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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 2
DECEMBER 21, 1966
PUBUCATION 808
NAi
MIS
FLORA OF PERU
BY
J. Francis Macbride
CURATOR, PERUVIAN BOTANY
BOTANICAL SERIES
FIELD MUSEUM OF NATURAL HISTORY
VOLUME XIII, PART IIU, NUMBER 2
DECEMBER 21, 1966
PUBUCATION 808
tlfT i:3RARY OF TV--
PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS
FLORA OF PERU
J. Francis Macbride
SAPINDACEAE. Soapberry Family
Reference: L. Radlkofer, Sapindaceae, Pflanzenreich IV. 165.
1931-34.
Plants usually ligneous, sometimes trees, often lianas, less fre-
quently herbaceous vines, ordinarily with tendrils and alternate
commonly compound leaves, those of the climbing genera often
stipulate. Flowers pseudo-polygamous or unisexual, regular or
irregular, bracteate and bracteolate in cincinni disposed in racemi-
form or paniculate inflorescences, generally small with 4-5 (rarely
none) often more or less connate usually imbricate sepals and 3-5
(or none) imbricate petals that not rarely are squamate or barbate
within. Disk annular, regular or unilateral between the petals and
the stems, sometimes obscure. Stamens 8 (5-10 or more), the
filaments filiform, the 2-celled anthers usually introrse. Ovary
ordinarily 3-celled and 3-lobed, sometimes 2-celled, with simple or
divided styles and 1-2 or rarely more anatropous ovules in each cell
affixed to the axis. Fruits capsular or drupaceous, variously de-
hiscent or indehiscent, often consisting of 2-3 samaras. Seeds
without endosp)erm sometimes with a fleshy aril. — In the following
compilation the more generic term panicle has been used to include
reference to the frequently thyrsoid inflorescence in order to avoid
the awkward English plural of thyrsus. As defined by the mono-
grapher the species of some genera are highly critical; some, dis-
tinguished primarily by presence or absence of papillosity or hypo-
derma, are especially open to question as suggested by Radlkofer
himself who now and then qualifies the characters, if not in his key,
often in his descriptions; equally tenuous are a number of other
distinctions, particularly those pertaining to the leaves which pre-
sumably mark specific lines but more likely point up the variations
or instability or both of a few entities that maintain fairly constant
populations but produce plenty of aberrations. It is my feeling too
that the high proportion of incompletely known species explains
in part the difficult (from a practical standpoint) taxonomy, devised
largely by the monographer and notable as basically sound, clearly
organized and meticulously presented.
291
292 Field Museum of Natural History — Botany, Vol. XIII
The Litchi Tree (Litchi chinensis Sonnerat) indigenous to China
and one of the most ancient of cultivated plants is also one of the
best known members of this world-wide family because of its nut-
like subglobose bright red clustered fruits, their papery warty shells
drying brown and enclosing a pulp-surrounded seed. Perhaps also
in cultivation is the Rambutan (Nephelium lappaceum L.), similar
but the brittle shell of the fruit covered with long soft processes.
The allied but native South American Mamoncillo or Genip (Meli-
cocca hijuga L.) has granulate-shelled plum-shaped fruits, the single
seed (edible when roasted) enclosed in a sweetish gelatinous pulp.
All these trees are in general related to the Peruvian Talisias and
illustrated, Chicago Nat. Hist. Mus. Pop. Ser. 26, and elsewhere.
Probably also in cultivation but primarily only for ornament
is the African Akee (Blighia sapida Koenig), a great tree with
beautiful pinnate foliage and large almost bizarre fruits ripening
red and yellow and opening into three parts each filled with lustrous
white or yellowish tissue (cooked when ripe but dangerous unless
taken at exactly the right stage) surrounding the large black poison-
ous seed.
Scandent plants, often with tendrils.
Calyx with 4-5 imbricate sepals, the two upper connate or the
outer smaller; leaflets rarely only 3.
Fruit a samara, usually 3 together; stems often rigid-ligneous
even above 1. Serjania.
Fruit often alate dorsally, never samaroid.
Fruit firm, capsular; stems ligneous even above . . 2. Paullinia.
Fruit membranous; stems rather herbaceous above.
Fruits little inflated, alate; leaflets 3 3. Urvillea.
FVuits inflated, not alate; leaflets several.
4. Cardiospermum.
Calyx 5-parted or -lobed, the equal segments valvate or sub-
imbricate; leaflets always 3.
Calyx cupulate, the lobes subimbricate; flowers pseudo-um-
bellate 5. Thinouia.
Calyx lobes valvate; flowers pseudo-racemose. . .6. Allosanthus.
Erect shrubs or trees.
Leaves compound (sometimes by reduction 1-f oliolate) .
Leaves 3-foliolate or the lateral leaflets rarely wanting.
7. Allophylus.
Flora of Peru 293
Leaves pinnate (or leaflets only 3 but pinnately disposed in one
Talisia) or bipinnate.
Leaves bipinnate 12. Dilodendron.
Leaves pinnate.
Fruits rather samaroid, the cells inflated (Peru); sepals
rounded, tomentose; leaflets entire.
10. Porocystis. 9. ToiUicia.
Fruits not samaroid; petals normally 5 (rarely 4 or more
than 5).
Calyx divisions free, broadly imbricate.
Leaflets entire; fruits indehiscent, usually glabrous or
puberulent.
Fruits not granular, usually 3-coccic; sepals petal-
oid; leaf rachis often winged 8. Sapindus.
Fruits granular, usually 1-celled; sepals subcori-
aceous; rachis emarginate 11. Talisia.
Leaflets serrate, unless C. scrobiculata; fruits capsular,
dehiscent 13. Cupania.
Calyx cupulate, 5-lobed; leaflets entire (Peru); capsules
dehiscent 14. Maiayba.
Leaves mostly simple; fruit a 3-lobed capsule 15. Llagunoa.
Leaves all simple; fruit a winged capsule 16. Dodonaea.
1. SERJANIA [Plum.] Schum.
Shrubs scandent by axillary or p>eduncular tendrils, the leaves
usually bitemate, often pellucid punctate, the stipules minute or
none. Flowers irregular, rather small in axillary racemes or panicles
with 4-5 persisting sepals, 2 sometimes more or less united, outer
2 smaller, the inner petaloid, and 4 petals, the scales of the lower 2
apically appendaged. Disk glands 4 or the 2 smaller ones some-
times obscure. Style 3-lobed. Fruit 3 samaras, broadly winged
toward the base, united along the axis and with indehiscent seed
cells at the apex, the seed aril small. — Secondary wood often com-
posite (abnormal) with 3-5 or 8-10 additional wood columns.
The fruit, while partly diagnostic, is incompletely known for many
Peruvian species, and much herbarium material is only in flower;
the following key therefore has been devised to lead (usually without
fruiting character) perhaps at least to some of the more commonly
encountered species.
294 Field Museum of Natural History— Botany, Vol. XIII
Leaves all simply 5-foliolate, the lower pair of leaflets undivided, or
3-foliolate.
Leaflets 5.
Petioles and rachis wing-margined S. dibotrya.
Petioles and rachis emarginate.
Leaflets rounded-obtuse or -retuse; inflorescences narrow.
S. elongaia.
Leaflets usually with short obtuse acumen; inflorescences
often panicled S. grandifolia.
Leaflets 3 S. grammatophora.
Leaves more compound; rachis rarely wing-margined (S. deltoidea).
Leaves all or mostly with only one pair of ternate or subternate
leaflets or obviously pubescent as S. longistipula; see page 298.
Indument of leaves (at least one side) or of stems conspicuous,
dense, usually in age and flowers large (4.5-6 mm. long);
fruit cells (known) often more or less compressed.
Branches slender, long-hispid; leaflets entire; wood simple.
S. altissima.
Branches slender, strigillose or glabrate as the green mem-
branous leaflets; wood composite.
S. communis var., S. dumicola.
Branches stout, soon 4 (3) -6 mm. thick, usually, as the firm
leaflets beneath, densely short-tomentose or hirsutulous.
Indument of branchlets short-hirsutulous; stipules ovate;
leaves biternate (Peru) S. diffusa.
Indument of branchlets in type sparse, lax or in var, dense;
stipules linear-subulate, leaves imparipinnate.
S. longistipula.
Indument at least of branchlets closely tomentose; stipules
ovate or "lanceolate-subulate" (S. mollis); leaves
biternate.
Wood composite; epidermis typically lacking mucus.
Panicles solitary or the upper panicled.
Terminal leaflets long-petiolulate (type); pedicels
2 mm. long S. mollis.
Terminal leaflets shortly petiolulate or subsessile
(type); pedicels to 5 mm. long. .S. sufferuginea.
Panicles paniculately congested (imperfectly known);
leaflets all subsessile S. peruviana.
Flora of Peru 295
Wood simple; epidermis containing mucus (as to tjrpes;
cp. following with S. mollis).
Fruit cells alulate; branches scarcely striate (type).
S. aluligera.
Fruit cells merely callose; branches lightly sulcate
(type) S. caUigera.
Indument never conspicuous unless in inflorescence; fruit cells
(known) more or less inflated except S. communis, S. in-
scripta, S. leptocarpa.
Flowers small, 2-3 (3.5) mm. long; sepals equally puberulent
and wood composite unless in the last four species and
S. rubicaulis with exception of S. pyramidata.
Leaflets coarsely serrate medially or nearly to base.
Upper stems soon glabra te; leaflets serrate, often below
middle.
Serrations subduplicate, approximate from below the
middle S. tenuifolia.
Serrations simple, irregular, remote, mostly from
middle S. memhranacea.
Upper stems short-hirsutulous; leaflets serrate only
above middle S. rufa.
Leaflets entire, subentire or serrations obtuse, rounded,
remote, few, or stems usually prickly.
Branches deeply 5-sulcate, more or less prickly; wood
simple S. rubicaulis.
Branches striate or lightly 8-sulcate, smooth; wood com-
posite except as noted above.
Leaflets ample, even the lateral usually several cm.
wide.
Leaflets remotely obtusely serrate, often slightly
pubescent.
Serrations coarse; fruit wings diaphanous, cells
lightly nerved S. memhranacea.
Serrations small or obscure; fruit wings firm-
membranous, cells deeply nerved.
Leaflets acute or acutely acuminate; wood
simple S. exarata.
Leaflets obtusely short-acuminate; wood com-
posite S. pyramidata.
296 Field Museum of Natural History — Botany, Vol. XIII
Leaflets entire or bidentate near obtuse acumen,
quite glabrous S. grandifolia.
Leaflets small, even the terminal rarely wider than
3 cm.
Leaflets subequal, all remotely serrulate above the
middle S. trirostris.
Leaflets obviously unequal, all entire or subentire.
Lateral leaflets subrotund, 2,5 X 2 cm., all
a little puberulent S. subrotundifolia.
Lateral leaflets subelliptic, 10 X 4-5 mm., all
glabrous except nerves S. Killipii.
Flowers large (for genus) normally at least 4 mm. long; sepals
in part soon glabrate or greenish or the leaflets entire or
obtusely few-dentate; wood simple only in S. exarata.
Sepals equally cinereous or fulvous puberulent except S.
caracasana and possibly forms of S. communis; leaflets
often entire or obtusely few-dentate.
Leaflets oblong-lanceolate, usually at least three times
longer than wide S. lethalis.
Leaflets ovate- or oblong-elliptic, often about twice lon-
ger than wide.
Leaflets (lateral) obtuse or obtusely acuminate, quite
glabrous (Peru) as outer sepals . . . .S. caracasana.
Leaflets acute or acutish, often a little pubescent as
outer sepals.
Stem angles, at least in part, very acute or even
winged.
Leaves subcoriaceous, lustrous; petioles often in
part margined; outer wood bodies triangular.
S. lethalis, S. paucidentata.
Leaves submembranous, often dull; petioles
emarginate; outer wood bodies lamellate.
S. nutans.
Stems 5-6-striate-angled; branches (early) as in-
florescence reddish-puberulent; petioles in part
margined S. inscripta.
Sepals, at least in part, soon glabrate, greenish (unless S.
glahrata, S. exarata); leaflets, usually at least lateral,
acutely, angulately, irregularly or coarsely serrate
Flora of Peru 297
often below the middle; wood compKjsite except S.
exarata.
Leaflets, unless the terminal, obtuse, glabrous (Peru)
as outer sepals S. caracasana.
Leaflets usually acute and as the outer sepals most often
slightly puberulent at least on nerves.
Fruit cells subcompressed, cristate; petioles emargin-
ate; leaflets unequal or dissimilar.
Leaflets remotely few-dentate or subentire.
Flowers roseate; leaflets pilosulous beneath,
similar S. leptocarpa.
Flowers yellowish; leaflets glabrate, diverse (var).
S. inscripta.
Leaflets repand-dentate at least to middle.
S. Alsmitkii.
Fruit cells subglobose; lateral leaflets often smaller
than terminal but rarely diverse; petioles ob-
scurely or not margined.
Leaflets lanceolate to somewhat ovate-lanceolate, to
about 3 cm. wide; upper stems 2-3.5 mm. thick;
fruit cells lightly nerved.
Petioles emarginate; leaflets small, ovate-lance-
olate.
Third and fifth sepals connate; stipules about
ovate S. communis.
Third and fifth sepals free; stipules subulate.
S. dumicola.
Petioles in part margined; leaflets narrowed at
each end S. sphaerococca.
Leaflets usually broadly ovate, often several cm.
wide; upper stems soon 4-5 mm. thick; fruit
cells deeply nerved.
Wood composite; petioles emarginate; leaflets
ample S. glabrata.
Wood simple; petioles in part submarginate;
leaflets 3.5 cm. wide (type) S. exarata.
Fruit cells inflated; petioles in part margined; leaflets
dissimilar, the much smaller lateral often sub-
lobulate-dentate S. inflata.
298 Field Museum of Natural History — Botany, Vol. XIII
Leaves usually imparipinnate, all or mostly with more than 1 pair
of ternate or subtemate leaflets, their divisions often only
1-2 cm, wide (cf. also the pubescent S. longistipula and
a doubtful specimen of S. diffusa might be sought here).
Lower pair of leaflets mostly 5-foliolate or pinnate; wood
abnormal except S. striolata; flowers 3.5 (3) -6 mm. long,
except S. deltoidea, unopened in S. striolata.
Leaf rachis wing-margined; fruit cells distended . . S. deltoidea.
Leaf rachis obscurely or not margined; fruit cells subinflated.
Leaflet divisions mostly or all about 1 (-1.5) cm. wide;
flowers large.
Inflorescences subspiciform; peripheric wood columns 5;
northern species S. brachyptera.
Inflorescences rather lax; peripheric wood columns 1-2;
southern species S. platypetala, S. squarrosa.
Leaflet divisions mostly or all about 2 cm. wide; flower
buds small; wood normal S. striolata.
Lower pair of leaflets merely ternate; wood normal except S.
parvifolia; fruit cells more or less compressed.
Leaflet divisions mostly about 2 cm. wide; flowers 3.5 (3)
-6 mm. long.
Ultimate leaf divisions rather oblong-elliptic; young fruit
and sepals (type) glabrous or nearly . . S. fuscostriata.
Ultimate leaf divisions obliquely obovate; fruit and inner
sepals puberulent S. striata.
Leaflet divisions mostly 1-1.5 cm. wide; flowers 2-2.5 mm.
long.
Leaves glabrous or nearly, lacking mucus; wood normal.
S. oxyphylla.
Leaves barbate in axils and lightly pubescent; wood
abnormal S. parvifolia.
Serjania Alsmithii Macbr., sp. nov.
Scandens, fruticosa; rami 5-angulares inter angulos mediocriter
canaliculati, juniores undique, adultiores ad angulos hirtelli; corpus
lignosum minus durum, compositum e centrali magno pentagono
cavitate medullari sat larga percoso et periphericis parvis angulis
centralis superpositis; foliis bitematis; petiolis canaliculato-striatis;
foliolis minutissime denseque pellucido-punctatis chartaceo-mem-
Flora of Peru 299
branaceis utrinque glabris vel in nervis supra obscure sparseque
hirtellis paullo nitidulis subtus pallidioribus oblongo- vel ovato-
ellipticis acutis, mucronatis, basi plus minusve attenuatis, sub-
petiolulatis, repando-dentatis, plerumque 6-7 cm. longis, 3^ latis,
lateralibus paullo minoribus; thyrsi solitarii laxe cincinnigeri,
cincinni breviter stipitati, pedunculo rachique tenere puberulis,
pedicellis fructiferis circa 4 mm. longis; floribus vix notatis ut
videtur 4-4.5 nmi. longis, sepalis subglabris circa 3 mm. longis. —
Fructus fere ovatus, glaber, valde compressus 3 cm. longus infra
medium circa 3 cm. latus basi et apice vix excisi infra loculos vix
constricti, alis paullo dilatatis.
Distributed as S. tenuifolia which it resembles considerably in
foliage but apparently it is rather a member of another section of
the genus, probably section Platycoccus Radlk. at least sens. lat.
It accordingly requires another name for purpose of cataloguing
in this work and, if it does not prove to be extra-Peruvian may
honor in the same group with S. Killipii the famous "botanical
brothers," to use the happy term of Croizat.
Loreto: Subligneous vine; fruit pinkish red, Yurimaguas, Killip
& Smith 28301, type.
Serjania altissma (Poepp. & Endl.) Radlk. Monogr. Serj. 251.
1875: 72. Cardiospermum aUissimum Poepp. & Endl. Nov. (Jen.
& Sp. 3: 38. 1844.
Scandent, suffruticose, the deeply 5-sulcate-costate branches
3.5 mm. thick and somewhat pilose-glandular as also setulose with
rusty spreading trichomes 3-3.5 mm. long, this sort of pubescence
in some degree extending to the inflorescence (this more glandular)
and the leaves, these glabrate in age; ligneous structure simple,
notably costate, in cross section sinuate or crenate-lobed ; stipules
subulate-linear, to 6 mm. long, setose, the petioles sparsely so,
8 cm. long, the intermediate ones 5, the lateral 1.5 cm., all striate-
sulcate; leaves bitemate, often 22 cm. long, 20 cm. wide, mem-
branous, brownish, paler beneath, rather to very obscurely pellucid-
punctate, epidermis containing mucus, above sparsely or scarcely,
beneath early more or less densely puberulent especially on nerves
and somewhat ciliate margins with many sessile glands intermixed,
the terminal leaflet long-attenuate to the petiolule, broadly elliptic
or subrotund, subobovate, to 9 cm. long, 4.5-5 cm. wide, the lateral
smaller, all abruptly and acutely acuminate and entire or rarely
with a tooth or two toward the tips; panicles solitary or paniculately
300 Field Museum of Natural History— Botany, Vol. XIII
congested, longer than the leaves, the approximate cincinni on
stipes 5-15 mm. long, the rather crowded pedicels 3-3.5 mm. long;
flowers about 5 mm. long, white; outer sepals puberulent and
articulate-glandular, the longer inner very minutely puberulent;
petals broadly obovate, the scales half as long, the upper obovate,
the lower dentiform; stamens pilose at base; fruit (barely half
mature) setigerous. — The fruit described by the authors does not
belong here, according to the monographer; in Williams JtllS it
measures 3 by 2.5 cm. scarcely 1.5 cm. at the cells, these setose,
carinate by the decurrent wings, the pericarp fragile, seed black.
F. M. Neg. 5533.
San Martin: Tarapoto, Spruce J^560; U5W; Mathews 1322; 1322
bis. Zepelacio near Moyobamba, 12-1,600 meters, Klug 3596 (det.
Standley). Chazuta, Klug Ji-lO^ (det. Standley).— Loreto: Yuri-
maguas, Poeppig 2U15, type; Williams U113; Killip & Smith 29071.
Serjania aluligera Radlk. Bull. Herb. Boiss. 1: 464. 1893; 64.
Canescent scandent shrub with subterete scarcely striate branches
— ligneous area simple— and upper temate or 5-foliolate-pinnate
leaves, the lower bitemate; stipules minute, ovate; petioles 3-4 cm.
long, the intermediate 2 cm. long, the lateral 6 mm. long; leaves of
the flowering branchlets 10-13 cm. long, 9-10 cm. wide, the oval
leaflets serrate-dentate above the middle, mucronulate, the terminal
4-6 cm. long, 2-3.5 cm. wide, attenuate into a petiolule 5-8 mm.
long, the lateral gradually smaller, sessile, all pinnate-nerved, sub-
chartaceous, brownish -green, opaque, microscopically glandular,
puberulent above, softly ashy pubescent beneath, epunctate, the
epidermis containing mucus; thyrsi subcorymbose-racemiform,
solitary, divaricate, the flowering part half as long as the peduncle,
the cincinni spreading, long (to 1 cm.) -stiped; bracts and bractlets
subulate, long-glandular; pedicels about 4 mm. long,in fruit to 7 mm.,
articulate at lower third; male flowers with inner sepals 5 mm. long,
the outer half as long, all ashy puberulent; petals oblong-spatulate,
6 mm. long, 2 mm. wide, glandular within, the scales villous at the
margin, the upper crest obcordate-bifid and with long deflexed
barbate appendage, the lower obliquely emarginate aliform; stamens
lightly pilose, the anthers glabrous; young fruit elliptic, ashy pube-
scent, rusty villous at apex, the cells compressed at sides and with
aluliform processes near the dorsal crest, these hidden in the tomen-
tum, the endocarp glabrous except very sparsely pilose in the angles.
F.M. Neg. 5964.
Cajamarca: Callacate, Jelski J^09, type.
Flora of Peru 301
Serjania brachyptera Radlk. Bot. Jahrb. 37: 145. 1905; 63.
Like S. squarrosa but the stems 8-angulate and particularly the
ligneous peripheral areas 5, the spreading panicles 7-26 cm. long
with only 1 cm. long about 5-flowered cincinni, the fruit cells gla-
brous or sometimes floccose within; petioles 5-10 mm. long; leaves
5-6 cm. long and broad with 3-4 pairs of leaflets, the lower temate
or pinnate, trichomes few; outer sepals ashy hirsute; fruit not mature,
subquadrate, glabrous, 1.5 cm. long, at base 1.2-1.4 cm. broad. — It
is possible that my 2291, incomplete, may belong here. F.M.
Neg. 5534.
Cajaniarca: Hacienda La Tahona near Hualgayoc, 2,600 meters,
Weberbauer ^057. — Hudnuco: Among herbs and shrubs, 3,200
meters, near Punchau, Prov. Huamalies, Weberbauer SSOIt, type.
Serjania calligera Radlk. Bot. Jahrb. 54: Beibl. 117: 42.
1916; 65.
Much like S. aluligera but the sordidly tomentulose tumid fruit
cells laterally entirely free scarcely cristate but at the lateral angles
above sometimes with a produced callus; branches 4-5-sulcate;
petioles 5-10 mm. long, the intermediate 2.5 cm., the lateral 3 mm.
long, all more or less rusty tomentulose as the leaves, the larger with
terminal leaflet 6.5 X 4 cm.; cincinni approximate on stipes only
2-3 mm. long, the pedicels 2-3 or in fruit 5 mm. long, articulate near
base; petals white, 6 mm. long, 3 mm. wide; ovary densely tomentose,
the cells villous within, the glabrous style trifid at apex; immature
fruit to 1.8 cm. long, 1.4 cm. broad, at the cells 8-9 mm. — Apparently
nearly S. mollis but the leaves smaller (Dahlem note) and certainly
doubtfully distinct from S. aluligera.
Cajamarca: Between Huambos and Mont^n, 2,500 meters,
Weberbau£r J^15, type. — Piura: Hacienda San Antonio, 1,300 meters,
Weberbauer 6011.
Serjania caracasana (Jacq.) Willd. Sp. PI. 2, pt. 1: 465. 1799;
91. PauUinia caracasana Jacq. Hort. Schoenbr. 1: 52. pi. 99. 1797.
Typically nearly glabrous even to the outer sepals, only the
branches (apically) and solitary or paniculate inflorescences micro-
scopically glandular and minutely pubescent, the former lightly
6-8-striate, 2-4 mm. thick; wood composite, peripheral areas often
about 8; stipules broadly triangular, 2 mm. long, puberulent, as the
6-12 cm. long emarginate petioles, especially at the articulation;
leaves usually bitemate but variable; leaflets oblong-lanceolate or
302 Field Museum of Natural History — Botany, Vol. XIII
subelliptic, mostly gradually and obtusely acuminate, or the lower
lateral ones shortly ovate and obtuse, all obtusely serrate-dentate
or subentire (Peru), membranous or subcoriaceous, lustrous above,
rarely barbate in the nerve axils, punctae and pellucid lines obvious,
epidermis lacking mucus, the terminal petiolule 1-1.5 cm. long,
its leaflets often 1 dm. long or longer, 3-5 cm. wide or wider; cincinni
more or less stiped, 5-11 flowered, the pedicels 3-5 (-7 in fruit)
mm, long, articulate above the base; flowers white; outer sepals
usually glabrous, inner sordidly puberulent, nearly glabrous within,
3.5-5 mm. long; petals long-clawed, 4.5-7 mm. long, 1,5-2 mm. wide,
laxly glandular within; scale-crests more or less emarginate, even
bifid or the lower dentiform; stamens lightly villous; fruit purplish
above, 2.2-3.8 cm. long, 2-2.5 cm. (at cells 8-9 mm.) broad, little
if at all constricted under the cells, truncate-rounded or shortly
cuneate at base, ecristate, glabrous except for scattered glands,
the endocarp glabrous or with a few trichomes at base and back of
cells. — The Peruvian forms are genuina Radlk. and elliptica Radlk.,
the elliptic leaflets subentire; cf. S. Ampelopsis PI. & Lind. under
S. inscripta.
Hudnuco: Mito, 1567 (distr. as S. striata). Without data,
Ruiz & Pavdn, fide Radlk. Widely distributed in South America,
north to Cuba and Mexico.
