deltoid, pedicels 1-2 mm. long, basally bibracteolate;
female inflorescences 1-flowered, pedicels 3-7 mm. long; flowers
mostly 4-merous; calyx 2 or in female flower and under drupe to
3 mm. wide, the obtuse lobes to 1 mm. wide at base, to 0.75 mm.
long; male corolla rotate, the oval petals 2.5 mm. long, in the female
ovate and free or nearly; stamens about one-fifth shorter than male
petals, or the almost free or free staminodia one-sixth shorter;
ovary obtuse, conical, the stout stigma obsoletely 4-lobed; drupes
obtusely angled, about 5 mm. long, 3.5 mm. broad, with 4-5 dorsally
sulcate pyrenes. Description of flowers after Loesener, Repert.
Nov. Sp. 1: 166. 1905. Forma glabra Loes. I.e. has glabrous shoots.
F.M. Neg. 13241.
Amazonas: Molinopampa to Chachapoyas, 2,000 meters, Weber-
bauer 4329; 4356; 263. Ecuador, Guiana.
Ilex sessiliflora Triana, Ann. Sci. Nat. ser. 5. 16: 378. 1872;
166.
Distinctly pilose, especially the branchlet tips, inflorescences,
these solitary, and the leaves beneath; older branchlets to 4 mm.
thick, sulcate, glabrescent; stipules subulate-filiform, about 1 mm.
long, persisting; petioles 4-7 mm. long, narrowly margined, dilated
at base; leaves ovate or ovate-elliptic, obtuse at base, rounded or
284 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
obtuse, rarely acute at tip, nigrescent-apiculate, 3-5.2 cm. long, 13-
29 mm. wide, the densely serrulate margin often revolute or recurved,
rigid-coriaceous, lustrous or opaque above where sparsely villous, at
least on impressed midnerve and nerves, these 5-7 (-9), prominent
particularly beneath as the reticulate veins; inflorescences simple or
dichotomous, 3-7-flowered or the female usually 1-3-flowered,
peduncles 6-23 mm. long, bracts deltoid, acute or acuminate,
pedicels 1.5-2.5 mm. long or the female flowers subsessile; flowers
4-merous, calyx more or less villous, the male about 1.5 mm. wide
with obtuse or acute ovate-deltoid lobes scarcely 1 mm. long, the
female 3-4.5 mm. wide, with rounded ciliolate lobes 1.5 mm. long;
corolla subrotate, petals elliptic to obovate, male to 2.5 mm. long,
female 3.5-5 mm. long; ovary truncate-subellipsoid, 4-celled; drupes
(immature) 6 mm. long, 5 mm. thick, with 4 pyrenes. The var.
Pearcei Loes., I.e. 168, has leaves sparsely villous or glabrate, petals
of female flowers about 3.5 mm. long, calyx lobes acute or acuminate.
F.M. Neg. 13248.
San Martin: Road to Vitoc, Ruiz & Pavdn. Puno: Sachapata,
Lechler 2588; 2589. Bolivia; Colombia.
Ilex tarapotina Loes. Monogr. Aquifol. 1: 393. 1901.
Glabrous unless the minutely pulverulent young branchlets and
the male inflorescences, these mostly fasciculate in the leaf axils;
stipules subulate, about 1.25 mm. long; petioles 8-12 mm. long,
angled and rugulose beneath, 1-1.75 mm. thick; leaves ovate-
oblong to sublanceolate, cuneate to obtuse at base, gradually acumi-
nate, 8.5-10.5 (13) cm. long, 2.5-3.5 (5) cm. wide, entire margin
recurved, fleshy-coriaceous, very lustrous above, paler and minutely
punctulate beneath; lateral nerves 8-10, anastomosing only at
margin, obsolete above as sometimes beneath; flowers 3-7 (male),
4-5-merous, peduncles 5-11 mm. long, pedicels 1-2 mm. long, bracts
less than 1 mm. long; calyx to nearly 2 mm. wide, glabrous or nearly
in type, the subulate deltoid lobes scarcely ciliolate; petals hardly
3 mm. long; anthers ovoid or ellipsoid. Here would be sought
I. laurina HBK., 360, origin unknown but probably from north
of Peru (Loesener), the leaves minutely serrulate, inflorescences
glabrous. Illustrated, Loes. I.e. pi. 10, fig. 1. F.M. Neg. 32570.
The leaves furnish a very good tea or "mate," according to Ule.
San Martin: Tarapoto, Spruce 4399, type. Junin: Enefias,
Pichis Trail, Killip & Smith 25685.
FLORA OF PERU 285
Ilex teratopis Loes. Monogr. Aquifol. 1: 93. 1901.
Glabrous except for the puberulent many-flowered (male)
axillary panicles of 4-merous fragrant flowers; branchlets stout, even
the annual 4.5-6 mm. thick; stipules persisting, deltoid, acute, to
2 mm. long; petioles margined, plicate-sulcate, 10-16 mm. long, 2-3.5
mm. thick; leaves obovate or broadly oval, subcuneate or rounded
at base, rounded at tip, about 9-15.5 cm. long, 5.5-8 cm. wide, fleshy
and rigid-coriaceous, margin serrulate, recurving, slightly lustrous
above, paler and densely punctate beneath, the midnerve bistriate
above, multi-striate and sulcate beneath, there the 12-15 lateral
nerves prominent and reticulate, little so above; female flowers 6 or 7,
the drupes fasciculate-congested at tip of 1.5-2.5 cm. long peduncle,
this to 4 cm. long in the male panicles, their calyces 3-5 mm. wide
with deltoid obtuse lobes 1.25-1.75 mm. long; petals to nearly 5 mm.
long, the shorter stamens with ellipsoid anthers; drupes globose or
depressed rugulose with prominent evidently 4-lobed stigma, mostly
7-8 mm. thick, with 4 biovulate pyrenes. Type of the subgenus
Yrbonia, unique by its biovuled ovary cells and its many-flowered
long-peduncled panicles of male flowers. Type a 3-7-meter tree
from Huaycani by Pearce. Illustrated, Loes. I.e. pi. 1, fig. 2.
Huanuco: Rio Pozuzo, Weberbauer 6780 (det. Dahlem). Pan
de Azucar, Sawada 62. Puno: Between Sandia and Chunchusmayo,
Cachicachi, 2,000 meters, Weberbauer 1313 (det. Loesener); 242.
Bolivia?
Ilex trachyphylla Loes. Monogr. Aquifol. 2: 277. 1908.
Branchlets, these 4-5 mm. thick, axillary solitary inflorescences
and leaves shortly scabrous hirtellous, the latter especially asperous
above and hirtellous beneath on the nerves; petioles 2-4 (6) mm.
long, about 3 mm. wide, sulcate above, angled and hirtellous beneath;
leaves ovate, oval or obovate, rounded both ends or sometimes
excised, 5.5-9 cm. long, 3.3-6.5 cm. wide, obscurely and remotely
if at all serrulate, rigid-coriaceous, lustrous above, less so and indis-
tinctly brunnescent-punctate beneath, the 7-9 lateral nerves and
reticulate venation there prominent; peduncle (female flowers)
2-3 cm. long in fruit, subdivaricate secondary axis 2-3 mm. long,
bracts to 5 mm. long, pedicels 1-1.5 mm. long, the 5-6-merous
flowers apically congested; calyx 5-6 mm. wide, nearly as broad as
the depressed globose drupe, this with 5-6 pyrenes that are obsoletely
few-striate dorsally.
Cajamarca: Shanyn, Jelski 1176, type.
286 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Ilex villosula Loes. Monogr. Aquifol. 1: 366. 1901 and 2: 288.
1908.
Branchlets and leaves beneath villous, the former to 4 mm. thick,
the minute stipules hidden in the pubescence; petioles 3-4 mm. long;
leaves usually elliptic to obovate, commonly cuneate-acute at base,
rounded and often emarginate, 3-4.5 cm. long, 1.5-2.2 cm. wide,
entire margin revolute, fleshy and rigid-coriaceous, sparsely short-
hirtellous above; lateral nerves 4-7, impressed above, prominent
beneath, reticulate both sides; inflorescence (female flower) usually
single at base of shoots, sometimes fasciculate in leaf axils (male
not seen), 1-flowered, hirtellous, pedicels 4-5 mm. long; flowers
4-5-merous, calyx about 1.25 mm. wide, lobes acute, petals less
than 2 mm. long; ovary 4-5-celled, the stigma 4-5-lobed; male
flowers 4-merous, 1-5, ordinarily fasciculate in leaf axils, hirtellous,
peduncles 2-5 mm. long, pedicels about 1 mm. long or, when only
one flower, to 4 mm. long, bractlets subulate at base of pedicels,
calyx lobes broader than long, shortly acuminate, barely 0.5 mm.
long, petals nearly free, oval or ovate, 1.5- nearly 2 mm. long,
about 1 mm. wide, the stamens, their anthers ovoid, about as long.
Sometimes a tree to 8 meters tall (Weberbauer). F.M. Negs. 13265;
23198.