Serjania communis Camb. in St. Hil. Fl. Bras. 1: 362. 1825; 51.
Slender sarmentose triangulate sulcate branches as the biternate
leaves variously pubescent to nearly glabrous; stipules ovate-
lanceolate; petioles 2-4 cm. long, scarcely margined; leaves about
14 cm. long, 12 cm. wide, the larger terminal leaflet subrhombic-
lanceolate, 5-8 cm. long, 2-3 cm. wide, more or less attenuate to the
petiole, the lateral subovate-lanceolate, rounded at base, all acute
or obtuse, mucronulate, serrate-dentate, often doubly, membranous
to chartaceous, rarely more than obscurely pellucid-punctate or
areolate; panicles mostly solitary, the lower twice as long as the
leaves, hirsutulous or laxly puberulent especially the small lance-
olate bracts and bractlets as also the 4 mm, long pedicels, these
equaling the ellipsoid buds; outer sepals half as long as inner, all
crisp-pulverulent; petals white, obovate, microscopically glandular
within, the upper 5-6 mm. long, 2-3 mm. wide, the half as long
scales emarginate or obtuse, sometimes obsolete as also the lateral
glands; stamens pilose-puberulent; ovary oblong, little narrowed at
base, minutely glandular; fruit cordate-ovate, 2-2.7 cm. long, 1.8-2
cm. broad, narrowed at the pubescent cells (these narrowly cristate),
Flora of Peru 303
emarginate at apex, the puberulent wings more or less dilated at
base; cotyledons subequal, suberect; seeds ellipsoid. — Radlkofer
named three variants in accord with the degree or character of the
pubescence, mollis Radlk., pilosula Radlk. and glabra Radlk. The
wood structure is central (large, with 3 smaller peripheral columns;
cf. the author, Monogr. Serj. Suppl. pl.l). I should like to exclude
this from Peru but if the following incomplete specimens actually
belong here probably some flowering material cited under S. inflata
does also, as originally determined. F.M. Neg. 36040.
San Martin: Tarapoto, Spruce (var. glabra, fide Radlk.). —
Junin: Puerto Yessup, Killip & Smith 26313 (det. Killip). To
eastern Brazil.
Serjania deltoidea Radlk. Monogr. Serj. 322. 1875; 203.
Branches obtusely or acutely triangular, lightly striate, the
younger reddish tomentulose with crisped trichomes; wood com-
posite with 3 peripheral areas; stipules small, ovate; common
petiole emarginate, caniculate above, the leaf-rachi in part winged,
the wings narrowed below; leaves deltoid in outline, often 16 cm.
long, 12 cm. wide, membranous, glabrous or nearly both sides,
lustrous above, subopaque and paler beneath, containing mucus,
minutely pellucid punctate, bi- or subtripinnate (upper pinnae
simple, the lower pinnate, the pinnules themselves in part pinnate)
with 3-4 pairs of ovate-oblong obtuse sessile coarsely dentate or
incised leaflets, except the terminal, these 3.5 cm. long, 1.5 cm. wide,
petiolulate, elongate-subrhombic with obtusish acumen; panicles
solitary or paniculate, slender and interruptedly floriferous, the
rachis reddish tomentulose, the dense cincinni subsessile 5-6-
flowered with 1 mm. long pedicels articulate at base; outer sepals
less tomentose than inner, these 2 mm. long; petals 2 mm. long,
half as wide, the upper scales crenulate, the lower subaliform; fruit
retrorsely subhirsute, retuse, contracted below the cells, 2.5 cm.
long, 2 cm. wide, the endocarp reddish villous, the seed basal.
F.M. Neg. 5539.
Hudnuco: Chicoplaya, Ruiz & Pavdn. Near Rfo Monz6n, 600
meters, Weberbauer 3606, part. Bolivia.
Serjania dibotrya Poepp. & Endl. Nov. Gen. & Sp. 3: 35.
pi. 2U2. 1844; 108.
Glabrate high-climbing liana, the stout somewhat trigonous and
striate branches early rusty setulose and microscopically glandular
304 Field Museum of Natural History— Botany, Vol. XIII
at the tips, 3-6 mm. thick, the ligneous structure composite with 1-3
peripheral areas; stipules to 2 mm. long, puberulent; petioles to
10 cm. long and rachi to 5, wing-margined (to 6 mm. wide), the
leaves 5-foliolate, the lower to 25 cm. long or longer, 2 cm. wide,
early especially on the lower surface with a few appressed setae,
subcoriaceous, transversely reticulate, densely pellucid punctate,
glabrous and lustrous above, containing traces of mucus, paler, dull
and glanduligerous beneath, the lateral leaflets to 11 cm. long, 7 cm.
wide, ovate, acute or shortly acuminate, the base rounded with
short petiolule, the terminal subrhombic little larger, attenuate at
base, all obtusely repand-denticulate or subentire; panicles solitary
and congested paniculately in the apex of the branchlets, more or
less reddish tomentulose, glabrate, densely cinniferous, the cincinni
shortly stiped, 7-9-flowered, the pedicels scarcely exceeding 1.5 mm.;
flowers hermaphrodite; sepals all tomentulose, the outer as the
interior nearly 4.5 mm. long; petals lanceolate-oblong, 5.5-6 mm.
long, 1.5 mm. wide, glanduligerous within, the less than half as long
scales little villous; anthers glabrous; fruit minutely setulose, about
3.5 cm. long, 2.3-3 cm. wide, cordate at base, obtuse, constricted
under the cells, these subglobose and ecristate, the endocarp glabrous
except at base of cells. F.M. Neg. 5542.
Hudnuco: In woods, Cuchero and Pampayacu, Poeppig, type.
Chinchao, Rivero 222. Pozuzo, Ruiz & Pav6n. — Junin: Colonia
Peren^, Killip & Smith 2520U (det. Killip). Bolivia.
Serjania diffusa Radlk. Monogr. Serj. 302. 1875; 194.
Branches subterete, 6-7-striate and more or less rusty hirsute-
tomentose as the solitary panicles, the rachi of these somewhat
angled; wood composite with 3 peripheral areas; petioles striate,
tomentose-hirsute as the leaf-rachi and the small ovate stipules;
leaves tritemate (in Mathews specimen bitemate), the subrhombic
terminal leaflet 8 cm. long, half as wide, with petiolule to 1.5 cm.
long, the upper lateral oblong, the lower ovate, sessile, all coarsely
incised-dentate, acute or acutish, thick-membranous, above sparsely,
on the nerves beneath densely pilose, pellucid punctae hardly
discernible, epidermis containing mucus; cincinni crowded on stipes
barely 2 mm. long; outer sepals lightly pilose, the inner canescent
tomentose, 3.5 mm. long; petals 4.5 mm. long, 2 mm. wide, the upper
scale crest emarginate or subentire, the lower dentiform; young
fruit oblong, cleft at apex, yellowish canescent, the endocarp white
floccose, the seed basal. — Mature fruits of Solis 10839, apparently
Flora of Peru 305
correctly named, are 15-18 mm. long, the hard cells densely setulose.
The author suggests the Mathews plant may not belong here.
Amazonas(?): Pariahuanca (Mathews 1202). Bolivia to Colom-
bia.
Serjania dumicola Radlk. Monogr. Serj. 111. 1875; 57.
In general like S. communis var. moUis but differing in structure
of flowers (female not described); stipules small, subulate; third
and fifth sepals free; petals 4.5 mm. long, 3 mm. wide; fruit cordate,
apically emarginate, little constricted medially, 2.8 cm. long, 2.5 cm.
wide, puberulent, the cells subcuneate at base, cristate dorsally. —
To be expected in adjacent Peru. Wood structure: medullary
cavities small or hardly any.
Peru (probably). Bolivia.
Serjania elongata Macbr. Field Mus. Bot. 11: 28. 1931.
Smooth liana, even the elongate solitary racemiform inflo-
rescences essentially glabrous, these laxly flowered and often 2-3 dm.
long; leaves 5-foliolate, the leaflets all entire or sometimes obscurely
1-3-undulate-dentate, broadly elliptic, very obtuse or rounded at
both ends except the cuneately based and petiolulately contracted
terminal, mostly about 6 cm. wide, 10 cm. long, chartaceous,
moderately reticulate and lustrous on both sides, scarcely pellucid
punctate or lineolate; rachis and petiole not at all margined; in-
florescence branchlets only 2-4 mm. long or in fruit apparently
about twice as long, the flowers usually 4 or few; pedicels slender,
barely 2 mm. long in flower; sepals about 2-2.5 mm. long, glabrous
except the slightly longer inner, these a little tomentulose; petals
obovate, about 3 mm. long, slightly barbate within at base; filaments
short-pilose; fruits obcordate, glabrous, about 2.5 cm. wide and
long, the cells compressed. — Description of fruit from Krukoff 10967
referred by A. C. Smith to an apparently unpublished species name
of Rusby, which appropriately defines the shape of the leaflets.
Junln: In sunny brush. La Merced, 5511, type. — Loreto: Trail
to Tarapoto, Ferreyra Jt95U. Bolivia.
Serjania exarata Radlk. Monogr. Serj. 166. 1875; 110.
Similar to S. membranacea but rather more hirtellous and especially
leaflets acute or acuminate, less coarsely, even obscurely serrate and
the fruit manifestly dilated at base, glabrous, the wings chartaceous,
the divaricate cells deeply exarate between the veins; fruit 2.5 cm.
306 Field Museum of Natural History— Botany, Vol. XIII
broad, 3.5 cm. long; inner sepals 3 mm. long; petals 4 mm. long,
2.5 mm. wide. — Simulates S. glabrata but presumably distinct by the
simple wood (as to type) ; compare also S. pyramidata. Klug 3707
only in flower may be distinct; the leaflets are abruptly acuminate,
about 1 dm. long, half as wide, obscurely dentate, inflorescences to
3 dm. long; without fruit doubtfully belonging here but for expedi-
ency it may be designated var. extensa Macbr., var. nov. foliolis
ad 5 cm. latis, obscure serratis abrupte subcaudato-acuminatis
inflorescentibus valde elongatis.
Huanuco: Pampayacu, Rio Chinchao, 50J^9 (distr. as S. sphaero-
cocca). — San Martin: Zepelacio near Moyobamba, Klug 3707? (det.
Standley, S. altissima). Amazonian Brazil.
Serjania fuscostriata Radlk. Bot. Jahrb. 37: 147. 1905; 171.
Allied to S. oxyphylla and in general similar but the somewhat
larger leaves imparipinnate with 3 pairs of unequally serrate-dentate
leaflets, the lower pinnae temate, the next mostly bifoliate, the upper
simple, sometimes one or the other as the terminal leaflet more or less
modified; cincinni stipes about 5 mm. long, the pedicels 4 mm. long,
articulate below the middle; inner sepals 4 mm. long, the outer
especially ciliate; petals 3.5-4 mm. long, the scales villous at margins,
the crest of the upper deeply bifid; fruit unknown. — To this I refer
with some doubt, the leaves incomplete and no flowers, my own
collection which once I determined as S. brachyptera, that, as to type,
with much smaller leaflets; the glabrous fruits are 2 cm. wide and
long, the cells subinflated, the rachis densely hirtellous, the upper
stems puberulent. F.M. Neg. 5548.
Ancash: Below Pampa Romas, among trees near brook, 1,900
meters, Weberbauer 3183, type; 163. — Hudnuco: On canyon shrubs,
2291.
Serjania glabrata HBK. Nov. Gen. & Sp. 5: 110. 1821; 113.
Older branches triangular, glabrate, the younger typically
sparsely pubescent but often densely and softly, 6-8-nerved;
ligneous structure composite with 3-5 peripheral areas; stipules
1-1.5 mm. long, ovate, puberulent; petioles 5-7 cm. long, the inter-
mediate 4-6 cm., bisulcate above, all emarginate; leaves finally
ample, bitemate, membranous, pale green, usually containing mucus,
obsoletely pellucid punctate and lineolate, glabrous above, laxly
to densely and softly pubescent beneath, the slightly larger terminal
leafiet sometimes 13-15 cm. long, 6-8 cm. wide, often smaller,
Flora of Peru 307
attenuate to petiolule 5-12 mm. long, subrhombic, the lateral
shortly contracted to the petiolules, all acute or subacuminate,
unequally and coarsely crenate, serrate or subrepand dentate;
panicles solitary and paniculately congested, the rachi sulcate-
angulate, the 5-16-flowered cincinni shortly stiped; pedicels 2-3 mm.
long, articulate near base or nearly at middle, puberulent; outer
sepals much shorter, glabrate or much less pubescent than the
3-4 mm. long whitish-tomentulose inner ones; petals white, 3.5-5
mm. long, 1-2 mm. wide, the upper scales subentire to dentate, the
lower bicomute; fruit usually 3-4 cm. long, 2-3 cm. broad, cordate-
ovate, hardly retuse, contracted under the cells, pilose or finally
glabrate, the cells transversely obovoid, divaricate or obliquely
erect, sulcate- and scrobiculate-exarate, the endocarp floccose. —
Forma genuina Radlk. is more glabrate than the commoner forma
moUior Radlk., the sepals distinctly diverse in degree of indument,
the pedicels articulate near base, the large fruit glabrate. F.M.
Neg. 5549.
Stems, macerated, used as a fish "poison" (Mexia); vine to
4 meters high, stems purplish, leaves soft, thin, glossy, paler beneath
(Woytkowski) ; flowers fragrant,
Piura: Canchaque, Prov. Huancabamba, Stork 11S97. — Caja-
marca: Colasai on the Rio Huancabamba, Bonpland, type. Calla-
cate, Jelski U15. Tambillo, Jelski 328. — San Martin: Tarapoto,
Spruce U1S9 (part) ; Ule 65SU; WiUiams 5^19; 5680; 585 Jt; 5908; 6261;
low woods and stream thickets, Woytkowski 35055; 35189 (both
det. Cuatrecasas). San Roque, Williams 72JtS; 7377; 775 J^; 7800.
Zepelacio, Kliig 3370 (det. Standley). — Hudnuco: White-flowered
vine in sunny shrubs, 4^56; Ruiz & Pav6n (the last det. Melchior). —
Junin: La Merced, 52^0- Near Peren6 Bridge, Killip & Smith 25^1
(in part S. ruMcauiis). — Loreto: Cachipuerto, Klu^ 3125 (det.
Standley, by slip of pen, "PauUinia"). Santa Rosa below Yuri-
maguas, Killip & Smith 28719 (det. Killip). Iquitos, Mexia 650 U
(det. Standley). Florida, Rio Putumayo, Klug 2080? (young).— Rio
Acre: Seringal San Francisco, Ule 9556; 9557. — Cuzco: Santa Ana,
Cook & Gilbert 16W' Paraguay to Ecuador. "Macote" (Williams;
Mexia); "verap" (Killip & Smith).
Serjania gramma tophora Radlk. Monogr. Serj. 258. 1875; 152.
Notable in Peru by the simply temate leaves, the three broadly
ovate (or the lateral, sometimes rather oblong-elliptic) leaflets
remotely repand-dentate or rarely sub-lobulate, all subsessile and
308 Field Museum of Natural History— Botany, Vol. XIII
mucronulately obtuse or shortly acute, shortly or the terminal
long-attenuate at base, not very unequal, often a dm. long or longer,
and about half as wide, early as the 6-7-sulcate stems, more or less
ashy pilose, glabrate in age, membranous, both sides minutely
glandular, pellucid punctate and lineolate, the epidermis lacking
mucus; wood simple; stipules minute; petioles emarginate, 4-5 cm.
long; inflorescences solitary, 6-12 cm. long, cincinni subsessile, ashy
tomentose including the sepals, the much longer inner 3.5-4 mm.
long, the basally articulate pedicels 1.5-2 mm. long; petals white,
5.2 mm. long, upper scales biappendaged; anthers puberulent. — The
type grown at Munich from seed, its origin unknown but similar
species are from the region of Guayaquil, Ecuador, S. longipes Radlk.,
151, and S. brevipes Benth., 151, in both of which the leaves are
obsoletely or most minutely punctate; in the former the elliptic
leaflets are subentire, the cincinni stiped, while in the latter
the leaflets are evenly 2-6-denticulate, the petals less than 3 mm.
long. F.M. Neg. 5968.
Peni(?) : See note above.
Serjania grandifolia Sagot, ex Radlk. Monogr. Serj. 166. 1875;
112.
Glabrous except the slightly pulverulent inflorescence including
the unequally, or in Peru subequally, puberulent sepals; wood
simple; stems 3-4 mm. thick; leaves bitemate but not rarely reduced,
the lower leaflets often simple; terminal leaflets well-petiolulate,
8-12 cm. long, 3.5-5 cm. wide, the lateral little smaller, all subentire
or entire, about oblong to broadly elliptic, obtusely short-acuminate,
coriaceous, lustrous at least above, obsoletely punctate and lineolate,
containing mucus; petioles canaliculate; petioles minute, broadly
ovate; inflorescences solitary, congested apically or on subleafless
branchlets; cincinni shortly stiped, the pedicels about 2 mm. long;
outer sepals in type subglabrous, nearly half as long as the tomentu-
lose inner, these scarcely 2.5 mm. long; petals nearly 3 mm. long,
glandular within; stamens basally pilose; fruits glabrous, reddish,
attenuate from cordate base, there about 3.5 cm. wide, at cells 1.4
cm. wide, to 5.5 cm. long, the ligneous sutures often bilamellately
produced. — The Peruvian liana seems to belong to this species
typically of the Guianas but may prove to be different when com-
pletely known; in flower the only marked distinction apparently is
the more pubescent outer sepals; for convenience it may now be
noted as var. pubisepala Macbr., var. nov. forma typica differt
Flora of Peru 309
sepalis plus minusve tomentulosis indumentum baud vel paullo
diversis.— 1)1% 965, type. F.M. Neg. 5550.
Loreto: Near Iquitos, KIilq 876; 88U; 965; Williams 8077.
French and Dutch Guiana; Amazonian Brazil.
Serjania inflata Poepp. & Endl. Nov. Gen. & Sp. 3: 36. 1844;
177.
Nearly glabrous, the branches acutely 6-angled and thus 6-
sulcate, sparsely crisp-pubescent, the flowering about 3 mm. thick;
wood composite with 2-3 small peripheral areas; stipules small,
ovate; petioles in part narrowly margined; leaves bitemate, about
1 dm. long, nearly as wide, the ovate-oblong subrhombic acute
terminal leaflets attenuate to petiolule, 5 cm. long, 2.5 cm. wide, the
others smaller, sessile, obtuse, even nearly orbicular, all remotely
serrate and at base sublobed, dark green, subcoriaceous, sparsely
crisped pubescent beneath and above on the nerves, more or less
pellucid with small dots, the epidermis containing mucus; panicles
solitary, minutely puberulent, the cincinni stipes 2-3 mm. long, the
pedicels 2-A mm. long, in fruit twice as long, articulate below or at
the middle; outer sepals slightly puberulent, the inner densely,
3.5-4 mm. long; petals 4.5 mm. long, 1.5 mm. wide, the scales about
two-thirds as long, the upper bifid, the lower emarginate; style
glabrous; fruit glabrous, cleft both ends, 3 cm. long, nearly 2 cm.
wide, inflated above, the seed below the middle of the cells. — S.
rigida Radlk., forma glabra Radlk., 175, of Brazil, has less acutely
angled branchlets, the leaf nerves very rigid beneath. Probably at
least sens. lat. should include S. rigida and apparently there is also
an Ecuadorean form. F.M. Neg. 5555.
San Martin: Tocache, Poeppig 1965, tjrpe. Tarapoto, Williams
6568. Caceres, Prov. Marisca, Ferreyra UU65. — Hudnuco: Near Rio
Monzon, Weherhauer 8606, part; 286. Pozuzo, Ruiz & Pav&n (in
herb, as PauMinia curassavica). — Junfn: San Ram6n, Killip &
Smith 2^756? (dist. as S. communis). La Merced, 5360; Killip &
Smith 25^08? (dist. as S. communis). — Loreto: Mishuyacu, Klug
2557 (det. Standley but aberrant). — Cuzco: Quillabamba, Prov.
Convenci6n, West 7197.
Serjania inscripta Radlk. Bot. Jahrb. 37: 145. 1905; 62.
Branches 5-6-angulate, canaliculate, 4-5 mm. thick, puberulent-
tomentulose at least above as the p>etioles and the solitary elongate
inflorescences; wood composite, the peripheric columns small, 3,
310 Field Museum of Natural History— Botany, Vol. XIII
angled; leaves bitemate, the leaflets oval-lanceolate, remotely
crenate-dentate, the terminal and upper lateral 8-9 cm. long,
3-3.5 cm. wide, the lower lateral 3-5.5 cm. long, 2-2.5 cm. wide,
obtuse, the others all acute or subacuminate, more or less attenuate
to the sessile base, membranous, glabrous above except the obliquely
ascending nerve, sparsely pilose beneath, notably marked with im-
pressed lines and pellucid punctae above, the epidermis containing
mucus, the intermediate petioles narrowly wing-margined; stipules
minute, deltoid; inflorescences with the long peduncle 1.5-nearly
3 dm. long; cincinni stipes about 1 cm. long; pedicels 3-4 mm. long,
to twice as long and medially articulate in fruit; flowers about or
little longer than 4 mm., the free sepals subequal and equally
tomentulose; petals glandular within the half as long scales with
deflexed villous appendage; fruits cordate, glabrous or the compressed
cells villous, at least 2.5 cm. long, about 2 cm. wide. — Suggests
S. leptocarpa with larger flowers and more strongly serrate leaflets.
The specimen from Cutervo has fruits 3.5 cm. long, the stems
glabrate, petioles obscurely margined, leaflets diverse, the roundish
lateral 1-3-dentate and may not belong here, even as a variant; it
suggests some specimens of S. Ampelopsis PI. & Lind., 58, of Ecuador
if that species is variable enough to include them but as to type it
seems to differ in its simple wood, subequal ovate-lanceolate acute
or acuminate leaflets all entire and even the lateral with petiolules
to 2 cm. long. F.M. Neg. 5556.
Ynes Mexia found the macerated stems used as a fish "poison."
Cajamarca: Izco, Prov. Cutervo, Stork & Horton 10210 (det.
Standley, S. pyramidata?) . — Loreto: Iquitos, Mexia 650 Jf. (det.
Standley, S. — by slip of pen, "Paullinia" — glabrata). Lower Rio
Nanay, Williams 612. Fortaleza near Yurimaguas, Klug 2812
(det. Standley, S. rubicaulis). Bolivia; adjacent Brazil. "Macote"
(Mexia).
Serjania Killipii Macbr., sp. nov.
Scandens, fruticosa; ramis superioribus petiolis pedunculisque
gracilis multistriatis minute subhirsutulo-puberulis; corpus lignosum
compositum e centrali majore et periphericis circa 8 paullo minoribus
et centrali cingentibus; foliis bitematis; foliolis integris glabris vel
subtus ad nervos obscure pubescentibus, minute pellucido-punctatis,
valde diversis superioribus oblongo-lanceolatis vel terminalibus
paullo obovatis ad_^ basin breviter attenuatis acutis vel interdum
obtusis 3.5-4 cm. longis, circa 1.5 cm. latis, lateralibus paullo
Flora of Peru 311
minoribus inferioribus similibus sed lateralibus vix 1 cm. longis,
4-5 mm. latis; thyrsi solitarii et in apice ramorum paniculatim
congesti dense cincinniferi, cincinni vix stipitatis; pedicellis circa
2 mm. longis; sepalis exterioribus 1.5 mm. longis dense puberulis
tamen sepalis interioribus; petalis anguste obovatis circa 3 mm.
longis; filamentis pilosis; fructibus ignotis.
There are notes by Killip on the t5T)e sheet suggesting that this
is near S. trirostris with "larger serrate leaflets," while the leaves
exactly match those of S. gracilis Radlk. of Brazil, a species with
larger flowers; in the absence of fruits its position of course is
uncertain but it does seem to be related to S. trirostris and S. sub-
rotundifolia; indeed it may prove with more collections to be a part
of the latter but in this case there must be considerable variation
in the size and shape of the leaflets.