Amazonas(?): Yambrasbamba, Mathews 1565, type. Libertad:
Valley of the Mixiollo, 2,200 meters, Weberbauer 7043. Huanuco:
Monzon, Prov. Huamalies, 1,700 meters, Weberbauer 3560. Puno:
Cachicachi, 1,900 meters, Prov. Sandia, Weberbauer 1155 (det.
Loesener).
Ilex Weberbaueri Loes. Repert. Nov. Sp. 1: 165. 1905.
Densely branched, 1 meter tall, the somewhat angled pulverulent
young shoots to 1 mm. thick, the older subterete with sordid gray
sulcate bark and at third year 3.5 mm. thick; leaf internodes to 7 mm.
long; petioles 1.5-3 mm. long, barely 1 mm. broad; stipules minute,
subulate; leaves ovate-oblong to obovate-oblong, rarely ovate-oval,
or obovate, mostly cuneate at base, rounded or obtuse at apex and
then often apiculate, sometimes subacute, 7-14 (-17) mm. long, 3-9
mm. wide, rigid-coriaceous, glabrous, 2-4-crenulate-serrulate, obvi-
ously punctate and 3-4-nerved beneath or the nerves obsolete, there
as above the reticulation inconspicuous; male inflorescences 1-
flowered, solitary in leaf axils, glabrous or essentially, pedicels 3-4
mm. long, medially bibracteolate; flowers 4-merous, calyx patelli-
form, 2-3 mm. in diameter, erect lobes longer than tube, acuminate,
FLORA OF PERU 287
subsetulose or punctiform pubescent, 1.3 mm. long, 1 mm. wide
at base; corolla subrotate, petals broadly obovate, 3 mm. long, 2 mm.
wide; pistil (male flower) subcylindric, the beak 4-sulcate. A meter
tall. Near /. ovalis (R. & P.) Loes. with pilose branchlets, 3-flowered
male inflorescences; other relatives different and Brazilian. F.M.
Neg. 13261.
Junin: Near Huacapistana, 3,000 meters, Weberbauer 2091, type;
251.
CYRILLACEAE. Cyrilla Family
Erect shrubs or small trees with alternate estipulate entire leaves
crowded toward the tips of the terete branches and actinomorphic
flowers in axillary or terminal simple or panicled racemes. Calyx
valvate and 5-parted or with 5 equal or very unequal persistent
sepals. Petals 5, more or less cohering at base, imbricate or con-
torted. Stamens 5 or 10, inserted on receptacle, the free filaments
subulate, the 2-celled anthers finally longitudinally dehiscent. Disk
annulate, cylindric or subcupulate, confluent with base of 2-4(5)-
celled ovary. Style developed or obsolete, stigma punctiform or
acutely lobed. Ovule solitary on inner angle or pendulous, then
several. Fruit capsular or drupaceous, sometimes winged, more or
less subtended by calyx; seeds 1-4 with endosperm, the straight
cylindric embryo small.
PURDIAEA Planchon
Reference: Fr. Mattick, Notizbl. Bot. Gart. Berlin 12: 395-397.
1935.
Costaea Richard, 1853. Alloiosepalum Gilg, Notizbl. Bot. Gart.
Berlin 11: 97-99. 1931.
Leaves sessile. Racemes few-flowered. Sepals unequal, often
conspicuously. Petals ovate to obovate. Stamens 10, apiculate.
Ovary globose. The single Peruvian species conforms closely to
the family character but it is striking by its strongly unequal veiny
sepals; Gilg placed it, with query, in the Ochnaceae, a not unreason-
able error discovered by Mildbraed.
Purdiaea Weberbaueri (Gilg) Mattick, Notizbl. Bot. Gart.
Berlin 12: 395, 398. 1935. Alloiosepalum Weberbaueri Gilg, Notizbl.
Bot. Gart. Berlin 11: 97. 1931.
Glabrous shrub, the bark of the younger branches grayish-black;
leaves obovate-oblong to oval, gradually narrowed to base, acute or
288 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
shortly apiculate, 3-4 cm. long, 1-1.5 cm. wide, coriaceous, opaque
both sides, the slender reticulation prominent only beneath; racemes
3-4 cm. long; bracts linear, 1 mm. wide, 5-7 mm. long; outer sepals
broadly ovate, rounded apically, obviously nervose, about 1 cm. long,
7-8 mm. wide, the lanceolate acute inner 6-7 mm. long, 2 mm. wide,
marginally ciliolate; petals reddish-lilac, 7-8 mm. long, 4-5 mm.
wide, ovate or broadly ovate, apiculate; filaments about 3 mm. long;
anthers 4 mm. long; ovary 2 mm. in diameter; style 5 mm. long;
stigma punctiform. Type was a 2-meter shrub. According to
Mattick, who gives a key to the known species, the Peruvian shrub
is distinguishable by a combination of leaf-size, completely glabrous
racemes and subequal sepals and petals, but apparently the author
himself had decided that it was referable to P. nutans Planchon of
Colombia, probably a too conservative interpretation unless future
collections show intergradation. Illustrated, Gilg, I.e. 98, fig. 3.
The data for this compilation were kindly supplied by "Roxy"
Ferris and Barbara Law of the Dudley Herbarium to whose staff,
it may be recorded, I am so often indebted.
Amazonas: In woods, Molinopampa between Chachapoyas and
Moyobamba, 2,100 meters, Weberbauer .477-4, type; also
This number of the Flora of Peru was in proof before I saw a
copy of Pflanzenfamilien ed. 2. 20b, which pertains to several of
the families treated. Some of my judgments might have been
altered and some probable errors and omissions avoided if this
excellent source book had been available during the preparation
of my account. However, I have been able in a few instances to
insert data from it, thanks to the prompt loan by Marjorie W. Stone,
Bibliographer of the Gray Herbarium, of their copy, which was
sent directly to Stanford at the kind suggestion of Professor G. F.
Ferris, then in charge of the Natural History Museum there.
INDEX
Synonyms in italic type
Acalypha, 121
Actinostemon, 197
Adenophaedra, 154
Alchornea, 146
Aleurites, 3
Alloiosepalum, 287
Anacardiaceae, 238
Anacardium, 240
Andrachne, 50
Angostylis, 118
Anthodon, 219
Aparisthmium, 150
Apodandra, 117
Aquifoliaceae, 270
Argithamnia, 92
Argothamnia, 92
Argyrothamnia, 92
Argythamnia, 92
Astrococcus, 119
Bernardia, 157
Briquetina, 227
Buxaceae, 220
Calatola, 222
Callitrichaceae, 235
Callitriche, 236
Caperonia, 96
Caryodendron, 153
Casimirella, 232
Celastraceae, 259
Cheiloclinium, 201
Chiropetalum, 94
Chonocentrum, 56
Citronella, 227
Cleidion, 155
Cnidoscolus, 163
Codiaeum, 3
Conceveiba, 152
Conceveibastrum, 151
Coriaria, 237
Coriariaceae, 237
Costaea, 287
Croton, 57
Cuervea, 216
Cunuria, 168
Cyrillaceae, 287
Dalechampia, 98
Discocarpus, 56
Discophora, 223
Ditaxis, 92
Dodecastigma, 176
Drypetes, 56
Elaeophora, 116
Eleutherostigma, 115
Emmotum, 225
Euphorbia, 8
Euphorbiaceae, 3
Haematostemon, 119
Haplorhus, 258
Hevea, 108
Hieronima, 51
Hieronyma, 51
Hippocratea, 216
Hippocrateaceae, 200
Huertea, 234
Humirianthera, 232
Hura, 199
Hyeronima, 51
Icacinaceae, 221
Ilex, 271
Jatropha, 158
Julianiaceae, 238
Julocroton, 57
Kippistia, 201
Kummeria, 223
Leretia, 230
Loxopterygium, 256
Mabea, 183
Mangifera, 241
Manihot, 168
Maprpunea, 198
Mauria, 245
Maytenus, 259
Megalostylis, 118
Nealchornea, 180
Omphalea, 179
Pausandra, 173
Pedilanthus, 34
Pera, 106
Peritassa, 204
Phyllanthus, 34
Pleurisanthes, 226
Plukenetia, 115
Pogonophora, 176
Poraqueiba, 229
Pristimera, 216
Purdiaea, 287
Rhacoma, 269
Rhus, 255
Richeria, 48
Ricinocarpus, 122
Ricinus, 114 Stillingia, 191
Styloceras, 220
Salacia, 208
Sapium, 192 Tapirira, 243
Schaefferia, 270 Tetrorchidium, 177
Schinopsis, 257 Thermophila, 208
Schinus, 251 Tontelea, 208
Sebestiania, 188 Tragia, 119
Securinega, 47 Turpinia, 233
Senefeldera, 181
Spondias, 242 Villaresia, 227
Staphyleaceae, 233 Villaresiopsis, 227
580. 5FB C001
FIELOIANA, BOTANYSCHICAGO
13:3-3A
30112009379048