Loreto: Masisea, Killip & Smith 26853, type (U. S. National
Herbarium).
Serjania leptocarpa Radlk. Monogr. Serj. 112. 1875; 57.
Scandent, subherbaceous, the younger subterete-trigonous
striate-sulcate, 6-costate branches, thyrsi, petioles and leaflets both
sides more or less pubescent with short curved trichomes, finally
glabrate; branches brownish, 2-3 mm. thick; stipules minute;
petioles 5-7 cm. long, the intermediate little shorter, the lateral
1.5-2 cm. long, the larger bitemate leaves 18-22 cm. long, 1&-18 cm.
broad, the basally attenuate terminal leaflets about 9 cm. long,
4.5-6 cm. wide, the basally rounded lateral ones 5-8 cm. long, 3-4
cm. wide, all ovate, acute or subacuminate, angulately bi- or tri-
dentate or subserrate, subpetiolulate, glabrescent, membranous,
drying brownish, obscurely pellucid-punctate or -areolate; thyrsi
solitary, elongate, exceeding the leaves, the smaller often panicu-
lately congested in branchlets; buds 4.5-5 mm. long subequaling the
nearly glabrous pedicels; flowers in tjrpe pale rose, the subglabrous
outer sepals half as long as the whitish puberulent inner, the third
and fifth barely coalescent at base; petals clawed, obovate, about
8 mm. long, 3.5 mm. wide; upper scales dilated, the lower dentiform;
stamens short-pilose; ovary obversely pyramidal triquetrous; fruit
glabrous except microscopically glandular, 4-^.5 cm. long, 2.5-3 cm.
broad, little constricted below the cells, excised base and apex, the
wings obscurely dilated at base. — Wood structure, Radlk. I.e.
Suppl. pi. 3, the medullary cavity large. The Peruvian material
placed here with some doubt. Illustrated, Mart. Fl. Bras. 13, pt. 3:
pi. 58.
312 Field Museum of Natural History — Botany, Vol. XIII
Hudnuco: Near Chinchao, Ruiz & Pavdn, (part; cf. S. pyrami-
data). — Rio Acre: Seringal Auristella, Ule 9558. Bolivia; Brazil.
Serjania lethalis St. Hil. Hist. PI. Remarq. Br^sil 1: 206. 1824;
138.
High climbing glabrate liana, the stems obtusely triangular, the
branches subglabrous to subtomentulose, about 3 mm. thick, more
or less 6-striate and obscurely trigonous; ligneous structure com-
posite, with 3-4 triangular peripheral areas; stipules ovate, minute;
petioles sulcate above, the intermediate especially often narrowly
margined; leaves bitemate, the lower 26 cm. long, 24 cm. wide, all
sometimes on nerves a little pubescent or ciliate marginally, other-
wise mostly glabrous both sides except for microscopic glands,
lustrous, coriaceous, obsoletely pellucid punctate, containing traces
of mucus; leaflets lanceolate, narrowed both ends, sessile, obtuse or
obtusely prolonged into a linguiform acumen, entire or 1-3-dentate,
the terminal 9-15 cm. long, 3-4.5 cm. wide, the lateral little smaller;
thyrsi solitary or congested, the subsulcate rachi tomentose, the
mostly well-'stiped cincinni 6-9-flowered, the pedicels 3-6 mm. long,
articulate near base; outer sepals shorter, the inner 3-4 mm. long,
all lanate- tomentose; petals 4-4.5 mm. long, 2-2.5 mm. wide, the
scales more or less bifid; fruit cordate at base, where 2-2.5 cm.
broad, 2-3 cm. long, little if at all contracted under the cells, these
subglobose, lanate-tomentose, scarcely cristate, the lustrous wings
nearly glabrous, the endocarp more or less villous. — Ex range the
Peruvian plant could be S. nutans or S. paucidentata, especially
the latter, which compare.
St. Hilaire and Radlkofer, Monogr. Serj. 229, have discussed the
toxic qualities of this plant, used as a "fish poison" and said to be
injurious to other animals including man,
San Martin: Tarapoto, Spruce ^892 (det. Radlk.). Bolivia to
eastern Brazil. "Timbo"; "Sacha."
Serjania longistipula Radlk. Bot. Jahrb. 37: 148. 1905; 177.
More or less pubescent, the 3-angled lightly sulcate branches
typically with only a few long subsetaceous deciduous trichomes or
in variant densely hirsutulous-pilose; wood composite with 3 peri-
pheral areas; stipules linear-subulate, pilose, 8-10 mm. long; petioles
3-5 cm. long, the lateral 1.5-2 cm. long, all emarginate, striate;
leaves imparipinnate with 3 pairs of oval obtuse sessile or nearly
sessile leaflets, the terminal attenuate to petiolule, 7-11 cm. long,
Flora of Peru 313
3-4.5 cm. wide, the lateral smaller, rounded at base, all subserrate
about the lower half, membranous-chartaceous, minutely pellucid-
punctate and lineate, containing mucus, above on nerves densely,
between them sparsely pubescent with subsetaceous curved ap-
pressed trichomes, the pubescence beneath shorter and softer; lowest
pair of leaflets temate or 5-foliate-pinnate; panicles solitary, the
cincinni shortly stiped, the pedicels 4 mm. long, articulate below the
middle; outer sepals minutely, the inner ashy puberulent, 4 mm.
long; petals 4.5 mm. long, the villous scales with deflexed appendage;
young fruit slightly emarginate, somewhat yellowish tomentose, the
endocarp white-villous. — Apparently there is a variant with more
pubescent stems and more acute leaflets, the Stork and Horton
specimen less extreme in these characters; it may be known as var.
aberrans Macbr., var. nov., foliolis plus minusve acutis, ramulis
dense piloso-hirsutulis. — Weberbauer 7878, type. F.M. Neg. 5561.
Apurlmac: Prov. Andahuaylas, 2,800 meters, Stork & Horton
107S5 (var. det. Standley, Paullinia). — Cuzco: Marcapata Valley
above Chilechile, Weberbau£r 7878 (type, var. det. Macbride, S.
diffusa). — Puno: Among shrubs, 2,100 meters, Weberbauer 518, type;
also 537; 237.
Serjania membranacea Splitgerber, PI. Nov. Surinam, in
Hoeven & Vriese, Tijdschr. Nat. Gesch. 9: 11 (105). 1842; 109.
Scandent, little pubescent, the younger branches slightly and
obtusely 5-6-angled, lightly pilose or at the apex hirtellous, the
older subterete, multistriate, glabrate; ligneous structure simple;
stipules minute, broadly triangular; petioles 5-6 cm. long, the inter-
mediate 2.5 and the lateral 1 cm. long, bisulcate and pubescent
above; leaves bitemate (sometimes reduced), minutely pellucid-
punctate, containing mucus, hirtellous only on the nerves above
where lustrous, beneath paler, dull and longer pubescent on the
nerves, the larger 14 cm. long, 12 cm. wide; leaflets about 5 cm. long,
3 cm. wide, shortly ovate, remotely and coarsely blunt-dentate, the
little larger terminal contracted to the petiolule, the lateral sub-
sessile, all obtuse or shortly with mucronulate acumen; panicles
solitary and paniculately congested, the cincinni stiped, 7-9-
flowered, the pedicels 2 mm. long, subglabrous, articulate near the
base or at about one-third; sepals all more or less puberulent, the
nearly twice as longer inner hardly 2.5 mm. long; petals 2.5-3 mm.
long, 1 mm. wide; scales about as in related species, tomentose at
margins; anthers glabrous; fruit slender, cordate at the little dilated
314 Field Museum of Natural History— Botany, Vol. XIII
base, 2 cm. broad, 3.5 cm. long, retuse, glabrous with diaphanous
wings, the divaricate cells depressed between the transverse nerves,
the endocarp floccose. — The Lechler specimen was possibly mis-
labeled as suggested by Radlkofer; if Peruvian, see S. exarata or
S. leptocarpa to which also the rest of the material cited here could
be referred except that it has the smaller flowers of S. memhranacea;
the simulating S. tenuifolia has sessile cincinni. F.M. Neg. 23670.
Loreto: Timbuchi on Rio Nanay, Williams 970 (distr. as S.
leptocarpa). Maquisapa, Upper Rio Nanay, Williams 1199 (distr.
as S. exarata). Manfinfa, Upper Rio Nanay, Williams 1091
(distr. as S. leptocarpa) . — Puno : San Govdn, {Lechler 2332) . Guiana;
Costa Rica. "Novia sisa" (Williams).
Serjania mollis HBK. Nov. Gen. & Sp. 5: 108. 1821; 67.
More or less rusty tomentulose — or the younger parts as the
sepals ashy — the stout 5-6-costate branches 4-6 mm. thick, obtusely
striate between the ribs; ligneous structure composite, the peripheral
areas 3-5; stipules lanceolate-subulate; petioles 3-4 cm. long; upper
leaves temate, to 14 cm. long, 12 cm. wide, the leaflets ovate, crenate
or subserrate-dentate, obtuse, rounded and subcordate at base or the
larger subtrilobed terminal ovate and decurrent into the 2 cm. long
petiolule, 9-11 cm. long, 7-8 cm. wide, the lateral little smaller, sub-
sessile, all tomentose-pilose, or in age scabrous above, most obscurely
if at all pellucid-punctate, epidermis lacking mucus; panicles equal-
ing the leaves, the cincinni shortly stiped; bracts and bractlets 4 mm.
long; inner sepals 5 mm. long, canescent both sides; petals not known,
pedicels 2 mm. long; fruit broadly cordate-ovate, 2-3 cm. long,
2.4-4 cm. broad, emarginate and with short hirsute style at tip,
deeply cordate at base by the long-produced wings, tomentose and
hirsute, the cells broadly cristate dorsally and usually comigerous,
the endocarp arachnoid-floccose, the seed affixed above the base of
the cells. — My specimen had fragrant greenish-white flowers with
yellowish centers; it is probably S. sufferuginea, as the following
except the Cajamarca specimens, if that is distinguishable. How-
ever, the monographer has .S^. mollis in his section Ceratococcus and
the latter in section Physococcus, the former with rather compressed
fruit cells, medially tumidulous, laterally comiculate, alulate or
callose produced while the latter section is described as having cells
inflated, scarcely ever cristate; these sections may be distinct as
regards other species but scarcely in respect to these plants. F.M.
Neg. 5564.
Flora of Peru 315
Cajamarca: Between Guerocotillo and Montdn, Bonpland, type.
Monte Seco, Prov. Hualgayoc, 1,800 meters, Soukup 3831 (distr.
S. ferruginea). — Hudnuco: Mufia, 3905. — Ayacucho: Aina, Killip
& Smith 22Jk55. — Cuzco: San Miguel, Urubamba Valley, Cook &
Gilbert 977.
Serjania nutans Poepp. & Endl. Nov. Gen. & Sp. 3: 36. pi. 2^2.
1844; 144.
With the habit and pubescence of the allied S. lethalis; branches
(typically) acutely 6-8-angled, the ligneous structure more com-
posite; larger leaves 32 cm. long or longer, 20 cm. wide, membranous,
glabrous or early with some scattered reddish trichomes on nerves
both sides, paler beneath, pellucid-punctate, sometimes not con-
taining mucus, the leaflets obovate, the lower lateral elliptic, the
terminal to 14 cm. long, half as wide, all cuneate at sessile base,
obtuse or shortly acuminate, subentire or remotely crenate-dentate;
panicles solitary, laxly flowered, reddish puberulent, the short cin-
cinni sessile or nearly, the pedicels 8 mm. long, articulate above the
base; sepals all ashy-tomentulose, the inner 5.2 mm. long; petals
9 mm. long, 4 mm. wide, the upper scales subentire to bifid; style
scarcely 1 mm. long, puberulent; fruit 3.5 cm. long, 2.2 cm. broad
(Bolivian species), 2.5-3 cm. long, the cells hispid (Peru), horizon-
tally truncate at base according to Poeppig who noted the petioles
as margined, emarginate fide the monographer. — It is possible this
may emerge with several other similar extra-Peruvian forms in-
cluding S. paucidentcUa and S. lethalis.
San Martin: Tocache, Poeppig 1897, type. — Loreto: Balsa-
puerto, Kliig 3081 (det. Standley, S. glabrata). Masisea, Killip &
Smith 26852. Yurimaguas, Killip & Smith 27573.— Rio Acre:
Seringal San Francisco, Ule 9559. Bolivia; Brazil.
Serjania oxyphylla HBK. Nov. Gen. & Sp. 5: 112. 1821; 170.
Scandent shrub, but the leaves beneath on nerves and margins
a little puberulent and the rachises of the solitary or congested
panicles tomentose-hirtellous; branches about 10-striate, 10-costate,
the ligneous structure simple; stipules small, ovate-acuminate;
petioles striate, emarginate or the secondary slightly margined;
leaves 11-16 cm. long, 9-14 cm. wide, bitemate or the terminal
leaflet 3-lobed or dissected, the lower pinnae temate, the others
simple; terminal leaflet 6 cm. long, 2.5 cm. wide with petiolule 12 mm.
long, the lateral gradually smaller, sessile, ovate-elliptic to sub-
316 Field Museum of Natural History — Botany, Vol. XIII
rotund, usually acute or acuminate, remotely and obsoletely to
conspicuously serrate or incised, membranous, glabrous and brown-
ish-green above, paler beneath, pellucid-punctate and lineolate,
lacking mucus, the few nerves oblique-erect; cincinni 6-9-flowered,
the lower with stipes 2 mm. long; pedicels glabrous, articulate at
base, becoming 2 mm. long; inner sepals 2-2.5 mm. long, all merely
ciliolate or fimbriate; petals 2 mm. long, about 0.7 mm. wide, the
upper scales crenulate or subbifid, the lower uncinate-corniform;
fruit shortly ovate or elliptic, cleft both ends, 1.5 cm. long, 1.2 cm.
broad, scarcely contracted below the cells or dilated at base, the
cells 7-8 cm. long, 6 cm. wide, inflated apically, glabrate, the endo-
carp glabrous. F.M. Neg. 5568.
Cajamarca: Near Querocotillo, Ja^n de Bracamores, Bonpland,
type. Callacate, Jelski UIO, part.
Serjania parvifolia HBK. Nov. Gen. & Sp. 5: 113. 1821; 172.
Resembles S. oxyphylla but the branches about 8-sulcate, the
ligneous structure composite with 3-5 peripheral areas, the tips as
the petioles, leaf-nerves beneath with some small trichomes, other-
wise glabrous or nearly; leaflets 3 pairs, the lower temate, the rest
simple, the terminal leaflet about 1.5 cm. long, 6 mm. wide, attenuate
to petiolule, acutish, the lateral obtuse, ovate or oblong, subsessile,
all mucronate, crenate-serrate, subcoriaceous, paler and in nerve
axils beneath barbate, obsoletely pellucid-punctate, the epidermis
containing mucus; stipules minute; petioles including the lateral
not margined; panicles solitary, the cincinni on stipes 1.5-3 mm.
long; fruiting pedicels 3 mm. long, articulate near base; inner sepals
1.8 mm. long, all nearly glabrous; fruit glabrous, 8-9 mm. long and
about as broad, the cells well-produced and inflated above, wing-
dilated below the middle. F.M. Neg. 5571.
Cajamarca: Tomependa, Bonpland, type. Callacate, Jelski
Jf-lO, part.
Serjania paucidentata DC. Prodr. 1: 603. 1824; 141.
Scandent essentially glabrous shrub, the younger branches, these
2.5-4 mm. thick, and inflorescences sparsely pubescent or glabrate;
branches canaliculate, 6-angulate, the alternate angles more acute
or rarely costately obtuse; wood composite, the 3 peripheral columns
small, oblong; stipules small, ovate; leaves bitemate, the leaflets
elliptic or oblongish, the larger terminal more or less abruptly and
marginally petiolulate often to a dm. long, about half as wide, all
Flora of Peru 817
typically with a short linguiform acumen with 1-4 depressed teeth
below it on each side, lustrous or subopaque, membranous-coriaceous,
obsoletely and minutely pellucid-punctate, the epidermis containing
mucus; petioles usually margined only above; inflorescences solitary
or apically panicled, sometimes 2 dm. long or longer, the cincinni
stipes 3-10 mm. long, 7- rather many-flowered; pedicels about 2 mm.
long; sepals all ashy-tomentulose, the longer inner 3.5 mm. long;
petals oblong, attenuate below, 4.5 mm. long, glanduliferous within,
twice as long as the scales; fruits triangular from the cordate-excised
base, nearly 3 cm. long, nearly 2 cm. wide or at the pilose ovoid or
ellipsoid cristate cells 7 mm. wide, finally glabrate. — Very much like
S. lethalis unless the acutely angled branches and other perhaps
minor characters serve to distinguish it; with that species it is the
earliest name for closely allied forms. F.M. Negs. 5616; 33410.
San Martin: Tarapoto, WiUianis 6100. — Loreto: Pumayacu,
between Balsapuerto and Moyobamba, Klug SI 96 (det. Standley,
Paullinia grandifolia). Amazonian Brazil to Trinidad and Tobago.
Serjania peruviana Radlk. Monogr. Serj. Suppl. 154. 1886;
193.
Branches as the paniculate panicles sordidly canescent tomentu-
lose, the former subterete but 6-sulcate, the rachises of the latter
terete, not sulcate-angled; wood composite with 3-4 peripheral
areas; stipules unknown; petioles all emarginate, ashy tomentose;
leaves bitemate, the terminal leaflet 6 cm. long, 3.5 cm. wide, sub-
rhombic, the smaller lateral ones ovate, all subsessile, subacute,
crenate-dentate, thick-membranous, pubescent above, tomentose
beneath, nearly epunctate, the epidermis lacking mucus; cincinni
stipes only about 3 mm. long, the pedicels as long, articulate near
the base, sepals all canescent, the inner 4.5 mm. long; petals 5.5 mm.
long, 3 mm. wide, the upper oblong scale crest approximately as long,
the lower small, wing-like; stamens hirsute. — Otherwise unknown
and not seen by me but without fruit its position remains uncertain
until re-collected; except for the "apically paniculately congested
panicles" it seems to be, probably, a part of S. mollis or (and)
S. sufferuginea; well-developed specimens of the latter at least have
paniculate inflorescences and the length of the cincinni stipes
appears to depend on age.
Amazonas: Chachapoyas (Mathews 309It, type, herb. Kew).
Serjania platypetala Radlk. Bot. Jahrb. 54: Beibl. 117: 42.
1916; 64.
318 Field Museum of Natural History — Botany, Vol. XIII
Resembles S. squarrosa; branches sulcate, 6-10 costate, with
only 1 peripheral ligneous area; stipules minute, subulate; leaves
membranous, with a few subsetaceous trichomes above, glabrate
beneath, the terminal 1.5-3 cm. long, 1-1.5 cm. wide, the lateral
not more than half as large, oval or suborbicular; panicles to 3 dm.
long including the 8-10 cm. long peduncles; pedicels to 6 mm. long,
articulate below the middle; outer sepals glabrous; the inner 5 mm.
long; petals 6-7 mm. long; lateral glands obsolete; fruit shortly
ovate-subquadrate, laxly puberulent at tip, endocarp sordidly
(instead of white as in related species) villous, 1.5 cm. long, 1.3-1.5
cm. broad. — The author distinguished this by the membranous
leaves, longer inflorescences and cincinni stipes, smaller ovate-
subquadrate fruit. It seems to me doubtful that these characters
are significant.
Ayacucho: Near Ayacucho, rocky slopes among shrubs, 3,000
meters, Weberbauer 5506, type.
Serjania pyramidata [R. & P.] Radlk. Monogr. Serj. 155.
1875; 99.
Character in general that of the similar S. sphaerococca but the
petioles all (or the intermediate obscurely margined) emarginate,
the leaflets ovate-elliptic, remotely and obtusely serrate above the
middle, a little pilose on the midnerve both sides as well as in the
axils beneath, the lower 2 dm. long, nearly as wide, the terminal
leaflet 1 dm. long, 4.5 cm. wide, all sessile or nearly; pedicels 1.5-2
mm. long, in fruit 2.5 mm. long; outer sepals sparsely puberulent,
the inner ashy-tomentulose, 2 mm. long; petals 2.5 mm. long, 1 mm.
wide, the scales villous at margins, the upper with deflexed append-
age; fruit 3 cm. long, 2.2 cm. broad, the coarsely veined cells ob-
viously obliquely cuneate at base instead of globose and obscurely
cuneate as in the related species. — In herbaria as Paullinia pyrami-
date R. & P. Here might be sought S. mucronulata Radlk., 173, of
southern Ecuador intermediate in character of fruit to two sections
(Radlkofer), among Peruvian species nearest perhaps to S. inflata
with rather similar leaflets but somewhat larger flowers and inflated
fruit cells; the acute leaflets and smaller fruits distinguish it; the
fruits when mature suggest those of S. exarata, in nervation of
the cells.
Hudnuco: Valley of the Monzon, 600 meters, Weberbauer 3610;
862^; 286. Cuchero and Chinchao, Ruiz & Pav6n, type; Dombey.
Ecuador; Venezuela.
Flora of Peru 319
Serjania rubicaulis [R. & P.] Benth. ex Radlk. Monogr. Serj.
254. 1875; 150.
Scandent, suffruticose, more or less aculeate, and pubescent or
puberulent; branches canaliculately 5-sulcate, the ligneous structure
simple; petioles emarginate or the intermediate ones slightly; leaves
bitemate, nearly glabrous except both sides punctiform-glandular,
and slightly pubescent near the nerves beneath with crisp trichomes,
membranous-subcoriaceous, reticulate- veined, greenish-brown, paler
beneath, lacking mucus, pellucid-punctate and lineate; leaflets
lanceolate-oblong, the lower lateral ovate, obtuse, the others acute
both ends, sessile and with 1-3 (-5) large or small teeth near the tip
on both edges, the somewhat larger terminal often 10-12 cm. long,
4-5 cm. wide, sometimes smaller, rarely much wider; panicles mostly
paniculately racemose, crisply pubescent; flowers small or medium,
the sepals all densely tomentulose, upper petal scales merely emar-
ginate or calloused; fruit section cordate-ovate, abruptly narrowed,
little if at all contracted below the ecristate cells, glabrous as also
the endocarp, 1.5-2 cm. long, about 1-1.5 cm. wide, the seed medially
affixed. — The species name was originally in Ruiz and Pav6n's
Journal under Paullinia.
Killip and Smith noted its use as a fish stupefier.
San Martin: Valley of the Rio Mayo near Moyobamba, Weber-
bauer ^532; 289. Zepelacio, Klug 8730 (det. Standley, S. glabrata).
Tocache, Ferreyra U3^; H60. Pongo de Cainarachi, Klug 2613
(det. Standley). Juanjui, Ferreyra U55U- Tarapoto, SpriLce U139
(in part S. glabrata). — Hudnuco: Near Pozuzo, Ruiz & Pavdn, type.
Woods, Pampayacu and Cuchero, Poeppig. Distrito Divisioria,
Woytkowski Sj^56 (det. Cuatrecasas). Churubamba, Mexia 8120;
8205; 8215 (all det. Standley).— Junin: Rio Paucartambo, Valley
near Peren^ Bridge, Killip & Smith 30682 (det. Killip) ; also 25 W
in part (apparently in part S. glabrata, as det.). — Cuzco: Potrero,
Dept. Convencion 1,250 meters, Vargas 2H7 (distr. as Paullinia
species). — Loreto: Wooded banks of Lower Rio Huallaga, Killip
& Smith 29011 (det. Killip). Lorenzo, between Rios Pastaza and
Huallaga, Dennis 29282 (det. Killip, S. glabrata?). Bolivia. "Verap"
(Killip & Smith), "anti-Christo" (Mexia).
Serjania rufa Radlk. Monogr. Serj. 324. 1875; 204.
Reddish hirsutulous with mostly crisped or curved trichomes
including the 6-8 angled-sulcate branches, the bitemate leaves on
both sides, especially on the nerves, the panicles even to the sepals
320 Field Museum of Natural History — Botany, Vol. XIII
but there the indument more reduced and subtomentulose; petioles
striate, emarginate or the intermediate in part margined; stipules
small, lanceolate; terminal leaflet obovate-cuneate, 8 cm. long, half
as wide, the smaller lateral rather obliquely oblong-elliptic, all
sessile, abruptly and acutely acuminate, coarsely serrate from the
middle, subcoriaceous, nitidulous above, opaque beneath, minutely
pellucid-punctate, containing mucus; panicles solitary or panicled,
often 1-1.5 dm. long, slender, the glomeruliform cincinni sessile,
the basally articulate pedicels scarcely exceeding 1 mm.; inner sepals
2 mm. long, the third and fifth united nearly to the middle; petals
2-2.5 mm. long, 1 mm. wide, the upper scales obcordate, the lower
aliform; ovary pilose. — Type in flower; fruit of Killip & Smith
25063, fruits 2 cm. wide, 2.5 cm. long, lightly pubescent especially
toward the subglobose cells, their pericarp subindurate, those of my
5554- similar but as wide as long. F.M. Neg. 5580.
Mashed stems used to stupefy fish (Killip & Smith) .
San Martin: Near Tarapoto, Mathews 1321, type; Spruce J!t559. —
Junin: On river bank brush near La Merced, 555 J^; Killip & Smith
238U5 (det. Killip). Colonia Peren^, KiUip & Smith 25063 (det.
Killip). Pichis Trail, Killip & Smith 25k2k. (det. Killip). Rio
Paucartambo Valley, KiUip & Smith 25376. "Verap" (Killip &
Smith).
Serjania sphaerococca Radlk. Monogr. Serj. 153. 1875; 96.
Allied to S. caracasana which it resembles except that the
younger stems, as the rachises of the inflorescences, are at least
moderately pubescent with sordid or yellowish crisp trichomes, the
leaflets lanceolate and barbate in the axils of the nerves, the inter-
mediate and lateral petioles usually margined; leaves often 16 cm.
long, 12 cm. wide, the upper gradually smaller; petioles 4-9 cm. long,
early hirtellous; leaflets narrowed both ends, equally crenate or
subserrate, the slightly larger terminal one 7-9 cm. long, 2-2.5 cm.
wide; panicles often congested at branchlet tips and much longer
than the reduced leaves; pedicels 2.5 mm. long, in fruit to 4 mm.;
inner sepals 4 mm. long; fruit 2.5-3 cm. long, about 2 cm. broad. —
Illustrated, Mart. Fl. Bras. 13, pt. 3: pi. 69.
Hudnuco: Chinchao, Ruiz & Pavdn. Cuchero, Poeppig, addenda
j^9. — San Martin: Juanjui, King 3800 (det. Standley, S. leptocarpa?) .
Tarapoto, Spruce U060. — Junin: Along Rio Peren^ near Colonia
Peren^, Killip & Smith 25109 (det. Killip). La Merced, Killip &
Flora of Peru 321
Smith 23 J^90?.— Ay 3LCucho: Aina, Killip & Smith 23099 (det. Killip).
Bolivia.
Serjania squarrosa Radlk. Monogr. Serj. 115. 1875; 63.
Scandent shrub with rusty-red 5-6-angulate branches 2.5 mm.
thick, the lateral lightly canaliculate, puberulent toward the tips;
central ligneous area large with only 1 or 2 small peripheral ones;
stipules lanceolate; petioles 1.5-2.5 cm. long, striate, narrowly
canaliculate above; leaves subbipinnate, 8-9 cm. long, 5-6 cm. wide,
the lower pinnae 5-foliolate-pinnate, the following ternate, the upper
simple; terminal leaflet 2 cm. long, half as wide, the lateral half as
large, all subtrinerved, subcoriaceous, paler beneath, the smaller
ovate, cuneate at base, sessile, crenate or subserrate, scabrous
above, below on nerves a few scattered long trichomes and barbate
in the lower axils, epunctate or minutely and very obscurely punc-
tate, the epidermis containing mucus; rachis margined above;
panicles 7 or 8 cm. long, squarrosely spreading, about one- third
floriferous, puberulent, the free sepals sordidly; bracts and bractlets
subulate; flowers "rather large"; petals attenuate-obovate, glanduli-
ferous within, flabellately nerved, the scales of the upper with
deflexed appendage; stamens a little pilose; fruit cordate-ovate,
2 cm. long, as broad below, sometimes suborbicular, excised base
and apex, the cells dorsally cristate, puberulent, the endocarp
whitish villous. — Male flowers in type imperfect from fungus.
Most of the following material was distributed as S. platypetala
and some of it at least may belong there if that form is separable.
F.M. Negs. 5979; 36051.
Apurimac: Abancay, 2,400 meters, Vargas 9Jt3. — Cuzco: Pachar,
2,900 meters, Pennell 13688. Huasao, Prov. Quispicanchi, Herrera
687. Ollantaitambo, Cook & Gilbert 603 (det. Killip, S. brachypterd) ;
Pennell 13685; Cook & Gilbert 389; Herrera 8j^9a.—Funo: Ollachea,
Soukup 500. Without locality, Gay, type. "Huecjucjilla," "be-
juchilla" (both, Cook & Gilbert).
Serjania striata Radlk. Monogr. Serj. 280. 1875; 172.
Scandent shrub, glabrous except for some scattered trichomes
on the imparipinnate leaves beneath on the nerves and at the
margins, barbellate in the nerve axils, the tomentulose inner sepals
and the apex of the young fruit; branches about 10-striate, 10-
costate, the ligneous structure simple; petioles and leaf-rachises
striate, emarginate, unless the lateral; stipules small, ovate; leaves
16 cm. long, 14 cm. wide, the ovate leaflets 5-7 cm. long, 3-4 cm.
322 Field Museum of Natural History — Botany, Vol. XIII
wide, 3 pairs, the lower pinnae temate, the next often tripartite,
rarely subentire, the lower obtuse, the rest acute or caudate, the
terminal leaflet often tripartite or pinnate, all dentate or nearly
lobed, rather obovate and plicate, membranous, somewhat lustrous
above, paler beneath, true pellucid-punctae lacking, epidermis
lacking mucus; panicles solitary on stipes 5-10 mm. long, the 3-5
mm. long puberulent pedicels articulate below the middle; outer
sepals glabrous, the inner 5 mm. long; petals 6 mm. long, 3 mm.
wide, both the upper and lower scales bicruriate; fruit (young)
oblong, the cells villous-tomentose with seed at base, the endocarp
glabrous. — The Goodspeed collection matches the photograph but
the leaflet divisions are smaller. F.M. Neg. 36052.
Apurimac: Rio Pachachaca, 3-4 meters in shrubs, flowers waxy
white, 2,000 meters, Goodspeed Exped. 10521 (det. Standley). —
Cuzco: Gay, type.
Serjania striolata Radlk. Bot. Jahrb. 37. 147. 1905; 170.
Like S. oxyphylla but the branches ashy-pubescent at tip,
minutely puberulent between the costae, the leaves imparipinnate
with 3-4 pairs, the lower pinnae mostly 5-foliate-pinnate, the next
temate, the upper simple, all multinerved, the nerves spreading;
punctae not obvious; terminal leaflet acutely subulate-caudate,
the lateral obtuse, all with 8-12 approximate nerves; panicles
solitary, ashy-pubescent except the 1.5 mm. long pedicels that are
articulate below the middle; sepals glabrous except ciliate-glandular;
buds 1.5 mm. long, the opened flowers unknown. — It seems possible
that this could be a vigorous state of S. oxyphylla. F.M. Neg. 5582.
Ancash: In wood near river, 2,200 meters, Huaraz, Weherhauer
3031, type; 172.
Serjania subrotundifolia Radlk. Monogr. Serj. Suppl. 118.
1886; 132.
Somewhat pubescent liana, the younger branches as the in-
florescence reddish hirtellous, the older glabrate, 8-sulcate, 8-costate,
the composite wood with as many peripheral areas; leaves biternate,
the upper 15 cm. long, 12 cm. wide, membranous-coriaceous, lustrous
both sides, brownish, containing mucus, pellucid-punctate, glabrous
above, puberulent beneath, especially on the arcuate-ascending
lateral nerves and at the margins; leaflets elliptic, the lower lateral
subrotund, retuse and with apical callose on the lower side, subentire,
attenuate or contracted at the subsessile base, the terminal 6.5 cm.
Flora of Peru 323
long, 3.5 cm. wide, the lateral 2.5 cm. long, hardly 2 cm. wide;
petioles all emarginate; panicles congested, the cincinni stiped, the
pedicels barely 2 mm. long; inner sepals less than 2.5 mm. long,
tomentulose, the half as long outer ones also a little hirtellous;
petals 2.5 mm. long, scarcely 1 mm. wide, the upper scale crest
dilated, the lower dentiform; stamens villous, anthers glabrous
as the rudimentary ovary. — The type number as in Kew is the
same number (maybe the same species?) as the dubious collection
of S. membranacea.
Puno: Lechler 2SS1, type {2332 in herb. Kew).
Serjania sufferuginea [R. &. P.] Radlk. Monogr. Serj. 299.
1875; 176.
Scandent shrub, the 6-striate subterete branches sufferrugineous-
tomentose, about 3.5 mm. thick; ligneous structure composite, the
small peripheral areas 3; stipules small, ovate; petioles all emarginate
tomentose as the bitemate leaves especially beneath, these 16 cm.
long, 14 cm. wide, rather fleshy membranous, minutely pellucid
with orbicular punctae, the epidermis lacking mucus; terminal
leaflet 9 cm. long, 4.2 cm. wide, subrhombic-lanceolate, the elliptic-
lanceolate lateral gradually reduced, all sessile or the terminal
attenuate to short i)etiolule, crenate, acute; panicles solitary (perhaps
not always), the peduncles 5-angled, the rachises terete, loosely
and interruptedly cincinniferous, at least the lower branchlets
verticillately congested on stipes about 12 mm. long, the upper with
stipes 3-4 mm. long; pedicels 5 mm. long, articulate above the base,
rusty-puberulent as all the sepals, the inner 4.5 mm. long; petals
nearly 5 mm. long, about 2.5 mm. wide, the upper scales obcordate,
the lower obliquely winged; young fruit subtruncate, tomentose and
with some spreading reddish long trichomes, the endocarp floccose,
the seed at the base of the cells.— Scrap of type at Chicago consists
of the very young fruit; the closely hirsute cells would doubtless
match later those of S. mollis as cited but type of that perhaps less
hirsute. The herbarium name was the same at Madrid by Ruiz,
under another genus. Here might be sought S. rigida Radlk. forma
hirta Radlk., 176, of adjacent Bolivia, the flowering stems only
2 mm. thick, partial petioles margined, leaflets beneath and branch-
lets reddish hirsute-tomentose with crisped trichomes, epidermis
containing mucus; there is a glabrous or nearly glabrous form; see
under S. inflata.
Hudnuco: Vitoc, Ruiz & Pavdn, tjrpe. Bolivia.
324 Field Museum of Natural History — Botany, Vol. XIII
Serjania tenuifolia Radlk. Monogr. Serj. 220. 1875; 132.
Essentially glabrous, indument when present a minute puberu-
lence on the midnerve of the rather coarsely and somewhat double-
serrate leaflets, especially above and in the inflorescences; wood
composite, the peripheral columns suborbicular; branches about 8,
sulcate-costate, not canaliculate, the flowering 3-5 mm. thick;
leaves biternate, thin- or rather firm-membranous; darkening in
herbaria, the ovate-elliptic or suboblong leaflets mostly lineately
pellucid but also somewhat punctate, epidermis containing mucus;
the basally attenuate terminal 10-15 cm. long, about half as wide,
the lateral often only half as large, all acuminate or more usually
obtuse and mucronate or barely acute; petioles emarginate; in-
florescences mostly terminal, the cincinni sessile, scarcely crowded;
pedicels slender to at least 3 mm. long; sepals all pulverulent, 2 mm.
long; petals nearly 3 mm. long; fruit (half mature in type) ovate,
glabrous including endocarp, style pilose, seed near base of cell,
in Woytkowski specimen 2 cm. broad and long, little constricted
at the obscurely veined subtrigonous cells. — Stem climbing about
6 meters, latex white, leaves thin, soft, the nerves paler, petals
greenish-white (Woytkowski).
San Martin: Hera near Moyobamba, Woytkowski 35313 (distr.
as S. brachyptera Radlk.). — Junin: Thickets along Rio Peren^,
Killip & Smith 25138 (probably, but young; distr. as S. inflata). —
Loreto: Yurimaguas, Poeppig, addenda 92, type.
Serjania trirostris Radlk. Notizbl. Bot. Gart. Berlin 6: 149.
1914; 200.
Glabrate or lightly puberulent as the tips of the nearly terete
striate branches; wood composite with 6 peripheral areas; stipules
small, deltoid; petioles all emarginate, the common one 3-4 cm. long;
leaves biternate, the terminal leaflet 9 cm. long, 4 cm. wide, all sub-
sessile, ovate-oblong-lanceolate, remotely and obtusely dentate from
the middle, subcoriaceous, glabrous above except the midnerve, very
sparsely crisped puberulent beneath, olive-green, epunctate, the
epidermis containing mucus; panicles dense, the short recurving
cincinni subsessile, the fruiting pedicels only 3 mm. long, articulate
below the middle; flowers white, 2 mm. long and broad, the sepals
all puberulent, the inner 2 mm. long, the third and fifth connate
below; upper scales with deflexed appendage, long-barbate, the lower
comiform; fruit sparsely crisped pubescent, 2 cm. long, nearly as
thick (doubtless larger at maturity) the cells dorsally produced into
Flora of Peru 325
erect-spreading beaks. — Resembles S. deltoidea with compoundly
divided leaves (Radlkofer). F.M. Neg. 9560.
Ayacucho: Afna, KiUip & Smith 23099? — Loreto: Yarinacocha
on the Rio Ucayali, Tessmann SSll, type; also SSI la. Bolivia.
2. PAULLINIA L.
With the habit of Serjania but flowers usually larger, leaves
often pinnately 5-foliolate and especially fruit a leathery 3-valved
capsule, the often reddish or yellowish 1-3-seeded valves winged or
ribbed, sometimes spiny. Flowers borne in axillary panicles that
are often racemiform or spiciform. Sepals 5, 2 of the inner more or
less united, imbricate. Petals 4, each with a crested hooded scale, the
upp)er ones with a deflexed appendage, barbate below. Disk glands 4.
Seeds more or less arillate. — Spruce noted the native names "Cu-
pana," "guarana" and "timbo" for some Amazonian species.
P. yoco Schultes & Killip and P. cupana HBK. are noteworthy
as containing enough caffein so that a stimulating beverage is made
from them — as noted below — while several other species are of
interest as serving the natives as fish stupefiers and one at least
(P. cururu L.) is in some kinds of curare poison.
Fruits echinate except P. neglecta; leaves all quinate or temate or
rarely 5-foliolate in P. neglecta.
Leaflets all or in part entire; fruits spinose.
Leaves often quinate; stems yellow setulose P. echinata.
Leaves temate and obscurely pubescent as the stems . P. hystriz.
Leaves temate and glabrous as the stems.
Leaflets reticulate veined; fmit firm, densely echinate.
P. paiUlinioides.
Leaflets subclathrate veined ; fmit spongy, sparsely echinate.
P. Sprucei.
Leaflets serrate toward tip; fmits smooth P. neglecta.
Fmits smooth, sometimes alate; leaves not simply quinate or temate
unless sometimes the uppermost temate.
Fmits alate (known); leaflets (Pern) acute or acutely short-
acuminate, unless P. Killipii.
Leaf rachis obviously margined, sometimes narrowly or the
leaves pinnate unless in P. enneaphyUa.
Leaves pinnate or lowest pair of leaflets temate or pinnate.
326 Field Museum of Natural History — Botany, Vol. XIII
Lower leaflets pinnate; plants setose-hispid P. hispida.
Lower leaflets often temate.
Leaflets nearly linear, 4-9 mm. wide P. linearis.
Leaflets lanceolate or ovate, wider.
Leaves pinnate with narrow leaflets.
Stems setulose P. tenera.
Stems glabrous P. fistuhsa.
Leaves bitemate with rather ovate leaflets.
P. serjaniaefolia.
Leaves 5-foliolate.
Stipules conspicuous, persisting; leaflets if dentate not only
at base.
Branches if setulose not angulately; wood simple.
Stems glabrous or minutely setulose.
Stipules often longer than 7 mm.; fruits obovate.
P. caloptera.
Stipules usually shorter than 6 mm.; fruits sub-
orbicular P. serjaniaefolia.
Stems conspicuously setulose P. setosa.
Branches barbate on angles; wood composite.
P. trilatera.
Stipules minute.
Leaflets often 1-2 dentate near base P. bidentata.
Leaflets closely denticulate P. enneaphylla.
Leaf rachis emarginate (see P. enneaphylla).
Indument clearly hispid-setose, often dense on petioles and
upper stems; panicles soHtary.
Stipules fimbriate, conspicuous P. fissistipula.
Stipules entire P. acviangula.
Indument nearly lacking to pilose-tomentose.
Wood simple; panicles soHtary or glomerate.
Panicles solitary (species ill-defined, may be ecotypes).
Stems lightly pubescent or glabrate P. nobilis.
Stems densely pubescent.
Leaves at maturity ample P. gigantea.
Leaves small, the leaflets less than 1 dm. long.
P. dasystachya.
Flora of Peru 327
Panicles glomerate in the leaf axils.
Leaves small, pilose beneath P. martinensis.
Leaves ample, glabrous.
Stipules conspicuous P. Killipii.
Stipules small, subulate P. exalata.
Wood composite; panicles unless uppermost openly fas-
cicled P. Alsmithii.
Fruits exalate (known); leaflets often obtusely short-acuminate
(merely apiculate in P. subroiunda), the rachis often dis-
tinctly wing-margined (in Peru) if leaflets notably acute.
Leaflets usually rounded or obtuse, often mucronate or callose
or if acutish, petioles and rachis not wing-margined.
Leaf rachis as often petioles at least narrowly margined; wood
simple unless in P. pinnata; panicles not glomerate.
Leaves subbitemate, the first pair of leaflets usually
temate P. pachycarpa.
Leaves 5-foliolate or imparipinnate and often 2-4 pairs.
Petioles emarginate; capsules subglobose.
P. spliaerocarpa.
Petioles at least narrowly margined.
Leaflets sharply dentate P. Josecuatrii.
Leaflets subentire or the teeth few, obtuse.
Leaves 5-foliolate; petioles margined.
Stipules minute.
Leaflets subentire; ovary as style puberulent.
Leaflets chartaceous, pale, subentire . P. laeta.
Leaflets rigid, drying dark, 1-2-dentate.
P. subauriculata.
Leaflets remotely dentate; ovary as style gla-
brous P. pinnata.
Stipules about 1 cm. long P. simulans.
Leaves usually with 2-4 pairs of leaflets; petioles
broadly winged.
Ovary glabrous; leaflets 2 pairs P. oUvacea.
Ovary tomentose; leaflets 3-4 pairs. P. pterophylla.
Leaf rachis as petioles emarginate; wood sometimes com-
posite.
Stipules entire, often deciduous or inconspicuous.
328 Field Museum of Natural History — Botany, Vol. XIII
Panicles solitary, sometimes disposed apically, elongate.
Leaf nerves subparallel, usually all less than 1 cm.
distant, venation clathrate; ovary tomentose.
Leaflet acumen entire P. elongata, P. faginea.
Leaflet acumen bidenticulate, short. .P. subrotunda.
Leaf nerves arcuate-spreading, variously distant.
Wood simple; plants often somewhat pubescent.
Leaflets consistently small, terminal rarely and
little longer than 1 dm.; stems striate or
costate; ovary tomentose (known).
Leaflets entire or subentire.
Stipules about 5 mm. long, persisting.
P. itayensis.
Stipules minute, caducous.
Panicles at most about as long as leaves.
Leaflets entire unless 1-2 callose near
base; panicle rachis nearly glabrous;
leaflet acumen broad . . P. uchocacha.
Leaflets subdenticulate toward tip;
panicle rachis puberulent; leaflet
acumen slender P. curvicuspis.
Panicles often exceeding leaves, mostly
borne apically P. capreolata.
Leaflets coarsely dentate, usually to below the
middle.
Leaflets firm; acumen obtuse.
Venation reticulate; panicle rachis to 1.5
mm. thick P. sphaerocarpa.
Venation clathrate; panicle rachis rather
stout P. obovata.
Leaflets diaphanous in herb.; acumen acute.
P. tenuifolia.
Leaflets consistently medium to large, lateral
usually well exceeding 1 dm.; stems often
sulcate; ovary or fruit glabrous.
Leaflets all long-cuneate to long petiolules.
P. cuneata.
Leaflets not cuneate-based except terminal.
Flora of Peru 329
Venation clathrate.
Petals 2.3 mm. long P. yoco.
Petals 4 to nearly 5 mm. long. .P. cupana.
Venation reticulate.
Stipules minute, caducous; stems striate-
costate.
Ovary pubescent; petals 3 mm. long;
acumen broad P. tarapotensis.
Ovary glabrous; petals 4 mm. long;
acumen narrow P. mazanensis.
Stipules to 8 mm. long, persisting; stems
angled, sulcate.
Flowers 3 mm. long, bracts minute;
leaflets acuminate P. simiUans.
Flowers 4.5 mm. long, bracts large;
leaflets obtusish P. Mariae.
Wood composite; plants glabrous or nearly, leaflets
medium size, reticulate; ovary glabrous.
Leaves 5-foliolate, pellucid lineolate ... P. spicata.
Leaves usually temate in part, obscurely pellucid.
P. neglecta.
Panicles glomerate, stout; ovary pubescent.
Leaflets glabrous, ample; wood composite. .P. exalata.
Leaflets pubescent, medium; wood simple.
P. martinensis.
Stipules stellately incised, persisting.
Stems short-hirtellous P. rugosa.
Stems long-hispid P. fissisHpiUa.
Leaflets acute or acutely acuminate and petioles or rachis wing-
margined (P. alata sometimes apiculate or obtusish).
Panicles solitary.
Panicles rather robust, the rachis several mm. thick; leaf-
lets about elliptic.
Leaves pilose beneath; fruits tomentose; wood composite.
P. eriocarpa.
Leaves subglabrous even beneath; fruits glabrous; wood
simple P. imberbis.
330 Field Museum of Natural History — Botany, Vol. XIII
Panicles slender, rachis to 1.5 mm. thick; leaflets oblong-
lanceolate; wood composite P. pinnata.
Panicles glomerate on stems or branches; wood composite.
P. rhizantha, P. alata.
Paullinia acutangula (R. & P.) Pers. Syn. PI. 1: 443. 1805;
324. Semarillaria acutangula R. & P. Prodr. 54. 1794. P. lactescens
Poepp. & Endl. Nov. Gen. & Sp. 3: 37. 1844.
High-climbing with 4-5-angled and thus 4-5-sulcate branches
sordidly or yellowish hirsute-hispid especially on the angles; wood
simple; stipules lanceolate-subulate, about 1.5 cm. long, 3 mm. wide
at base, hirsute beneath, glabrous above as the 5-foliolate-pinnate
leaves; petioles 4-8 cm. long, hirsute, more or less 3-4-angled but
emarginate as also the somewhat shorter rachis, both pilose-hirsute;
leaflets 10-15 (20) cm. long, 3-7 (9) cm. wide, or the terminal
a little larger and cuneate at base, the upper lateral broadly acute,
the lower rounded, all subsessile, oblong-elliptic-lanceolate to rather
broadly elliptic-obovate, cuspidate-acuminate, above the middle
remotely serrate-dentate, the teeth mostly mucronulate, mem-
branous, somewhat hispidulous beneath especially on nerves and
veins, above punctate-pellucid, lacking mucus; panicles solitary,
hirsute, the many-flowered cincinni stiped, the pedicels very short;
inner sepals about 4 mm. long, glabrous, the outer puberulent;
petals obovate; ovary hirsute; capsule obcordate-subrhomboid,
shortly stiped, 2 cm. long, half as wide, hirsute within and without,
the wings abruptly attenuate above the middle, the seeds apparently
glabrous. — Leaflet acumination in type to 12 mm. long, very acute.
Stems about 5 cm. thick, lactescent, the cinnamon-colored bark
rigid-hirsute (Poeppig). F.M. Neg. 23639.
Huanuco: Muiia, Ruiz & Pavdn, type. In woods at Pampayacu,
Poeppig 1758 (type, P. lactescens). — San Martin: San Roque,
Williams 71^7^- — Junin: Cahuapanas on Rio Pichis, Killip & Smith
26760 (det. Standley, P. nohilis). — Loreto: Yurimaguas to Bal-
sapuerto, Killip & Smith 28259 (det. Killip) ; 29050; 28662. Bolivia.
Paullinia alata (R. & P.) G. Don, Gen. Syst. 1: 660. 1831; 242.
Semarillaria alata R. & P. Fl. Peruv. 4: pi. 3^0. 1802.
Younger scandent branches triangular, more or less 6-costate and
sparsely crisped puberulent, the older soon twice as thick (6-8 mm.)
or larger and 3-sulcate; wood composite with 3 peripheral areas;
stipules 4 mm. long, lanceolate-subulate; petioles and leaf-rachises
winged, the former 4-6 cm. long, the latter 2-3 cm. long, the wings
Flora of Peru 331
each side 1.5-2.5 (-5) mm. wide, petiolules scarcely 2 mm. long,
leaves 5-foliolate-pinnate, the leaflets in type oblong-lanceolate but
apparently often broadly elliptic in variants, remotely dentate
(Peruvian) terminal 8-13 cm. X 3-5 cm., rest smaller or subentire,
membranous, nitidulous both sides, above on nerves, all over beneath
sparsely pubescent (rarely rather densely) and in the nerve axils
pilose, in tjrpe glabrescent in age, microscopically glandular and
subpellucid-lineolate, the epidermis lacking mucus but lower sides
sparsely crystallophorous; panicles 1.5-2 cm. long, appressed
yellowish-puberulent, borne on the branches, often fasciculate-
glomerate, the lower cincinni stiped, the upper sessile, the pedicels
3-4 mm. long, articulate above the middle; flowers 3.5 mm. long;
2 outer sepals less than half as long as the subglabrous inner, rotund
ovate, appressed puberulent; upper petals obovate-oblong, much
longer clawed than the lower oblong ones; ovary yellowish pilose;
capsule apparently subglobose, in var. loretana obovoid, about 2 cm.
long, shortly stiped, sparsely puberulent, the seeds medially arillate.
— See also P. rhizantha which may be distinct, the type of P. alata
having lanceolate leaflets. Stems to 6 meters tall and 2 cm. in
diameter; red fruit valves with black lustrous white-arilled seed
(Stork, Horton & Vargas).
All of the material from Loreto has much larger leaves but
apparently is not otherwise different; it may represent P. rhizantha
but I hesitate to use that name as the description does not quite
accord and some specimens approach the typical form; for con-
venience this then may be designated P. alata, var. loretana Macbr.,
var. nov., foliolis late ellipticis vel paullo obovatis interdum fere
1.5 dm. longis, 6-8 cm. latis. More distinctive is a collection that is
softly pubescent and may prove to be distinct but in the absence of
fruit may be treated as var. pubens Macbr., var. nov., ramulis
foliisque praecipue subtus molliter pubescentibus cum pilis flavis;
foliolis late ellipticis vel subobovatis 5-8 cm. latis, 10-12 cm. longis,
solum mucronato-apiculatis. P. largifolia Radlk., 241, and P.
densiflora Smith, 241, of the Amazon and Colombia respectively,
have temate leaves, the former with the panicles at defoliate nodes,
the latter in the axils of young leaves. P. fascicukUa Radlk., 242,
is more like P. alata var. loretana but has lustrous membranous
nearly glabrous leaves.
According to Kanehira used as a fish poison.
San Martin: Tarapoto, WiUiams 6092. Tingo Maria, Allard
2051 S. — Hudnuco: Chicoplaya, Ruiz & Pav&n, type. Pampayacu,
332 Field Museum of Natural History — Botany, Vol. XIII
Kanehira US. Prov. Huamalies, Stork & Horton 9564^ ^YP^, var.
pubens. Zepelacio, Klug 3263 (more pilose). — Loreto: Caballo-
Cocha, Williams 2339, type, var. loretana. Balsapuerto, Klug 2879
(det. Standley). Iquitos, Killip & Smith 27^21. Yurimaguas,
Williams J/Sl 7. Florida, Klug 1 993. — Cuzco : Deep woods, Echarate,
Goodspeed Exped. 104-5^ (det. Standley). Brazil to Panama.
"Macote" (Kanehira).
Paullinia Alsmithii Macbr., sp. nov.
Scandens fruticosa; ramis valde sulcato-costatis ramulis petiolis
paniculisque crispe pulverulento-strigillosis floriferis circa 7 mm.
crassis (corpus lignosum compositum); stipulis subulatis 6-10 mm.
longis, 0.5-1 mm. latis; petiolis costato-striatis et rhachis emargin-
atis, petiolulis 5-10 mm. longis; foliolis 5 late oblongo- vel ovato-
ellipticis basi plus minusve inequaliter rotundatis, apice breviter
subacute acuminatis plerumque 8-15 cm. longis, 4-7.5 cm. latis char-
taceis pellucido-punctatis-lineolatis remote repando-denticulatis
subclathrato-reticulato-venosis supra glabris subtus molliter puber-
ulo-pilosis; paniculis racemiformis 4-7 cm. longis 1-5 in axillis folio-
rum sessilibus vel superioribus interdum longe pedunculatis; bracteis
et bracteolis minutis; cincinnis breviter stipitatis remotis; pedicellis
3-5 mm. longis basi articulatis; sepalis subglabris interioribus circa
4 mm. longis; petalis oblongo-ellipticis; capsulis subrotundatis triala-
tis circa 1.5 cm. longis et latis glabratis, abrupte plus minusve stipi-
tatis. — Seems to be well-marked, possibly related to P. nobilis.
Loreto: Iquitos, Killip & Smith 26939, type; 27170; 27290; 2720k-
Paullinia bidentata Radlk. Monogr. Paull. 195. 1895; 276.
Nearly glabrous (but inflorescence unknown) scandent shrub
with terete branches, the wood simple; stipules minute, deltoid,
axillary, geminate, pilose as margined petioles; somewhat winged
leaf rachises, each about 3 cm. long, the 5-foliolate-pinnate leaves
as broad as long, about 18 cm., the upper leaflets 11-12 cm. long,
3 cm. wide, the lower 8 cm. long, 2.5 cm. wide, all lanceolate, gradu-
ally acuminate, the upper acute at base, sessile, the lower subtruncate,
subauriculate-bidentate, shortly petioled, pinnate-nerved, the nerves
arcuate-ascending, reticulate-veined, membranous, sparsely barbate
in the nerve axils, epunctate, containing mucus. — Placed by the
author with P. suhauriculata. F.M. Neg. 31037.
Cajamarca: Tambillo, Jelski kl2, type.
Flora of Peru 333
Paullinia caloptera Radlk. in Engl. & Prantl, Pflanzenfam.
3, 5: 304. 1895; 329. P. WiUiamsi Macbr. Field Mus. Bot. 11: 27.
1931.
Scandent, subglabrous, the branches triangular, the leaves
5-foliolate-pinnate with 4-sulcate or somewhat margined petioles
2-5 cm. long, the prominent linear-lanceolate acuminate stipules
7-17 mm. long; rachis often conspicuously winged, 5 mm. wide, the
sparsely pulverulent and ciliolate-hirsutulous leaflets ovate-lanceo-
late, subsessile or shortly petioled, acutely acuminate, nearly entire
or remotely 3-5-dentate especially toward the tip, laxly reticulate,
dull both sides, minutely punctate beneath, 3-3.5 cm. wide, 8-10
cm. long; inflorescence solitary, 2-3 cm. long, softly puberulent, the
very short ramuli only 3-5-flowered; sepals minutely pulverulent,
the outer 1.5, inner 3 mm. long; petals pilose at base. — With the
appearance of P. laeta and P. subauriculata but the leaflets acutish-
acuminate, the stipules large, the stems trigonous. The Peruvian
plant is slightly more puberulent than the typical form from Brazil
and Venezuela and when fruits are known may prove to be dis-
tinguishable at least varietally. P. emetica Schultes, Caldasia 2:
420. 1944, of southeastern Colombia has thin-membranous leaves
and erect lax inflorescences to 2 dm. long; an infusion of the leaves is
known to have been used as an emetic. Illustrated, Mart. Fl. Bras.
13, pt. 3: pi. 89.
Loreto: Rio Nanay, Williams 1196 (type, P. WiUiamsi). Mishu-
yacu, Kliig 91 S; IJ^SS. Timbuchi, Rio Nanay, Williams 982.
Venezuela; Amazonian Brazil. "Sapu-wasca" (Williams).
Paullinia capreolata (Aublet) Radlk. Monogr. Serj. 70. 1875;
300. Enourea capreolata Aublet, PI. Guian. 1 : 587. pi 235. 1775.
Liana, essentially glabrous except for the puberulent younger
branches, these terete, lenticellate, soon 3-4 mm. thick, and the
solitary or sometimes paniculate panicles, these 1.5-2.5 dm. long,
slender, (rachis 1-2 mm. thick), laxly flowered, the cincinni sessile
or shortly stiped; wood simple; petioles and rachis emarginate;
leaves pinnately 5-foliolate, the upper leaflets elliptic or elliptic-
lanceolate, the lower ovate, all subentire or rarely denticulate
apically, often undulate, shortly j)etiolulate or subsessile, 5-10 cm.
long, obtusely acuminate, rounded or acutish at base or the terminal
basally acute, lustrous both sides, micro-glandular, epidermis con-
taining mucus, laxly transversely veined, few-nerved, chartaceous or
subcoriaceous; bracts and bractlets minute, the pedicels about 2 mm.
334 Field Museum of Natural History — Botany, Vol. XIII
long, articulate medially; outer sepals scarcely a third as long as the
more or less connate subpetaloid inner, these 2.5 mm. long, all
ashy-tomentulose without; petals oval-obovate, the scales more or
less barbate including the bifid appendage; filaments complanate,
densely reddish-long pilose, the anthers glabrous; capsules depressed
globose, subsessile, glabrate without, lanate within, about 1.5 cm.
long, nearly 2 cm. broad; seed surrounded by the fleshy farinaceous
aril. — The branchlets of the Loreto specimens are notably lenti-
cellate; the minutely and sparsely puberulent subglobose fruits are
about 12 mm. in diameter, stipes 3 mm. long, pedicels 4 mm. long.
P. firma Radlk,, 299, is similar but the leaves are rigid coriaceous.
Loreto: Mishuyacu, Klug 695; 759; 870. Florida, Klug 1979
(det. Standley). Ucayali, Tessmann Jt.162. To Venezuela and
British Guiana. "Tingui" (Amazonian), "enourou" (Guiana).
Paullinia cuneata Radlk. Notizbl. Bot. Gart. Berlin 6: 150.
1914; 281.
Nearly glabrous, suffrutescent, the younger branches sulcate,
sparsely puberulent, the wood simple, the leaves 5-foliolate-pinnate;
stipules linear, lightly pilose, to 15 mm. long; petioles as rachis
emarginate, the former a dm. long or longer, the lower leaflets with
petiolules to 14 mm. long, the upper subsessile, obovate-cuneate,
10 cm. long or longer, 7-9 cm. wide, very shortly obtuse-acuminate,
remotely subrepand-dentate from the middle, the short teeth
spreading, nerves obliquely erect, prominent both sides, lustrous,
glabrous, epunctate; panicles solitary, dense, sessile; bracts subulate,
2 mm. long; outer sepals minutely puberulent, the broadly obovate
inner 2 mm. long, glandular-ciliolate, otherwise glabrous; petals
oblong, 2.5 mm. long, the scales with puberulent deflexed appendage.
— Similar to P. cupana HBK., 281, scandent or suberect shrub of
Venezuela and Amazonian Brazil — possibly also Peru — source of
"Guarana" but that with lateral leaflets ovate, rounded at base,
scabrous-glandular beneath, sepals sparsely setulose-pilose. — Type
from Cobijn, Bolivia (Ule 9571) near Madre de Dios.
Peru (no doubt; cf. note above). Bolivia; Brazil?
Paullinia cupana HBK. Nov. Gen. & Sp. 5: 117. 1821; 281.
Scandent or suberect liana, the apically brownish-pilose branches
soon glabrous and deeply 4-5 sulcate, the flowering 4-8 mm, thick,
the wood simple; stipules 2-3 mm. long, petioles and rachis emar-
ginate, canaliculate above, convex and lightly striate beneath;
Flora of Peru 335
leaflets 5, the upper oblong, the lower ovate, the short acumen more
or less obtuse, the terminal leaflet acute or subcuneate at base, the
lateral rounded or more or less petiolulate and remotely subrepand-
dentate, the dentations sometimes obscure, mostly prominent and
obtuse, usually 1-2 dm. long, 4.5 to 9 cm. wide, coriaceous, obscurely
clathrate-veined, glabrate both sides, obsoletely pellucid-punctate,
epidermis lacking mucus; panicles solitary, sessile or peduncled, the
subvillous rachis about 2 mm. thick, the rather remote cincinni
sessile, few-flowered; bracts subulate, 1-1.5 mm. long, pedicels
articulate below the middle, 4-5 mm. long; inner sepals 3 mm. long,
submembranous, all laxly hirsutulous, free; petals oblong, 5 mm.
long; filaments pilose, anthers glabrous; ovary glabrous, ~ stiped,
ellipsoid as the apiculate capsule, this brownish-tomentose within,
2-3.5 cm. long, deep red at maturity, the stipe finally 6-8 mm. long,
seed about 12 mm. long, glabrous. — Var. sorbilis (Mart.) Ducke,
Rodriguesia 3: 155-156. 1937 and Archiv. Inst. Biol. Veg. Rio Jan.
4: 47. 1938, differs in having tendrils, especially in the inflorescence,
often subentire leaflets, slightly smaller flowers, much smaller fruits,
these ovoid or spheroid, 15-18 mm. long, brilliant red, lustrous
(Ducke). Here may be mentioned Mexia 6297 with fusiform ellip-
soid fruits about 3 cm. long, leaflets remotely dentate, 1-1.5 dm.
long, referred to P. tarapotensis by Standley but with the venation
of P. cupana: it may not be related here but the native names
"Ycanchem" (Huitoto) and "lucumia" may be recorded. Illus-
trated, Radlkofer, I.e. 282; Ducke, Rodriguesia I.e.
The seeds of P. cupana and var. sorbilis are pulverized and mixed
with cassava flour (Manihot esculenta Crantz) and formed into
molds which are known as "pasta guarana" and dissolved as desired
in hot or cold water; the caffein content of the "pasta" is three to
six per cent, tannin two to three per cent so the beverage is astringent
(Schultes). Cf. Radlkofer I.e. for bibliography pertaining to medi-
cinal use.
Peru (possibly, see note above). Colombia to Venezuela and
Amazonian Brazil. "Cupana," "guarana."
Paullinia curvicuspis Radlk. Monogr. Paull. 237. 1895; 301.
Branches terete, substriate, sordidly pulverulent, finally sub-
glabrous, the wood simple, the leaves 5-foliolate-pinnate; petioles
and rachises emarginate, the former 3-6 cm. long, the petiolules 2-5
mm. long; leaflets 5-10 cm. long, elliptic-lanceolate, with narrow,
elongate, obtusish curved acumen, the terminal cuneate at base.
336 Field Museum of Natural History — Botany, Vol. XIII
the lateral subacute, all remotely denticulate above the middle,
petiolulate, few-nerved, narrowly transversely veined, subchar-
taceous, nitidulous and glabrous both sides but often barbulate in the
nerve axils beneath, microscopically glandular, containing mucus;
panicles solitary, the remote cincinni shortly stiped or subsessile,
the pedicels 2 mm. long, medially articulate; flowers medium in size,
all sepals canescent puberulent without; scales villous, the upper
bifid; filaments with some long reddish trichomes; otherwise known.
Puno: San Govdn, (Lechler 2358, type; 8277).
Paullinia dasystachya Radlk. Monogr. Paull. 270. 1895; 325.
Closely allied to and apparently not specifically distinct from
P. gigantea but the pubescence hirsute-tomentose; petioles 2-6 cm.
long, tomentose; leaflets 5-8 cm. long, 2-3.5 cm. wide, ovate-oblong,
acute or acutish, chartaceous, sparsely and minutely pubescent
above, slightly pellucid punctate-lineolate, coarsely serrate-dentate,
the few teeth obtuse; panicles yellowish-tomentose, cincinni long-
stiped, the fruiting pedicels about 4 mm. long, articulate at base:
sepals tomentose; capsule 3-winged, emarginate, the stipe about
3 mm. long, hirtellous-tomentulose without, pilose within, the seeds
lightly pilose. — Mexia 6773 from near Guayaquil, determined as P.
quitensis Radlk. 325, is probably better placed here. It is another
segregate, similar in pubescence but ovate leaflets remotely or
obsoletely dentate, epunctate, the tomentose capsule truncate on
stipe 1 cm. long.
Loreto: Florida, Rio Putumayo, Klug 2091; 2186? (det. Standley,
P. coloptera). Iquitos, Killip & Smith 27131. Pebas, Williams
18J^3. Bolivia; Ecuador.
Paullinia echinata Huber, Bol. Mus. Goeldi 4: 582. Aug. 1905;
303. P. echinata Radlk. Bot. Jahrb. 37: 153. Sept. 1905.
More or less reddish setulose high climbing liana, the pubescence
extending to the petioles, leaves both sides, especially on the nerves
and subclathrate veins, and the 3-5 fasciculately aggregate panicles
that are borne on the older branches; petioles 1-2 dm. long, subterete,
the sulcate petiolules 1.5-2.5 cm. long; leaves ternate or quinate,
the obovate leaflets shortly and narrowly acuminate, 15-25 cm. long,
6-10 cm. wide, entire or appearing repand when unequally revolute,
rigid-coriaceous, in age merely somewhat scabrous above, nitidulous
both sides, containing mucus; panicles 1-2 dm. long, the sessile
cincinni 4-6-flowered, the pedicels medially articulate, 3 mm. long,
Flora of Peru 337
8 in fruit; sepals tomentulose, the inner 4.5 mm. long; petals obovate
oval, glandular both sides; filaments rufous villous; capsule ellipsoid-
globose, on stipe 5-7 mm. long, densely echinate and setulose, the
spines rather rigid, about 4 mm. long. — Flowers pale rose (Ule).
F.M. Neg. 5597.
Loreto: Cerro de Canchahuaya, common in woods, on the
Ucayali, Huber IW, type. Yurimaguas, Ule 6865; Kuhlmann
209S1; Williams S992; Jt952. Puerto Arturo, KiUip & Smith 27880.
Brazil.
Paullinia elongata Radlk. Monogr. Paull. 238. 1895; 302.
Scandent terete striate branches yellowish-tomentulose with
a hirtellous indument; wood simple; stipules conspicuous, elliptic-
lanceolate, coriaceous, tomentose both sides, about 5 mm. long;
petioles and rachises emarginate, the former 5-8 cm. long, the
petiolules 1-2 mm. long; leaves 5-foliolate-pinnate, the leaflets
elliptic, obtusely acuminate, the terminal cuneate at base, the others
acute, 7-10 cm. long, about half as wide, all coarsely serrate-dentate,
coriaceous-chartaceous, clathrate-venose, glabrous above except for
the puberulent midnerve, yellowish puberulent-pubescent, especially
nerves, and microscopically glandular beneath, epidermis containing
mucus; panicles solitary, the bracts about 2 mm. long, the cincinni
sessile; pedicels 2 mm. long; sepals tomentulose, the inner nearly
4 mm. long; petals oblong; scales villous, the upper bifid with de-
flexed appendage; filaments whitish pilose. — This collection only in
flower was placed by the monographer with P. clathrata Radlk., 302,
of the Rfo Negro, with subentire leaflets, the pubescence less hirtel-
lous, both forms in contrast to the "glabrous crenate-dentate"
leaflets of P. faginea which compare; from material that has accumu-
lated it is not clear that these differences are not highly variable.
F.M. Neg. 23643.
Hudnuco: Vitoc, Ruiz & Pavdn, type.
Paullinia enneaphylla (R. & P.) G. Don, Gen. Syst. 1: 662.
1831; 330. Semarillaria enneaphylla R. & P. Fl. Peruv. 4: pi. SUl.
1802.
Obsoletely lenticellate scandent striate subterete branches
puberulent, finally glabrescent; wood simple; stipules small, deltoid;
petioles and rachis scarcely if at all margined, both about 3-4 cm.
long; leaves bitemate, the leaflets elliptic-lanceolate, acute both ends
(terminal, acuminate), 5-9 cm. long, 2-4 cm. wide, subsessile, rather
338 Field Museum of Natural History— Botany, Vol. XIII
densely serrate-dentate, chartaceous, glabrous above, barbate in the
nerve axils beneath, pellucid-punctate, lacking mucus; panicles
solitary, puberulent, the cincinni in flower subsessile, in fruit on
stipes 3-4 mm. long, the pedicels 1-2 mm. long, articulate at base;
flowers pubescent, whitish, the oval inner sepals 3 mm. long, all
sparsely glandular-ciliolate; petals obovate-oblong; ovary appressed
tomentose; capsule 3-winged, obcordate, apiculate, about 1,5 cm.
long, 1.8 cm. wide, glabrous without, puberulent within, the globose-
obovoid seeds glabrous, arillate nearly to the middle.— Capsule
sessile, that is estipitate, in contrast to that of the more northern
P. fuscesens HBK., 330, and P. navicularis Radlk., 335, both perhaps
variants found as near as Ecuador; both are more pubescent, con-
taining mucus, the leaf rachis more or less margined, barely in the
latter, the capsule stipe of the former short, of the latter 3 mm. long.
F.M. Neg. 29690.
Hudnuco: Chinchao, Ruiz & Pavdn, type. — Cajamarca: Between
Huambos and Montan, 2,500 meters, Weberbauer U21U-
Paullinia eriocarpa Tr. & Planch. Ann. Sci. Nat. s^r. 4. 18:
353. 1862; 265. P. eriantha Benth. ex Radlk. Monogr. Serj. 75.
1875.
Scandent branches glabrate to hirsute, 4-5-angulate-sulcate, the
wood composite, the peripheral areas 2 or 3; stipules appressed
pilose, about 8 mm. long, 3-5 mm. wide; petioles 2-5 cm. long, the
rachises shorter, both with wing-margins 4-6 mm. wide; leaves
5-foliolate-pinnate, sometimes temate, the leaflets 5-15 (20) cm.
long, 2.5-6 (9) cm. wide, elliptic-lanceolate, upper cuneate at base,
lower acutish, all acuminate, remotely repand-dentate above the
middle, shortly petiolulate or subsessile, subcoriaceous, typically
glabrous or often densely pilose beneath, lacking mucus but sparsely
crystallophorous; panicles solitary with short even headlike flowering
portion only 2-4 cm. long, the bracts unusually broad, scariose,
7 mm. long, nearly as wide, sericeous beneath; cincinni sessile, the
flowers subsessile with sericeous sepals, the broadly ovate inner
7-8 mm. long, 5-6 mm. wide; petals tomentulose below, sepaloid,
8-9 mm. long, 3-5 mm. wide; capsule Hgneous, about 2.5 (3) cm.
long, 1.7 cm. wide, ovoid, shortly acuminate, yellowish-tomentose,
the long-arillate seed lanate. — The more northern and eastern
ranging P. leiocarpa Griseb., 264, has leaves merely barbate in the
nerve axils and subulate bracts; the southeastern Colombian P.
splendida Schultes, Caldasia 2: 421. 1944, said by author to belong
to section Pleurotechus Radlk. but apparently comparable to P. ,
Flora of Peru 339
eriocarpa, has bracts about 2 mm. long and broad, papjrraceous
leaflets to 2 dm. long; KiUip & Smith 2H71 from Rio Itaya might
be referable to it. F.M. Neg. 5598.
San Martfn: Tarapoto, Spruce UU15 (type, P. eriantha); WiUiams
58It9; 6770. — Loreto: Yurimaguas, WiUiams 1^802. To Panama.
Paullinia exalata Radlk. Bot. Jahrb. 37: 150. 1905; 244.
Glabrous liana even to the sepals or a few trichomes in the axils
of the nerves on the 5-foliolate-pinnate leaves beneath; older stem
3-angled, enlarged at the cirrose nodes, the triangular canaliculate
branches 6-costate; wood composite with 3 peripheral areas; stipules
subulate, 3 mm. long; petioles and leaf-rachises not winged, the
former 6-16 cm. long, the latter 4-6 cm. long, the terminal leaflet
with petiolule 1 cm. long, 16-18 cm. long, the elliptic-ovate shortly
acuminate lateral leaflets little smaller, all obtusely and remotely
subrepand-dentate, chartaceo-subcoriaceous, opaque both sides,
lateral nerves prominent beneath, lacking mucus; panicles glomerate
on older stems, 2-3 cm. long, puberulent, the lower cincinni shortly
stiped, the pedicels 5-6 (-8 in fruit) mm. long, articulate above the
middle; flowers 5 mm. long, nearly glabrous except for the appressed
pilose ovary, the outer sepals with a few trichomes, the inner gla-
brous, 5 mm. long; petals 5.5 mm. long, the upper scales barbate
and parted, the lower bifid and aliform; ovary pilose; capsule gla-
brate, 8 cm. long, the seed part 1.5 cm. long and wide, the valves
little enlarged, the lustrous seeds 1.5 cm. long. F.M. Neg. 5599.
Junin: La Merced, 1,000 meters, Weherhauer 1910; 282. Brazil.
Paullinia faginea (Tr. & Planch.) Radlk. Monogr. Serj. 76.
1875; 301. Enourea faginea Tr. & Planch., Ann. Sci. Nat. s6r. 4.
18: 379. 1862.
Similar to P. elongata especially to the var. pubescens Cuatr. of
Colombia but typically less pubescent; flowering branches terete,
reddish-puberulent, 3-4 mm. thick; wood simple; petioles and rachis
emarginate; leaflets 5, oval, obtusely acuminate, the terminal acute
at base, the lateral acutish or subrounded, all with petiolules 2-4 mm.
long, remotely crenate-dentate above the middle, to about 12 cm.
long, 5 cm. wide, chartaceous or submembranous, multinerved,
narrowly and obliquely clathrate veined, glabrous or the midnerve
above and the nerves beneath somewhat puberulent and barbate
in the nerve axils, epidermis containing mucus; panicles solitary or
paniculate, the upper much exceeding the leaves, shortly peduncled;
340 Field Museum of Natural History — Botany, Vol. XIII
rachis about 2 mm. thick, puberulent-tomentulose, laxly flowered,
the sessile cincinni contracted, the pedicels only about 1 mm. long;
inner sepals 3 mm. long, tomentulose, subpetaloid; petals oblong;
filament trichomes yellowish; ovary globose, sessile, reddish-
tomentose. — P. clathrata Radlk., 302, seems to be a form with more
broadly elliptic leaflets, the dentations less pronounced, and may be
represented by King 2001. F.M. Neg. 23644.
Loreto: Rio Mazdn, Jos^ Schunke 37 (det. Standley). Florida,
Rio Putumayo, King 2001 (det. Killip) ; Klug 2323. To Colombia
and Amazonian Brazil.
Paullinia fissistipula Macbr., sp. nov.
Scandens fruticosa; ramis thyrsigeris fere teretibus leviter cos-
tatis 1 cm. crassis utrinque sed praecipue in costis cum petiolis
rhachisque emarginatis pedunculisque 3.5-15 cm. longis conspicue
cum pilis rufis ad 2.5 mm. longis setoso-hispidis; corpus lignosum
simplex; stipulis subrotundatis ad medium stellato-incisis adpresse
hirsutis 2-2.5 cm. longis; petiolis striatis, petiolulis 2 mm. longis;
foliolis 5 late ellipticis vel interdum paullo obovatis breviter acuteque
acuminatis 12-15 cm. longis, 6-8 cm. latis, ad apicem repando-
dentatis (dentibus nervo excurrente calloso-apiculatis glabris) sub-
coriaceis pellucido-punctatis tenuiter clathrato-reticulato-venosis
supra (nerviis 12-15 approximatis exceptis) glabris, subtus sparse
setulosis; paniculis densifloris circa 1 dm. longis, 2 cm. latis; bracteis
ubique adpresse pilosis oblongo-acuminatis ad 1 cm. longis; pedicellis
ad 8 mm. longis supra mediam articulatis; sepalis adpresse cinereo-
pilosis interioribus circa 3.5 mm. longis; petalis circa 4 mm. longis;
ovario rufo-hirsuto. — Suggests the Brazilian or Amazonian P. ruhi-
ginosa Camb., 266, and P. stipularis Benth., 267, both with oblongish
leaflets and linear-subulate bracts, but probably is related to P.
fimbriata Radlk., 288, of Central America with much shorter denser
indument, larger flowers.
Loreto: Balsapuerto, Klug 3056, type (det. Standley, cf. P.
gigantea) .
Paullinia fistulosa Radlk. Monogr. Paull. 259. 1895; 319.
Completely glabrous even the short glomerate panicles borne at
the defoliate nodes of the older branches, these scandent, 5-angled
to subterete, and somewhat fistulose by the enlarged medular
cavities, the wood simple; stipules linear-lanceolate, falcately re-
curving, nearly 1 cm. long; petioles all emarginate, the common
Flora of Peru 341
7-10 cm. long, sulcate, costate-striate, the lateral 1.5-2 cm., the
rachis-segments marginate-winged above, 2.5-6 cm. long; leaves
imparipinnate, the lower of the 3-5 pairs of lanceolate long-acumi-
nate leaflets temate, the terminal leaflet attenuate at base, the
lateral acutish, sessile, all entire or with a more or less prominent
tooth on each edge near the base, membranous^ lacking mucus,
8-12 cm. long, 2.5-3 cm. wide; panicles 1-1.5 cm. long, the fruiting
pedicels about 3 mm. long; inner sepals 3.5 mm. long; capsules
obovoid, conspicuously attenuate into stipe 5 mm. long, the wings
above 2-3 mm. wide, glabrous without, pubescent within, the
somewhat pilose seed one-third arillate. — P. medullosa Radlk., 319,
of Brazil, the leaflets somewhat acuminate, the lateral petioles
marginate-winged above, is probably a variant. F.M. Neg. 36026.
Loreto: Mission de Sarayacu, Castelnau, tjrpe.
Paullinia gigantea Poepp. & Endl. Nov. Gen. & Sp. 3: 37.
1844; 324. P. quitensis Radlk. Monogr. Paull. 269. 1895, at least
as to Peru.
Resembles the related P. aciUangula but more robust; stipules
scarious, gradually subulate-acute from the base, about 3 cm. long,
nearly a third as wide; petioles 15-25 cm. long, subterete; leaflets
15-30 cm. long, 10-18 cm. wide, or apparently on newer branchlets
often only a dm. long, less than half as wide, elliptic or the lower
broadly ovate, the lateral rounded at base, all remotely repand-
dentate or sometimes subentire, sessile or shortly petiolulate, char-
taceous, clathrate- venose, typically epunctate; cincinni sessile or
shortly stiped, in t3rpe few-flowered, the pedicels 4-5 mm. long;
inner sepals about 4 mm. long, tomentulose without; petals oblong-
elliptic; capsule said to be subglobose with short wings, hirsute. — As
interpreted here the cited material includes plants showing consider-
able variation in degree of pubescence — nearly glabrous to shortly
hirsute-tomentose — shape and size of leaves; this varies, it seems,
even on the same plant. The capsules, as seen on a few of the
collections, are puberulent-hirsutulous, wings narrowed to attenuate
more or less stipitate base, narrower than capsule, to 3 cm. long,
2-2.5 cm. wide including wings. The Schunke specimen shows
leaves from near top of stem and from below; the former are only
a third as large; the stipules also run from lanceolate-subulate, 1 cm.
long, to ovate-lanceolate, 2.5 cm. long, nearly 1 cm. wide. See
P. nohilis, apparently a less pubescent state, and P. dasystachya,
a tomentose form. Scandent to the tops of trees, probably over
342 Field Museum of Natural History— Botany, Vol. XIII
30 meters long, very woody, a dm. thick (Poeppig). F.M. Negs.
5601; 5619 (P. quitensis).
San Martin: Juanjui, Klug ^292 (det. Standley, P. nohilis). —
Loreto: Iquitos, Killip & Smith 27230. Gamitanicocha, Rio Mazdn,
Josi Schunke 2J^2. La Victoria, Williams 2698; 3127; 2898. Yuri-
maguas to Balsapuerto, Killip & Smith 2828j^.
Paullinia hispida Jacq. Hort. Schoenbr. 3: 9. pi. 268. 1798; 321.
Liana, notable among Peruvian species by the conspicuous long
rigid yellowish trichomes that densely clothe the younger 4-5 sulcate
branches, petioles and petiolules; wood simple; stipules scarious,
setose-ciliate, ovate-lanceolate, acute or acuminate, 2-3 cm. long,
5-15 mm. broad; rachis more or less margined in the upper segments;
leaves subbipinnate with 4-6 pairs of lanceolate or lanceolate-
elliptic acute or acuminate subsessile or shortly petiolulate remotely
dentate subchartaceous subglabrous pellucid-punctate leaflets, 6-20
cm. long, 2-6 cm. wide, the lowest pair of leaflets 5-foliolate, the
second temate; panicles 5-12 cm. long, fasciculate at defoliate nodes
and in the leaf axils, sometimes solitary and elongate in the axils
of upper leaves; peduncles with two tendrils; cincinni more or less
stiped; bracts and bractlets minute, pedicels 2-3 mm. long; sepals
glabrous, the oval inner ones 4 mm. long, half as wide; petals oval-
oblong; filaments shortly pilose at base; ovary hispid tomentulose;
capsules with stipe nearly 2.5 cm. long, 12-14 mm. wide including
wings, these about 5 mm. wide at excised tip, strongly narrowed to
stipe, glabrous, reddish, pilose within, seed nearly glabrous.
Loreto: Iquitos, Killip & Smith 27 ^^^3. Soledad, Killip &
Smith 29686. Mishuyacu, Klug 752. Alto Rio Itaya, Williams
3353. To Panama and Venezuela.
Paullinia hystrix Radlk. Notizbl. Bot. Gart. Berlin 6: 152.
1914; 304.
Allied to P. echinata and in general similar but merely scabrellous
or somewhat hirtellous on the leaf nerves, the elliptic leaflets with
1 or 2 callose obtuse teeth, the panicles solitary on leafy branches,
rusty villous, to 3 dm. long; inner sepals 5 mm. long; petals oblong,
6 mm. long; capsule puberulent and densely echinate with flexible
spines 6 mm. long or shorter, the stipe 2 mm. long. — The other
related species are P. paullinioides (Spruce) Radlk., 303, of Brazil
and P. granatensis (Planch. & Lind.) Radlk., 304, of Colombia, the
former with long-acuminate reticulately veined glabrous leaflets.
Flora of Peru 343
capsule stipe 4 mm. long, spines 1 cm. long, the latter with leaflets
of P. hystriz but glabrous except barbellate in nerve axils, the
capsule subsessile, the spines about 1 cm. long. These species, all
as yet known from only a few collections, may be found to be
variants of one and distributed in Peru, particularly in Loreto and
Madre de Dios. F.M. Neg. 5604.
Rfo Acre: Seringal San Francisco, Ule 9561, t)rpe.
FaulHnia imberbis Radlk. Monogr. Paull. 177. 1895 et Engl.
& Prantl, Pflanzenfam. 3, 5: 304. 1895; 263.
Glabrous scandent shrub, the 5-foliolate-pinnate leaves not even
barbate in the nerve axils beneath, only the solitary panicles min-
utely tomentulose; branches 4-5-costate, the wood simple; upper-
most leaves sometimes temate; leaflets elliptic-lanceolate, acuminate,
subrotund or acute at base, the terminal cuneate, 8-15 cm. long,
3-7 cm. wide, all shortly petiolulate, above the middle remotely
serrate or subrepand, subcoriaceous, lacking mucus, sparsely
crystallophorous; petioles (2-8 cm. long) and rachises broadly
winged, the wings 3-5 mm. wide each side; stipules linear-lanceolate,
5-12 mm. long; panicles solitary, cincinni sessile, pedicels 2-3 mm.
long, articulate above the middle; sepals ashy tomentulose both
sides, the outer suborbicular; petals about 5 mm. long, 2 mm. wide;
capsule pyriform, shortly stiped (stipe less than 1 cm.), red, the
seeds 9-12 mm. long. — P. ingaefolia Rusby, Mem. N. Y. Bot. Gard.
7: 291. 1927, from northern Bolivia is apparently similar but larger
in all parts; if a valid species it requires a new name. Illustrated,
Mart. Fl. Bras. 13, pt. 3. pi. 81.
San Martin: Tarapoto, Ule 686 j^. — Loreto: Cerro de Cancha-
huaya, Ucayali (Huber 1j^7, doubtful, fide Radlk.). Caballo-Cocha,
Williams 2363. Brazil; Guiana.
PauUinia itayensis Macbr., sp. nov.
Fruticosa; ramis teretibus obscure striatis parce piloso-puberu-
lentis ut etiam petiolis inflorescentiisque; stipulis lineari-lanceolatis
circa 5 mm. longis; petiolis rhachisque emarginatis; foliolis 5,
oblongo-ellipticis breviter et obtuse acuminatis, basi subacutis,
integris chsirtaceis praeter nervis supra utrinque glabris et glau-
centibus, tenuiter reticulato-venosis, supra nitidulis, circa 4 cm.
latis et 8 vel 10 cm. longis, breviter petiolulatis; pedicellis fructiferis
1 cm. longis; capsulis subglobosis apiculatis breviter stipitatis sub-
dense strigosis subcoriaceis ecostatis circa 1 cm. crassis. — Section
344 Field Museum of Natural History — Botany, Vol. XIII
Enourea. Imperfectly known and identity obscure but given a
name for convenience in the vicinity of P. curvicuspis.
Loreto: Along Rio Itaya, WiUiams 75, type.
Paullinia Josecuatrii Macbr., sp. nov.
Scandens fruticosa; caulis costato-teretibus petiolis paniculisque
plus minusve dense puberulis; corpus lignum simplex; stipulis ut
videtur caduceis ovatis circa 3 vel 4 mm. longis; petiolis rhachis-
que conspicue marginatis 3-4 mm. latis; foliolis 5 subsessilibus
oblongo-ellipticis basi acutis terminalibus cuneato-attenuatis apice
breviter obtuseque acuminatis plerumque 12-15 cm. longis, 5-6 cm.
latis, argute repando-mucronato-serratis, chartaceis, supra opacis
glabris, nervis 12-15 puberulis approximatis exceptis, subtus nitidulis
puberulis praecipue nervis venisque clathrato-reticulatis subpel-
lucido-lineolatis; paniculis solitariis spiciformis breviter pedunculatis
1.5-2.5 dm. longis, cincinni sessilibus contractis, pedicellis vix 2 mm.
longis; sepalis puberulis interioribus circa 2 mm. longis; petalis subob-
longis eroso-denticulatis 2.5 mm. longis; capsulis longe (circa 5 mm.)
stipitatis subglobosis extus sericeo-puberulis intus pilosis circa 1 cm.
longis, 8 mm. crassis intense sanguineis. — Simulates P. castaneifolia
Radlk., 269, of southeastern Brazil to which Jos^ Cuatrecasas
referred it but that apparently closely related species has acute or
acutely acuminate leaflets with emarginate petioles and rachis and
subsessile fruits. In the margined leaf-rachis it suggests the similar
P. seminuda Radlk., 268, of southern Brazil but that species is more
pubescent and also with acutely acuminate leaflets. It is true that
some species as P. spicata Benth. with normally emarginate petioles
may exceptionally have them margined but here this character is
associated with the obtuse leaflet acumen and therefore is probably
a stable significant species indicator. The collector noted the liana as
5 meters tall, the leaf-nerves above pale brown, flowers white,
fruits brilliant red.
Loreto: Aguaitia, Woytkowski S^^^^S, type.
Paullinia Killipii Macbr., sp. nov.
Fruticosa glabra; ramis floriferis sulcatis 7-15 mm. crassis (corpus
lignosum simplex) subfistulosis; stipulis papyraceis ellipticis acutis
2-3 cm. longis, 1-1.5 cm. latis; petiolis emarginatis, petiolulis 5-15
mm. longis; foliolis 5, oblongo- vel late ellipticis subacutis suba-
equalibus 1.5-2.5 dm. longis, 6-14 cm. latis, reticulato-venosis
chartaceo-membranaceis minutissime pellucido-punctatis; paniculis
in axillis foliorum glomeratis 1-2.5 cm. longis, pedicellis 2-4 mm. Ion-
Flora of Peru 345
gis; floribis glabris circa 3 mm. longis; capsulis ad apicem versus late
trialatis breviter stipitatis circa 2 cm. longis et latis (alis ad 5 mm.
latis). — May be near P. apoda Radlk., 318, of Colombia with three
pairs of leaflets but the stipules and fruits suggest those of P. hispida
while the leaves resemble those of P. nobilis to which I once referred,
carelessly it seems to me now, the type.
San Martin: Juanjul, KIilq j^OO (det. Standley, P. nobilis).
— Junin: Puerto Yessup, KiUip & Smith 263 83, type. — Loreto:
Santa Rosa below Yurimaguas, KiUip & Smith 28876; 287^1.
Paullinia laeta Radlk. Monogr. Paull. 195. 1895; 277.
Differs, ex char., from P. subauricidata: branches densely
lenticellate; leaves about 12 cm. long and broad; petioles and leaf-
rachis broadly winged, the wings obovate-cuneate, about 3 mm.
wide; leaflets elliptic-lanceolate, to 7 (9) cm. long, 3 (5) cm. wide,
1-2 mm. petiolulate, subentire, chartaceous, pale green, lustrous,
obsoletely barbate in nerve axils beneath, minutely pellucid-punc-
tate, containing traces of mucus; stipes of cincinni 2-3 mm. long, the
pedicels as rachis pilosulous; sepals petaloid, glabrous, the ovate
inner 3-4 mm. long; scales bifid, very shortly villous; ovary puberu-
lent. — Kliig 21^1 has apically borne spreading panicles about 1.5
dm. long, puberulent young fruits fusiform, stipes 4-5 mm. long,
leaflets broadly elliptic, 7-9 cm. long, 3.5-5 cm. wide. F.M. Neg.
23650.
Amazonas: Chachapoyas, Mathews, type. — Loreto: Liana in
dense forest, Florida, mouth of the Rio Zubineta, Klug 21^1-
"Imino-o" (Klug).
Paullinia linearis Radlk. Monogr. Paull. 223. 1895; 291.
Incompletely known but apparently distinctive by the 5-6 pairs
of narrowly linear leaflets, the lowest pair temate, the others acute
both ends, entire, sessile, 5-7 cm. long, 4-9 mm. wide, the margined
common petiole 4-5 cm. long, the rachis narrowly winged; stipules
about 2 mm. long; leaves imparipinnate, glabrous both sides except
with immersed glands beneath, epunctate, membranous; wood
simple; flowers and fruit unknown. — Said to be a small shrub.
F.M. Neg. 31038.
Loreto: Yurimaguas, Poeppig, type.
Paullinia Mariae Macbr., sp. nov.
i Liana glaberrima inflorescentia et stipula excepta; corpus lig-
nosum simplex; ramulis trigonis 5 mm. crassis costato-sulcatis; stip-
346 Field Museum of Natural History— Botany, Vol. XIII
ulis oblongo-lanceolatis 1-1.5 cm. longis, 4 mm. latis, adpresse
puberulis; petiolis rhachisque emarginatis leviter costatis, petiolulis
4-5 mm. longis; foliolis 5, late ellipticis obtusis terminalibus ad
18 cm. longis, 8 cm. latis lateralibus 9-12 cm. longis, 5-6 cm. latis
coriaceo-chartaceis nitidulis tenuiter reticulato-venosis obscure re-
moteque ad apicem dentatis vel subintegris paullo vel vix pellucido-
lineolatis; paniculis solitariis circa 1 dm. longis puberulis, cincinnis
sessilibus, bracteis oblongo-lanceolatis 7 mm. longis, pedicellis
5 mm. longis apice articulatis; sepalis 4 mm. longis; petalis obovatis
circa 4.5 mm. longis. — ^Apparently near P. tarapotensis to which
Standley referred it but the leaflets are obtuse and the stipules as
the bracts are conspicuous. The name commemorates the typist of
several numbers of the work, Mrs. Mary Fisher, who noticed with
characteristic alertness that I had used originally an untenable
name for this beautiful vine.
San Martin: Juanjul, Klug 3912, type.
PauUinia martinensis Cuatr. Fieldiana: Bot. 27, No. 2: 82.
1951.
Pubescent even to the outer calyx lobes except the leaves above,
the wood simple, the greenish-gray branchlets subterete, the in-
florescences forming sessile glomerules 1-1.5 cm. broad in the axils;
leaves 5-foliolate-pinnate, 1.5-2.5 dm. long, the rachis as petiole
emarginate, the chartaceous leaflets ovate or oblong-ovate or the
terminal subrhombic, all acute, the lateral rounded to truncate at
base, entire or coarsely 2-3-dentate above, 4.5-8 cm. long, 2.5-5 cm.
wide, the 5-6 lateral nerves prominent the laxly reticulate veins less
so, subglabrous above unless the principal nerves; petiolules 2-6 mm.
long, the terminal to 12 mm. long; stipules linear, 5-10 mm. long;
pedicels 0.1-3 mm. long; outer sepals broadly ovate, 3 mm. long,
inner little longer, puberulent; petals 4, glabrous, elliptic-oblong,
obtuse; scales half as long, the broader appendage deflexed, lightly
barbate, the upper crest bilobed, lower entire; tomentose disk with
2 oblong glands barbellate below; filaments pilose; ovary densely
hispid, the 3 styles glabrous. — Like P. alata (R. & P.) G. Don but
the branches are not winged and contain a simple wood system; it
differs from other species by its narrow linear stipules and pubescent
leaves, their rachis emarginate. (Description as remark after the
author.)
San Martin: Tarapoto, Woytkowski 85160, type.
Flora of Peru 347
PauUinia mazanensis Macbr., sp. nov.
Scandens fruticosa glabra (inflorescentia excepta) ; ramulis valde
costato-sulcatis; stipulis ignotis; petiolis rhachisque emarginatis
striatis, p)etiolulis circa 3 mm. longis; foliolis 5, ellipticis basi plus
minusve acutis apice abrupte tenuiter acuteque acuminatis ple-
rumque 10-13 cm. longis, 4.5-5 cm. latis chartaceis dense reticulato-
venosis obscure pellucido-puncticulatis; inflorescentiis solitariis vel
paniculatis 8-15 cm. longis puberulis, cincinnis nunc 1- nunc 3-7-
floris racemose dispositis; bracteis minutis; pedicellis 1 mm. longis;
sepalis interioribus 4.5 mm. longis; petalis oblongis; ovario glabro.
Without fruit but perhaps as suggested by Standley in herbaria
comparable as well to P. tarapotensis as to any other species and
apparently undescribed.
Loreto: Gamitanacocha, Rio Mazdn, Josi Schunke 2^, type.
Paullinia neglecta Radlk. Monogr. Serj. 71. 1875; 254. Sema-
rillaria nitida R. & P. Fl. Peruv. 4: pi. 339. 1802, not P. nitida HBK.
Essentially glabrous liana, the branches triangular or 5-6-
costate; wood composite with 3 peripheral areas; stipules small,
lanceolate; petioles and rachises of the leaves emarginate, the
former 2-4 cm. long, the petiolules 4-8 mm. long, all sparsely and
laxly puberulent above; temate (rarely 5-foliolate) leaves 12-15 cm.
long, the pinnate ones longer, all nearly as wide, the terminal
leaflets 8-13 cm. long, 4-6 cm. wide, obtusely acuminate as the
nearly as large lateral, all subacute at base, petiolulate, elliptic-
lanceolate, toward the tip more densely repand-dentate, or sub-
entire, subcoriaceous, lustrous, the veins prominent beneath, the
nerve axils barbate, pellucid-punctate, lacking mucus; panicles
sparsely pubescent, solitary, the cincinni sessile, 4-5-flowered, the
pedicels 3 mm. long, articulate at base; sepals subequal; capsule
glabrous, subglobose, abruptly contracted to stipe 3 mm. long,
about 1 cm. long and broad at seed, this 8 mm. long, glabrous, red.
— See remarks under P. tarapotensis; the characters used to separate
these several plants are doubtfully stable. The temately leaved
P. cururu L., 245, scarcely to be expected but possibly, as widely
distributed, has margined leaf rachis. F.M. Neg. 5612.
San Martin: Tarapoto, Williams 6626; 6752. Juan Guerra,
WiUiams 6851, 6911. — Hudnuco: Vitoc and Hudnuco, Ruiz & Pav&n,
type. Ganso Azul, Rio Pachitea, Sandeman 3381 (det. Standley).
— Junln: Chanchamayo, Isem 2381. Colonia Peren^, KiUip &
Smith 25205 (det. Killip).— Ayacucho: Aina, KiUip & Smith 22743
348 Field Museum of Natural History— Botany, Vol. XIII
(det. Killip). — Loreto: Canchahuaya on the Ucayali (Huber IS 89).
Bolivia.
PauUinia nobilis Radlk. Monogr. Paull. 271. 1895; 326.
Indument a minute puberulence, the scandent branches un-
equally 4-5-costate and sulcate, the wood simple, the 5-foliolate-
pinnate leaves with oval-oblong leaflets; stipules subulate, 7-9 mm.
long; petioles and rachises emarginate, about 5 cm. long, sulcate,
puberulent, the petiolules 2-3 mm. long; leaflets about 10 cm. long
and about half as wide, the lower lateral ovate, all shortly acuminate,
acute or rounded at base, subentire or remotely and rather coarsely
repand-dentate, chartaceous, nearly glabrous above except the
mid-nerve, minutely puberulent beneath and punctate-lineolate,
often branched pellucid, lacking mucus; panicles solitary, pulverulent,
the cincinni shortly stiped, to 7-flowered, the pedicels in fruit 4 mm.
long, articulate at base; sepals minutely puberulent, the inner oval
4 mm. long; petals apparently oblong; capsule 3-winged, obovate,
emarginate, puberulent without, pubescent within, the stipe 6-8
mm. long; seed somewhat pubescent, nearly to the middle arillate.
— P. boliviana Radlk., 327, has greenish instead of reddish-brown
branches, serrate oblong-lanceolate leaflets 7 X 2-3 cm.; smaller
(about 2 cm. long) glabrate capsule with 5 mm. long stipe, glabrous
seed; P. caloptera Radlk., 329, allied, of Brazil and Venezuela is
nearly glabrous, the leaf-rachis narrowly margined, leaflets densely
pellucid-punctate, stipules lanceolate, seeds pilose. Since so many
specimens have been placed here in herbaria the author's description
is included but the plant seems to be only a glabrate state of P.
gigantea. F.M. Neg. 5613.
Loreto: In woods, Yurimaguas, Poeppig, type; addenda 93.
Leticia, Ule 619 Jf, fide Radlkofer. Ecuador; Brazil.
PauUinia obovata (R. & P.) Pers. Syn. PI. 1: 443. 1805; 259.
Semarillaria obovata R. & P. Prodr. 54. 1794.
Nearly glabrous liana, only the younger parts slightly puberulent
and the leaves often barbate beneath in the axils of the rather
prominent nerves; branches three-angled and 5-6 costate, 3-5 mm.
in diameter; wood simple; petioles and rachis emarginate; leaves
5-foliolate, the terminal leaflets somewhat obovate, the lateral
oblong-lanceolate, subrounded at base, all acuminate, rather coarsely
dentate, petiolulate (petiolules to 5 mm. long), subchartaceous,
clathrate veined, to 11 cm. long nearly 5 cm. wide, usually smaller,
Flora of Peru 349
the epidennis lacking mucus; panicles solitary, dense, 10-15 or
sometimes to 45 cm. long, the sulcate rachis 3-4 mm. thick, tomentu-
lose with sessile contracted 5-7 flowered cincinni; bracts and bract-
lets minute deltoid; pedicels articulate at base; sepals densely ashy
tomentulose, ovate, p>etals oblong; capsules pyriform, subligneous,
finally glabrescent, about 3 cm. long and half as broad, the stipe
about 1 cm. long; seed nearly completely arillate, about 1 cm. long
and half as broad. — This is the earliest name in a group of closely
allied plants. Illustrated, Ruiz & Pav6n, Fl. Peruv. 4: pi. 338.
F.M. Neg. 23655.
The Indians eat the fleshy white sweet arils that half cover the
seeds (Ruiz & Pav6n).
Hudnuco: Macora, Ruiz & Pav&n, type. — Loreto: Gamitan-
acocha, Rio Mazdn, Jos^ Schunke 118 (det. Standley). Brazil.
"Monte lucuma" (Ruiz & Pavon), "patgo-huayo" (Schunke).
Paullinia olivacea Radlk. Notizbl. Bot. Gart. Berlin 6: 151.
1914; 291.
Scandent nearly glabrous but the terete striate or lightly sulcate
branches lenticellate apically and evanescently puberulent-hirtel-
lous; wood simple; leaves imparipinnate with 2 pairs of oval or
suboblong-lanceolate obtusely acuminate leaflets, the terminal one
cuneate at base, the lateral acute, all sessile, remotely and obtusely
dentate, membranous-chartaceous, reticulate- veined, glabrous, lus-
trous, olive-green above, lightly puberulent especially on the nerves
beneath and microscopically stipitate glandular, densely lineolate
and punctate pellucid, lacking mucus, 6-12 cm. long, 2.5-5 cm. wide,
the winged rachis 1.5-2.5 cm. long, the common petiole 3-5 cm.
long with wings 2-3 mm. wide; panicles solitary, puberulent, the
slender rachis dense, the cincinni sessile, the pedicels and small
bracts canescent as at least the sepals in bud, these in part connate
medially, the inner 2.2 mm. long; petals oblong, glandular both sides,
small; ovary glabrous, globose, long-stiped; fruits of Killip & Smith
specimen to nearly 1.5 cm. thick, the stipe to 7 mm. long. — As
suggested by the author the species is very near P. pterophylla Tr, &
PI. but in the few specimens seen the fewer leaflets and glabrous
fruits seem to be concomitant characters. F.M. Neg. 5614.
Loreto: Santa Rosa below Yurimaguas, Killip & Smith 28966.
— Puno: San Govdn, Lechler 2332a tjrpe, pt. — Rio Acre: Seringal
San Francisco, Ule 9563; 9576.
350 Field Museum of Natural History— Botany, Vol. XIII
PauUinia pachycarpa Benth. in Hook. Jonrn. Bot. Misc. 3:
196. 1851; 296.
Glabrous except for the rusty tomentulose tips of the slightly
three-winged or subterete lenticellate branches, the flowering 3-5
mm. thick, and the solitary or paniculate sessile or pedunculate
panicles; wood simple; petioles and rachis wing-margined; stipular
scars broad, semiamplexicaul; leaves imparipinnate, the lower pair
of leaflets temate; leaflets subelliptic or oblong with a more or less
elongate obtuse acumen, all sessile, remotely serrate-dentate, rarely
subentire, coriaceous-chartaceous, transversely veined, subopaque
both sides, rather densely pellucid-punctate and lineolate 6-20 cm.
long, the lowest lateral smaller, epidermis lacking mucus; panicles
congested, stout, the stiped cincinni more or less contracted, the
elliptic lanceolate bracts 4-6 mm. long; pedicels short; sepals
canescent tomentulose, the high-connate inner 5 mm. long, sub-
coriaceous; petals 7 mm. long, oval-oblong; filament trichomes long,
whitish, abundant; capsules globose, 1.5-3 cm. long with stipe 5-15
mm. long, velvety tomentulose without, pubescent within, the peri-
carp thick; seed nearly enclosed by the aril. — The Peruvian material
was distributed as P. grandifolia Benth. ex Radlk., 294, weakly
distinguished by the scarcely emarginate rachis; both plants may
prove to be a part of P. ingaefolia Richard, 295, with, however,
sessile cincinni. F.M. Neg. 5617.
San Martin: Tarapoto, Williams 67JfS. — Loreto: Timbuchi on
Rio Nanay, Williams 86 U. Manfinfa on Rio Nanay, Williams 1086.
Iquitos, Tessman 3609. Mishuyacu, Klug 825; 1369. To Venezuela.
Paullinia pauUinioides [Spruce] Radlk. Monogr. Serj. 75.
1875; 303.
Glabrous, the terete branches 2-3 mm. in diameter above; wood
simple; petioles emarginate; stipules minute, triangular; leaves ter-
nate; leaflets elliptical, subacute at base, rather long acuminate,
entire or the lateral usually with a callosed tooth at least on the
inner revolute margin near the base, all long-petiolulate, coriaceous,
closely reticulate-veined, subopaque above, slightly lustrous be-
neath, often 8-14 cm. long, the epidermis containing mucus; panicles
solitary or panicled, sparsely puberulent, the rachis scarcely 1 mm.
thick, the sessile or shortly stiped cincinni contracted, pedicels about
3 mm. long, articulate medially or lower; sepals tomentulose, the
inner connate, about 4 mm. long, subpetaloid; petals oblong-oval;
filament trichomes white; capsules ellipsoid-globose, apiculate,
Flora of Peru 851
2.5-3 cm. long, the abundant spines to 1 cm. long, the pericarp
about 3 mm. thick, glabrous within, subglabrate without, the stipes
about 4 mm. long; seed glabrous, arillate to the middle. — This may
be the proper name for P. SpriLcei since that appears not constant
or not recollected; at least most of the following material has leaflets
with one or two calluses on one or more leaflets or the fruits are
closely echinate; Killip & Smith collections det. Killip. The species
name was unpublished by Spruce under Castanella. Illustrated,
Radlkofer, I.e. 221 (fruit).
Loreto: Yurimaguas, Williams 7S7U; S858; Killip & Smith 27570;
27981. Santa Rosa, Killip & Smith 28762. Near mouth of the Rio
Tigre, Rio Marafi6n, Killip & Smith 27526. Northern Brazil.
Paullinia pinna ta L. Sp. PI. 366. 1753; 247.
Nearly glabrous, the scandent trigonous branches 5-6-costate,
usually sparsely pubescent or subtomentose on the angles; wood
typically composite with 1-3 peripheral areas; stipules linear-
subulate, 3-7 mm. long, rarely lanceolate and to 15 mm. long;
petioles and rachises commonly broadly winged, the former 2-6 cm.
long or longer, the latter 2-3 cm. long; leaves 5-foliolate-pinnate, the
terminal leaflet 7-12 cm. long or longer, 3-5 cm. wide, the lateral
little smaller, all ovate, oblong or lanceolate, obtuse or acute both
ends, shortly petiolulate, remotely serrate, subcoriaceous, lustrous,
sometimes sparsely pubescent and barbate in the nerve axils, us-
ually pellucid-punctate and lineolate, lacking mucus; panicles
solitary, usually racemiform, pubescent, the cincinni subsessile, the
2-4 mm. long pedicels articulate near the base; flowers 3-5 mm. long;
sepals obscurely if at all costate, the outer appressed puberulent;
scales with deflexed appendages (the upper), the crest of the lower
aliform; capsule clavate, 2-3 cm. long, 10-14 mm. broad, apiculate,
sometimes more or less crenate; seed 12-15 mm. long. — The similar
and widely distributed P. cururu L., 245, has temate leaves.
The plant contains a toxic alkaloid (as P. cururu) and is used as
a "fish poison."
Hudnuco: Pozuzo, Ruiz & Pav6n (fide Radlk.). Widely distri-
buted in warm America; Africa. "Timbo."
Paullinia pterophylla Tr. & PI. Ann. Sci. Nat. s^r. 4. 18:
354. 1862; 290.
Glabrous or glabrate except the younger parts, the flowering
branches lenticellate, 3-6 nmi. in diameter; wood simple; stipules
352 Field Museum of Natural History— Botany, Vol. XIII
linear-lanceolate, 4-5 mm. long; petioles and rachis broadly wing-
margined; leaves imparipinnate with 3-4 pairs of lanceolate-oblong
acuminate sessile leaflets, the terminal cuneate at base, the lateral
acute, all serrate above the middle, chartaceous, reticulate-veined,
lustrous, obscurely pellucid-punctate and -lineolate, to 11 cm. long,
5 cm. wide, usually 5-9 cm. long, 2-3 cm. wide, drying, fuscescent,
paler beneath, the epidermis lacking mucus; panicles half shorter
than the leaves, solitary, the bracts and bractlets subulate, scarcely
1 mm. long, the slender rachis about 1 mm. thick with sessile con-
tracted cincinni, the pedicels 1.5 to 2 mm. long; inner sepals more
or less connate, 2,5 to 3 mm. long, subpetaloid, appressed puberulent
without, the half as long outer two subcoriaceous; filament trichomes
sparse, whitish; capsules globose, abruptly contracted to stipe 4-6
mm. long, shortly tomentose without, laxly pubescent within, the
subglobose seed nearly completely arillate. — P. olivacea probably
will prove to be a variant but no intermediates seen. Fruit edible.
F.M. Neg. 23654.
Pasco: Oxapampa, Soukup 1823.
Paullinia rhizantha Poepp. & Endl. Nov. Gen. & Sp. 3: 36.
pi. 2J^. 1844; 243.
Similar to P. alata and not clearly distinct but according to the
authors the older branches narrowly wing-angled, the younger,
according to the monographer, deeply 6-sulcate, 6-costate, the
costae hirtellous; leaves glabrous; petioles 7-15 cm. long, rachis
3-4.5 cm. long, wings 1.5-2.5 mm. wide both sides; petiolules 3-5
mm. long; terminal leaflets 14-18 cm. long, 6-10 cm. wide, the
lateral smaller, elliptic-ovate to obovate, angulate- or subrepand-
dentate; panicles 2-2.5 cm. long, the pedicels 6-8 mm. long, medially
articulate. — Flowers according to Spruce, slightly larger. It is
possible that most or all of the specimens from Loreto cited under
P. alata var. loretana should be placed here, as they have the wide
leaflets of this species; this character seems to vary, as also degree
of pubescence.
Loreto: Yurimaguas, Poeppig 2239, type; also addenda 50. — Rio
Acre: Ule 9565. Brazil; Colombia.
Paullinia rugosa Benth. ex Radlk. Monogr. Serj. 75. 1875; 287.
More or less hirtellous liana with 4-5 sulcate branches 4-6 mm.
in diameter, the wood simple; stipules conspicuous, suborbicular,
3-4 mm. long, densely pubescent without, stellate incised; petioles
Flora of Peru 353
and rachis emarginate; leaves 5-foliolate, the broadly elliptic or sub-
orbicular leaflets typically obtuse, the lateral sometimes cordate at
base, all shortly petiolulate, obscurely repand-dentate or entire,
6-18 cm. long, 4-10 cm. wide, coriaceous, drying reddish beneath,
laxly and obscurely clathrate-veined and obscurely pellucid-punc-
tate, the epidermis lacking mucus; panicles much shorter than the
leaves, solitary or panicled, sessile or peduncled, 5-12 cm. long, the
rachis 2-3 mm. thick, the sessile contracted cincinni approximate;
bracts 2-3 mm. long, nearly as wide, the lanceolate subulate bractlets
smaller; pedicels 4-5 mm. long, nearly apically articulate; sepals
ashy-tomentulose, almost entirely free, the inner about 4 mm. long,
coriaceous; petals obovate, about 5 mm. long; filament trichomes
brownish, abundant; capsules shortly stiped, trigonous subglobose,
densely reddish-hirtellous within and without. — The Peruvian
species is only in flower but seems to belong here; it has, however,
a slight difference in leaflets and may become var. peruviana
Macbr., var. nov., foliolis breviter obtuseque acuminatis. F.M.
Neg. 5624.
Loreto: Yurimaguas, KiUip & Smith 27H7. Northern Brazil.
Paullinia serjaniaefolia Tr. & PI. Ann. Sci. Nat. s^r. 4. 18:
356. 1862; 339. Paullinia selenoptera Radlk. Monogr. Paull. 303.
1895; 339.
Scandent shrub, glabrous except the puberulent petioles and
panicles, these solitary, about 5 cm. long; branches triangular,
lightly 3-4-sulcate, wood simple; stipules linear-subulate, 5-6 mm.
long; petioles 0.5-3 cm. long, sulcate above, the rachis 1-2 cm. long,
wings both sides scarcely 1 mm. wide; leaves bitemate or typically
with sometimes two pairs of leaflets, the lowest ternate, the terminal
leaflets about 5 cm. long, 2-2.5 cm. wide, long-attenuate at base,
acuminate, the lateral gradually smaller, acute, all sessile, remotely
serrate, submembranaceous, scarcely nitidulous, glabrous, punctate
and lineolate-p)ellucid, lacking mucus; panicles short, peduncled and
solitary or sessile on older branches; flowers apparently small,
puberulent; capsule broadly 3-winged, the broad wings semicircular,
subsessile, excised at apex, glabrous, within pubescent; seeds ellip-
soid, pilose, one-third arillate. — The forma settUigera Radlk. of
P. selenoptera from Brazil has peduncle-angles, stipules and leaf-
margins setulose. From the Peruvian material and the Colombian
from the region of the type it does not seem that the number of
leaflets is significant since at least one Peruvian collection (Killip
354 Field Museum of Natural History — Botany, Vol. XIII
& Smith 28065) has two pairs of leaflets; and material with sessile
panicles is neariy P. pterocarpa Tr. & PI., 338, but the leaflets are
not entire and the rachis is margined. Illustrated, Mart. Fl. Bras.
13, pt. 3: pi. 90. F.M. Neg. 23657 (P. selenoptera) .
Loreto: Yurimaguas, Poeppig, type; Killip & Smith 28065;
Williams 7879; 53U1; 4.196. Sierra del Pongo, 500 meters, mature
fruit rose-red, Mexia 6277 (det. Standley). Balsapuerto, Killip &
Smith 28666. Mishuyacu, Klug 10; 583; 807. Brazil. "Curuba-
huasca," (Williams).
PauUinia setosa Radlk. Notizbl. Bot. Gart. Berlin 6: 154. 1914;
327.
Sulcate branches, peduncles of the solitary bicirrose panicles,
petioles and petiolules densely and more or less fasciculately long-
setose; stipules ovate-lanceolate, scariose, marginally setulose, 1.5
cm. long, 5-7 mm. wide; petioles emarginate, 8 cm. long, rachis
5 cm. long, narrowly wing-margined (1.5 mm. each side); leaves
5-foHolate-pinnate, the oval or suboblong leaflets 10-16 cm. long,
4.5-8.5 cm. wide, the terminal obovate-subrhombic, attenuate at
base, the upper lateral acute, the lower rounded at base, all acumi-
nate, remotely few-dentate, membranous, finely reticulate, both
sides on nerves, veins and margins, sparsely and finely setulose,
slightly punctate and lineolate-pellucid, lacking mucus; bracts 8 mm.
long, similar to stipules, the pedicels as long; sepals elliptic; petals
6 mm. long, 3 mm. wide, little glandular; ovary (male flowers)
pilose. — Suggests both P. gigantea and P. caloptera (cf. under P.
nobilis), and until fruit is known its position uncertain (Radlkofer).
If a branch of Krukoff 90^1 from Sao Paulo de Olivenga belongs to
the sterile specimen as preserved at Chicago the fruits are about
2 cm. wide and long, the wings nearly as broad at base as apex and
thus resemble somewhat those of P. serjaniaefolia and are congested
at defoliate nodes. The species thus must have either sessile or
peduncled panicles as P. caloptera. P. scaberula R. E. Schultes,
Bot. Mus. Leaflets Harvard. 13: 271. 1949, from the same locality
and therefore probably to be found within Amazonian Peru ap-
parently would be sought here in flower but the fruits are exalate
and the branchlets are "black scaberulent." F.M. Neg. 5625.
Rio Acre: Seringal San Francisco, Ule 9562, type. Brazil?
Paullinia simulans Macbr., sp. nov.
Liana glabra vel glabriuscula; corpus lignosum simplex; ramis
valde costato-sulcatis, trigonis, 5 mm. crassis; stipulis persistentibus
Flora of Peru 355
oblongo-lanceolatis circa 1 cm. longis striatis; petiolis rhachisque
emarginatis; petiolulis 3 mm. longis; foliolis 5, oblongo-ellipticis
breviter obtuseque acuminatis 8-15 cm. longis, 3.5-6 cm. latis ad
apicem remote denticulatis chartaceis paullo nitidulis vix pellucido-
punctatis reticulato-venosis; paniculis sessilibus vel longe peduncu-
latis interdum bicirrosis 4-8 cm. longis densifloris, cincinnis sessilibus
pulverulentis; floribus ut videtur circa 3 mm. longis; pedicellis 1 mm.
longis; bracteis 2.5 mm. longis subulatis; ovario puberulo. — Seem-
ingly allied to P. tarapotensis but the conspicuous stipules per-
sisting; it suggests also in general appearance P. spicata with
deciduous stipules, composite wood, glabrous ovary.
Loreto: Wooded banks of Rio Itaya above Iquitos, Killip &
Smith 29547, type (det. Killip, P. tarapotensis?). Mishuyacu, Kliig
U5. Florida, Rio Pishingo, Klug 2101?
Paullinia sphaerocarpa Rich, ex Juss. Ann. Mus. Nat. hist.
Nat. Paris 4; 348. 1804; 298.
Nearly glabrous liana, the younger lenticellate subterete branches
2-3 mm. in diameter, the wood simple; petioles emarginate, the
rachis wing-margined or in Peru obscurely; stipules minute, broadly
triangular; leaves 5-foliolate, the oval or oblong leaflets acuminate,
the terminal attenuate-cuneate at base, the lateral acutish or
rounded, all remotely, sometimes rather coarsely crenate-dentate,
4-12 cm. long, sessile or petiolulate, membranous-chartaceous,
laxly subclathrate- veined, lustrous both sides but paler beneath, the
epidermis containing mucus; panicles solitary or panicled, puberu-
lent, more or less peduncled, to 3 dm. long, the laxly flowered rachis
about 1 mm. thick with sessile or shortly stiped cincinni, the pedicels
2 mm. long; bracts and bractlets minute; sepals ashy-tomentulose,
the membranous inner about 2.5 mm. long, all free; filaments
hirsute-pilose; capsules subsessile, lanate within, finally glabrate
without, about 2 cm. in diameter. — Illustrated, Radlkofer, I.e. 221.
F.M. Neg. 5626.
Loreto: Iquitos, Williams S6U1. Mishuyacu, Klug 190. To the
Guianas.
Paullinia spicata Benth. in Hook. Joum. Bot. Misc. 3: 193.
1851; 256.
Liana, glabrous or essentially except the solitary spiciform
panicles, the branches subtriangular or 4-6 costate, about 5 mm. in
diameter, the wood composite with three smaller peripheral columns;
356 Field Museum of Natural History— Botany, Vol. XIII
petioles and rachis emarginate, at least in Peru; leaflets 5 or rarely
the uppermost leaves temate; leaflets oblong or oblong-lanceolate,
acuminate or acute, the terminal cuneate below the middle, the
lateral rounded or acutish at base, all shortly petiolulate, repand-
dentate or remotely serrate, subcoriaceous or chartaceous, very lus-
trous both sides, faintly reticulate- veined, barbate beneath in the
nerve axils, conspicuously pellucid-punctate and lineolate, the epider-
mis lacking mucus, the terminal leaflets about 1 dm. long and half as
wide or in Peru somewhat larger, the lateral little smaller; petiolules
2-5 mm. long; panicles subsessile or long-pedunculate, the tomentu-
lose rachis deeply sulcate, the subulate bracts about 5 mm. long, the
many-flowered cincinni sessile; pedicels stout, 2 mm. long, articulate
above the base; sepals tomentose puberulent, the inner about 4 mm.
long, 3 mm. wide; petals narrowly oblong-acutish; filaments pilose;
capsules 2.5 cm. long, 1 cm. broad, strongly spongy thickened within,
glabrous; seed compressed ellipsoid, nearly completely enclosed in
the aril, about 1 cm. long, 3-4 mm. wide. — The Peruvian specimen
matches one by Claussen determined by Radlkofer at Paris (KiHip).
Acumen in Peru obtuse. F.M. Neg. 5998.
Loreto: Between Yurimaguas and Balsapuerto, Killip & Smith
28167 (det. Killip). Brazil to Ecuador and the Guianas.
Paullinia Sprucei Macbr. Candollea 6: 12. 1934; 305. P.
riparia Spruce ex Radlk. Monogr. Serj. 75. 1875, not HBK., 1821.
Glabrous with scandent terete branches, simple wood, ternate
leaves; petioles 4 cm. long, terete but narrowly sulcate, the petio-
lules 2-4 mm. long; leaflets elliptic, subacute at base, shortly acumi-
nate, the acumen obtuse, entire but revolute-margined, coriaceous,
subclathrate venose, nitidulous, containing mucus, 9-12 cm. long;
panicles solitary to 3 dm. long, the cincinni sessile, the fruiting
pedicels 3-4 mm. long; capsule globose, the stipe about 3 mm. long,
sparsely echinate with rigid spines 4-7 mm. long; seeds about 12
mm. long, two-thirds arillate. — Distinct from the other allied species
(P. echinata, etc.) by the spongy instead of compact mesocarp. The
name was Castenella riparia Spruce, in herb.
Loreto: Yurimaguas, at the junction of the Rio Huallaga and
the Rio Maranon, Spruce 3883, type.
Paullinia subauriculata Radlk. Monogr. Paull. 196. 1895; 276.
Scandent shrub, the trigonous-subterete branches lightly yellow-
ish-pilose at tips, the wood simple, the 5-foliate-pinnate leaves about
Flora of Peru 357
16 cm. long and as broad; stipules minute, deltoid, axillary, gemi-
nate; petioles margined or winged above, leaf-rachis winged, both
3-4 cm. long, the petiolules 2-4 mm. long; upper leaflets 9-10 cm.
long, 4 cm. wide, elliptic, shortly and obtusely acuminate, acute at
subsessile base, the lower about 7 cm. long, 3.5 cm. wide, ovate,
subauriculate-bidentate, all rigid-coriaceous, olive-green, subopaque,
epunctate, containing mucus, the lateral nerves arcuate-ascending,
reticulate- veined, especially beneath where sparsely barbate in the
nerve-axils, the nerves above and marginally puberulent; panicles
solitary, rachis pilose, pedicels puberulent, 3 mm. long; cincinni
subsessile; sepals free, sparsely appressed puberulent (male buds);
petals oval, the scales villous; ovary rudimentary, the style puberu-
lent. F.M. Neg. 6000.
Cajamarca: Tambillo, Jelski UlS, type.
Paullinia subrotunda (R. & P.) Pers. Syn. PI. 1: 443. 1805;
260. SamariUaria subrotunda R. & P. Prodr. 54. 1794.
Scandent, the striate or ultimately 4-5 sulcate branches minutely
rusty-tomentose, the wood simple, the leaves 5-foliolate-pinnate;
stipules unknown; petioles 5-7 cm. long, petiolules 3-8 mm. long
and as the rachises emarginate, somewhat puberulent; terminal
leaflet 9-14 cm. long, 5-7 cm. wide, cuneate at base, the somewhat
smaller lateral rounded, the upper obovate (-oblong), the lower
ovate-subrotund, all rounded at base, shortly apiculate-acuminate,
minutely serrate toward the bidenticulate short acumen, chartaceous,
clathrate-veined, more or less barbellate beneath and tomentulose
or finely argenteous (Poeppig) with laterally affixed trichomes,
sparsely pellucid-punctate, lacking mucus and not crystallophorous;
panicles solitary with dense sessile cincinni, the 2-3 mm. long
pedicels articulate; sepals densely puberulent, the inner subrotund,
3-4 mm. long; petals oblong, scales of related P. tarapotensis; fruit
pyriform, subglobose, green, finally glabrous, larger than a walnut
(Poeppig). — This seems to be too near P. faginea, the only apparent
difference being the almost minute acumination with two dentations
and the subrotund lateral leaflets; the latter character is approached
in some material of P. faginea but I have seen no specimen with the
same acumination. Illustrated, Ruiz & Pav6n, Fl. Peruv. 4: pi. SS6.
F.M. Neg. 5629.
As in some other species, for instance, P. ohovata, the Indians eat
the fleshy arils that half cover the seeds (Ruiz & Pav6n).
Hudnuco: Vitoc, Ruiz & Pavdn, type. Cuchero and Pampayacu,
Poeppig 1S27. "Lucumas de monte."
358 Field Museum of Natural History — Botany, Vol. XIII
Paullinia tarapotensis Radlk. Bot. Jahrb. 37: 151. 1905; 258.
Scandent shrub, completely glabrous except the somewhat
tomentulose solitary panicles including the rachis, sepals and ovary;
branches sulcate, 6-costate, the ligneous structure simple; stipules
deciduous, not known; leaves 5-foliolate-pinnate; petioles and leaf-
rachises emarginate, the former 5-8 cm. long, the terminal petiolule
6-8 mm. long, its obovate-subcuneate leaflet 10-12 cm. long, 5 cm.
wide; lower leaflets ovate, the intermediate oval, all narrowly and
obtusely caudate, remotely and obtusely crenate-dentate, petiolu-
late, membranous, glabrous, the arcuate-ascending lateral nerves
prominent beneath, reticulate, epunctate, lacking mucus; lower
panicles subsessile, spiciform, ecirrose, the upper long-peduncled,
bicirrose, the short cincinni sessile, the pedicels scarcely 2 mm. long,
articulate below the middle; petals oblong, 3 mm. long, nearly
equaled by the inner sepals, the upper scale crest obcordate and with
deflexed appendage, the lower aliform. — As remarked by the author,
intermediate to P. elegans Camb., 255, and P. sjyicata Benth., 256,
both known from adjacent lands and to be expected; both have
composite wood (but the related P. pinnata may rarely have simple
wood, suggesting that the character alone may not always be
significant), the former with narrower subacute leaflets barbate in
the axils, larger flowers, the latter with all panicles spiciform,
slightly larger flowers, leaflets barbellate in nerve-axils. It is pos-
sible that P. neglecta should include this in spite of the simple wood
and five leaflets as to types. F.M. Neg. 5630.
San Martin: Juan Guerra, Ule 6613, type. Tarapoto, Williams
6633.
Paullinia tenera Poepp. & Endl. Nov. Gen. & Sp. 3: 37. pi 2^3.
1844; 317.
Lightly sulcate 5-6-angled younger branches as also the sulcate
petioles (2-4 cm. long) setose-hirsute with reddish trichomes;
stipules subulate-lanceolate, ciliate, 5-10 mm. long, 1-2.5 mm.
wide; wood simple; leaves imparipinnate, the lower of the 4 pairs
of lanceolate-sublinear leaflets temate; rachis segments about 2 cm.
long; leaflets subsessile, acute at base, sharply acute at tip or with
1-2 teeth near base, entire, subchartaceous, glabrous, lustrous,
green, epunctate, lacking mucus, 7-11 cm. long, 1-2 cm. wide;
panicles glabrous except the apically setulose small bracts, about
1 cm. long, congested at defoliate nodes, the pedicels about 4 mm.
long, sepals glabrous, the inner 3 mm. long; petals oblong, the scales
Flora of Peru 369
with barbate deflexed appendage, the filaments white-pubescent;
capsule obovate with narrow terminal wings, glabrous, the seed with
white bilobed aril. — This type was scandent, creeping among mosses
and defoliate while Klug noted it as a liana. It seems probable that
this may be the earliest name for several similar plants as P. hi-
dentata, P. fistulosa and P. linearis, all little known. F.M. Neg.
5631.
Hu^nuco: Cuchero, Poeppig 1090, type. — San Martin: Tarapoto,
Ule 5817. Juanjui, Klug U9S (det. Standley).
PauUinia tenuifolia Standley, sp. nov. in herb.
Scandens fruticosa glabriuscula; corpus lignosum simplex; ramis
teretibus circa 4 mm. crassis (cortice canescente); ramulis subher-
baceis leviter costatis vix 2 mm. crassis sparse ciliato-pilosis; stipulis
lineari-subulatis 5-7 mm. longis; petiolis rhachisque tenuis minute
costato-striatis, costis plus minusve dense ciliato-pilosis; foliolis 5,
subsessilibus vel terminalibus basi attenuato-petiolatis ovato-
oblongis vel ovatis anguste caudato-acuminatis remote arguteque
dentatis glabris membranaceis plerumque &-7 cm. longis 2.5-3.5
cm. latis tenuissime laxe reticulatis nervis lateralibus circa 7; pani-
culis 1-3-fasciculatis racemiformis 8-10 mm. longis minute pu-
berulis; cincinnis subsessilibus; pedicellis circa 1 mm. longis; floribus
ut videtur vix 1.5 mm. longis; capsulis glabris obovoideis 12 mm.
longis, 7 mm. crassis. — Since there is no indication by the author as
to the probable relationship of this delicate liana I may, as perhaps
often in other cases, be doing him a disfavor in publishing it for him
as undescribed; it seems to me to be distinctive in its extremely thin
leaves and small flowers, mostly borne on the older and leafless
nodes.
Loreto: Fundo Indiano near Iquitos, climbing to 4 meters in
dense forest, Mexia 6S9U, tyi)e.
Paullinia trilatera Radlk. Monogr. Paull. 254. 1895; 314.
More or less hirsute scandent shrub, the sharply three-sided
branches about 1 cm. in diameter and notably setose at the angles;
wood composite; stipules lanceolate, about 1 cm. long, 3 mm. wide;
rachis submargined, the nearly triangular petioles subhirsute on the
angles; petiolules 2-3 mm. long or the terminal longer; leaflets 5,
the terminal subrhombic, the upper lateral oblong-lanceolate to
15 cm. long, 6 cm. wide, often somewhat smaller, the lower ovate,
all long-acuminate, remotely and obsoletely serrate, membranous.
360 Field Museum of Natural History — Botany, Vol. XIII
glabrous above, somewhat hirsute beneath, not punctate or con-
taining mucus; panicles glomerulate on older branches on the leaf
axils, scarcely 1 cm. long, the 7-8 mm. long pedicels articulate
toward the base; outer sepals very small, the obovate inner about
3 mm. long, glabrous; petals obovate-oblong, more or less unguicu-
late; stamens glabrous; capsules broadly obovate or obcordate,
attenuate to the slender stipe, this about 1 cm. long, conspicuously
3-winged, 2,5-3 cm. long, the wings about 5 mm. wide on each side,
glabrous without, densely pilose within; seed ellipsoid, slightly
pilose, 10-12 mm. long, nearly as broad.
Peru (probably). Rio Putumayo, Peru-Colombia boundary,
Klug 1606. Amazonian Brazil.
Paullinia uchocacha Macbr., sp. nov.
Fruticosa praeter inflorescentibus glaberrima; ramis teretiusculis
leviter striatis; petiolis rhachisque circa 6-striatis baud marginatis;
foliis 5-foliolato-pinnatis; foliolis ellipticis basi acutis apice obtuse
subcaudato-acuminatis, lateralibus breviter, terminalibus longe
petiolatis integris vel interdum ad basin grosse bidentatis, plerumque
6-8 cm. longis et circa 4 cm. latis, chartaceo-coriaceis sparse pellucido-
lineolatis utrinque nitidulis et conspicue reticulato-venosis subtus
glandulis immersis adspersis; inflorescentiis solitariis ad 1 dm. longis,
parce pulverulentis; bracteis subulatis, parvis; floribus circa 3-
fasciculatis, tomentulosis circa 3 mm. longis; sepalis 2 exterioribus
1-1.5 mm. longis, subglabris, ovalibus; pedicellis circa 2 mm. longis;
capsulis subligneo-coriaceis depresse subglobosis 1.5 cm. crassis
breviter abrupteque stipitatis extus et intus glabris exalatis sub-
verrucosis ut videtur rubris. — If the detached fruit belongs to these
flowering specimens this seemingly is a very distinct species, as in
foliage it is quite different from others in the section Enoura to
which the fruit-character apparently best refers it. While most
comparable in Peru to P. curvicuspis the leaves bear some resem-
blance also to those of P. reticulata Radlk., 272, of Amazonian
Brazil but the leaflets in that species are acutely acuminate and
subsessile. The name proposed is a union of the native names as
recorded by the collector, omitting "huasco" (i.e. liana) ; cf. Williams,
Field Mus. Bot. 15: 536. 1936.
Loreto: Maquisapa on the Rfo Nanay, Williams 1211, type;
1209. "Uchohuasco," "cacha."
Paullinia yoco Schultes & Killip, Bot. Mus. Leaflets Harvard
10: 302. 1942.
Flora of Peru 361
Widely spreading liana, the robust scabro-lenticellate stems
attaining 12 cm. in diameter, the younger branches pulverulent and
with approximate stout tendrils, these becoming woody; leaves
usually 5-foliolate to 3.5 (4.5) dm. long and nearly as wide, the
sparsely puberulent rachis sulcate but emarginate; leaflets elliptic
(upper obovate), all shortly and obtusely acuminate, entire, cori-
aceous, chartaceous, glabrous (except puberulent nerves), 1.5 to
2.5 dm. long, 8-11 cm. wide, dark green but lustrous, drying yellow-
ish-brown beneath, clathrate-veined, the 7-9 nerves prominent;
panicles solitary, sparsely pulverulent, 10-15 (25) cm. long, the
rachis 3 mm. thick, axillary, racemiform, sometimes with a tendril;
pedicels 3-8 mm. long, minutely hirtellous as the acuminate bracts;
outer 2 sepals subcoriaceous, tomentulose without and ciliate as the
larger membranous inner three; petals entire, obovate, minutely
pilose within, 2.3 mm. long; filaments lanate; ovary globose, gla-
brous, the stigma deeply trifid; fruits subdrupaceous, obliquely
ovoid, 10-14 mm. long, 4-7 mm. wide, attenuate into stipe, red.
— Appears to be most closely related to P. clathrata Radlk., 302, of
Amazonian Brazil which is much larger and has pubescent sub-
dentate leaflets; there are also several floral differences (authors).
In Peru it suggests greatly P. cupana sens. lat. including the var.
sorbilis.
Klug and Schultes found all the softer tissues of the bark, stems
and wood used to extract the white or brownish sap which in 1927
was analyzed as containing 2.73 per cent caffeine (cf. Rouhier and
Perrot, probably this plant. Bull. Sci. Pharm. 33. 537-539: 1926;
Trav. Lab. Med. 17, pt. 6. 1926; Compt. Rend. 182: 1494. 1926;
Chem. Zentrbl. 1: 138. 1927. According to the authors, the Indians
in adjacent Colombia regularly used this product as a breakfast
beverage.
Loreto: Near the upper Putumayo, fide Schultes, I.e. 321.
Adjacent Colombia and Ecuador. "Yoco," "yoco bianco," "yoco
Colorado," "huarmi yoco."
3. URVILLEA HBK.
Cirrose scandent shrubs similar in habit to the more slender
species of Serjania but the petioled leaves nearly always 3-foliate,
sometimes the terminal leaflet triparted, the lateral deeply lobed.
Fruit samaroid-capsular, membranous or chartaceous, 3-winged,
the cells reticulate, the wings radiately venose.
362 Field Museum of Natural History — Botany, Vol. XIII
Urvillea ulmacea HBK. Nov. Gen. & Sp. 5: 106. 1821; 357.
Scandent, sparsely pubescent, the adult stems deeply trisulcate,
the flowering 1.5-2 mm. thick; stipules small, ovate-lanceolate;
leaves normally temate; petioles to 5 cm. long, terminal leaflet to
9 cm. long and 4,5 cm. wide, attenuate to petiolule, the lateral little
smaller, abruptly contracted to short petiolules, ovate or ovate-
lanceolate, acute or acuminate, mucronulate, unequally or sub-
duplicate serrate-dentate, above, especially on the nerves, puberu-
lent, beneath more or less canescent, rarely glabrate, epunctate or
somewhat pellucid-punctate, containing mucus; panicles exceeding
the leaves, puberulent, the cincinni sessile, the white flowers about
4 mm. broad; sepals glabrous or puberulent; petals obovate-spatu-
late, sparsely and microscopically glandular within; capsule elliptic
or obovate, shortly stiped, narrowly or broadly winged, subexcised
apically, glabrous or puberulent without, glabrous or glandular
within, the cells inflated; seeds 2-3 mm. long, ellipsoid. — In herbaria
by Ruiz and Pavon as "Paullinia dentata."
San Martin: Spruce 3215 (det. Radlkofer). — Hudnuco: Pozuzo,
Jf569; Ruiz & Pavdn; Rivero. — Junin: On sunny brush. La Merced,
5590. — Rio Acre: Seringal Auristella, Ule 956 Jt. Texas and Mexico
to Paraguay and Argentina.
4. GARDIOSPERMUM L.
Suffrutescent or annual with the habit and general character of
Urvillea but the membranous or subchartaceous incompletely 3-
celled fruit inflated. Lower disk glands obsolete.
Flowers 6-8 (-10) mm. long, the upper disk glands corniculate;
terminal leaflets usually wider than 3 cm C. grandiflorum.
Flowers 4-6 mm. long, the disk glands orbicular; terminal leaflets
rarely 2.5 cm. wide.
Terminal leaflets usually distinctly petiolulate; seed hilum typi-
cally small C. Corindum.
Terminal leaflets usually decurrently petiolulate if at all; seed
hilum typically large C. Halicacabum.
Gardiospermum Corindum L. Sp. PI. ed. 2. 526. 1762; 397.
C. Corindum L. forma villosum (Mill). Radlk. I.e. 401 fide Radlk.
Habit of C. grandiflorum and as variable, especially in pubescence;
leaves often more or less subbipinnate or subtritemate; leaflets
Flora of Peru 363
ovate-lanceolate to linear, the terminal at least well-petiolulate,
little larger, 4-7 cm. long, 1-2 cm. wide, all incised dentate to crenate
or subentire; capsule to 4 cm. long and broad, sometimes half as
large; seed of C. grandiflorum but only 2.5-4 mm. in diameter. —
Peruvian forms are lozense (HBK.) Radlk., stems white lanate,
leaves bitemate, leaflets ovate-oblong, dentate, yellowish appressed
setulose above, canescent beneath; moUe (HBK.) Radlk., stems
villous, leaves decompound, the small leaflets appressed pilose, the
large fruits softly hirsute; subsetulosum Radlk., crisply hirtellous to
subglabrous, the leaves to bipinnate, setulose on nerves and veins,
fruit large and pubescent or small and glabrate; etc. — Half-shrub of
drier valleys, 800-3,000 meters (Weberbauer). Standley and
Steyermark, Fieldiana: Bot. 24, no. 6: 240. 1949, reduced this, per-
haps correctly, to C. Halicacabum. Yellow and light brown mature
fruits in pretty festoons among cactus and acacias of dry Pampas
River plain, the most conspicuous plant March 1, 1939 (Stork &
Horton).
Piura: Amotape Hills, (Haughl & Svenson 11525). — Lima:
Amancaes, Ruiz & Pav&n; Wilkes Exped. Matucana, on slide rock,
29j^. — Hudnuco: Near Hudnuco, on shrubbery, 2061; Ruiz & Pav&n;
Haenke; Stork & Horton 9380 (4 meters high on Schinu^, det. Stand-
ley); Soukup 2229.— Junin: San Rafael, 2^8; Sawada P lU.
Cabello, 1335. Amazonas: Chachapoyas, Mathews; Weberbauer
4361; 29 Jt; 1335; 2061; 2^8. Near Tupen, Weberbauer It78It; 5^67;
155. — Arequipa: Prov. Camana, Worth & Morrison 156^2. Mejla
(Gunther & Buchtien 258, det. Bruns). — Apurfmac: Prov. Anda-
huaylas. Stork & Horton, 10661; Rio Pampas & Chincheros, 10788;
10735. Abancay, Soukup 759. — Cuzco: Prov. Urubamba, Vargas
11059. Valley Yucay, Herrera 1368. Widely distributed in warm
regions.
Cardiospermum grandiflorum Swartz, Prodr. 64. 1788; 372.
C. elegans HBK. Nov. Gen. & Sp. 5: 99. pi. JtS9. 1821. C. hispidum
HBK., I.e. 101.
Rather stout, the 5-6 costate-sulcate flowering stems about 3
mm. in diameter, early typically hirsute-tomentose as the petioles
and long rachises, the bitemate leaves softly pubescent beneath, or
in variants all these parts subglabrous or more or less hispid ; leaflets
ovate, the upper lateral suboblong, the terminal subrhombic, this
7-8 cm. long, 3.5 cm. wide, with petiolule 1.5 cm. long, the upper
lateral 5.5 cm. long, 2.5 cm. wide, the lower 2.&-3 cm. long, about
364 Field Museum of Natural History— Botany, Vol. XIII
2 cm. wide, all petiolulate, coarsely or incised dentate, membranous,
yellowish-green; sepals 4, often hispidulous, about as long as the
obovate or elliptic petals (said to be 8-10 mm. long, rarely seen
fully developed); upper disk glands erect, corniform; capsule char-
taceous, obovate or elliptic, trigonous, acute both ends, reticulate,
to 6.5 cm. long, 3.5 cm. broad, shortly stiped, pilose, setulose or
subglabrous; seeds globose, 7.5 mm. in diameter, the whitish hilum,
scarcely exceeding 1.5 mm., suborbicular. — The forma hirsutum
(Willd.) Radlk. is the hirsute-setose stemmed race, (possibly dis-
tinct?) the forma elegans (HBK.) Radlk., (t3T)e locality simply
"Peru") the nearly glabrous variant as to stems, leaves and fruit.
Most of the herbarium material seen lacking flowers is placed here
on the basis of leaf-character; the Williams' collections were dis-
tributed by me as C. Corindum, form.
Cajamarca: Tomependa, Bonpland (type, C. hispidum). —
San Martin: Tarapoto, Ule 66Jt.2; Spruce U217; Williams 5535; 552^.
Vaco Pozo, Woytkowski 351 UA; 35169 (both det. Steyermark).
Morales, Williams 566U- Juan Guerra, Williams 68^2. Juanjui,
Ferreyra ^553 (f. hirsutum). Near Saposoa, Ferreyra Ji.621. — Junin:
Chanchamayo, Isern 2101. La Merced, 5Jlf67 (det. Macbride, C.
Corindum, form). Puerto Bermudez to Cahuapanas, Killip &
Smith 26703 (det. Killip). Puerto Yessup, Killip & Smith 26301
(det. KiUip). — Huanuco: Mission Tocache, Poeppig. — Cuzco: Alto
Urubamba, Diehl & H err era 2Jf90. Without locality, Gay. Warm
America; Africa. "Casha huasca" (Williams), "achocha-china"
(Ecuador, Mexia).
Cardiospermum Halicacabum L. Sp. PI. 366. 1753; 379.
Climbing annual herb, glabrous or minutely puberulent, the
upper leaves biternate, about equaled by the slender bicirrose
panicles; terminal leaflet rhombic-lanceolate, 4-8 cm. long, 1.5-2.5
cm. wide, the lateral smaller, the lower ovate, the upper oblong,
obtuse or subacute both ends, or acuminate, all sessile, closely
incised-dentate or obtusely lobed, sometimes parted, especially on
the nerves and margins, both sides crisply or setulosely pubescent;
flowers 4 mm. long, equaled by the pedicels; sepals 4, glabrous or
sparsely pubescent; disk glands short; capsule subsessile, sub-
globose or turbinate-trigonous, 3-4 cm. long and broad, or smaller,
pubescent; seeds globose, about 5 mm. thick, said to be equaled by
the whitish cordiform bilobed hilum. — The var. microcarpum (HBK.)
Blume is the plant with smaller capsules. Illustrated, Radlkofer,
p. 380 from Pflanzanfam. 3, Abt. 5: 308 (fruit and flower).
Flora of Peru 365
San Martin: Chazuta, Kltug 3997 (det. Standley).— Hudnuco:
Pozuzo, Jt6Jt5. — Junin: Colonia Peren^, KiUip & Smith 25U16 (det.
Killip). — Loreto: Punchano near Iquitos, WiUiams 1815; Killip &
Smith 27SA1 . Iquitos, Williams 1 U6; 795S. Cachiperto, King 8117
(det. Standley). Balsapuerto, Klug 2881 (det. Standley).— Aya-
cucho: Kimpitiriki, Killip & Smith 2285 J^ (det. Killip). — Cuzco: In
patio at Illapani, 700 meters, Biies. In all tropics and subtropics.
"Baillarina" (Williams).
5. THINOUIA Tr. & PI.
Habit of Paullinia or UrviUea but the fruit consisting of 3 seed-
cells with terminal samara-like wings joined at one edge to a central
axis and the nearly regular flowers notably pseudo-umbellate.
CalyTc 5-parted. Petals 5, the scales bifid or 2. Leaves always
trifoliate, the leaflets petiolulate.
Thinouia obliqua (R. & P.) Radlk. Sitzungsb. Akad. Wiss.
Munchen 8: 282. 1878; 425. Paullinia obliqim R. & P. ex Klotzsch
(?), Bot. Zeit. 5: 393. 1847. T. repanda Radlk. in Pflanzenfam. 3,
Abt. 5: 308. 1895; Mart. Fl. Bras. 13, pt. 3: 456. 1897, at least as to
Peru. T. myriantha Tr. & PI. Ann. Sci. Nat. s6r. 4. 18: 369. 1862,
at least to Peru.
Nearly glabrous scandent shrub with terete lightly striate
lenticellate branches, scarcely pulverulent apically; stipules minute;
petioles 4-7 cm. long; terminal leaflet 8.5-10 cm. long, excluding
petiolule 1-nearly 2 cm. long, rounded at base, the little smaller
lateral truncate or subcordate at base, all ovate, acute, mucronu-
late, entire or obsoletely bi-tridentate apically (lateral more or less
inequilateral), chartaceous-coriaceous, subtransversely reticulate-
veined with 4 lateral nerves, these pilose in the axils beneath,
otherwise glabrous both sides, containing mucus and impressed
glandular punctate; panicles cirrose or ecirrose, puberulent, the
pedicels medially articulate; fruit 6 cm. long, the cells rather tumid,
inconspicuously reticulate- veined, 1 cm. long, 8 mm. wide, glabrous,
glandular within, the wings 4 cm. long, 1.5 cm. wide at middle where
broadest, constricted at base. — With flowers unknown in type,
generic position doubtful (Radlkofer) but from the materials now
accumulated it seems doubtful if there is any specific difference in
the relative lengths of petals and petal scales. However this may be,
the small petals may barely exceed the calyx but may be exceeded
by the scales. F.M. Neg. 5639.
366 Field Museum of Natural History— Botany, Vol. XIII
Hudnuco: Pozuzo, in woods, Ruiz & Pav&n, type. — Junin: Santa
Rosa, 650 meters, Killip & Smith 26139 (det. Killip, T. repanda).
—San Martin: Juanjui, Klug ^293. Chazuta, Klug 4058; 4102.
— Loreto: Rio Mazdn, Quebrada Andrade, Jose Schunke 183 (det.
Standley, T. repanda). On the Rio Ucayali, Tessmann 3314; 3500
(det. Dahlem, T. myriantha vel aff. T. obliqua). — Ayacucho: Near
Kimpitiriki, Killip & Smith 22969. — Cuzco: Echarate, 900 meters.
Stork, Horton & Vargas 10496 (det. Standley, T. repanda).
6. ALLOSANTHUS Radlk.
Scandent shrub simulating closely AUophylus but the regular
flowers (only the male described) with 5-lobed calyx, the 5-deltoid
lobes valvate and scarcely exceeding the patellate disk, this with
subconvex center and elevated free margin. Petals 5 with 2 inner
densely villous scales. Stamens 8, densely villous-tomentose below,
exserted, the globose anthers papillose, excised at base, the pollen
grains globose. — Fruit unknown but genus placed by the author in
the vicinity of Cupania with dehiscing capsules.
Allosanthus trifoliolatus Radlk. Pflanzenreich IV. 165: 1157.
1933.
Glabrous, except the ashy-puberulent or pulverulent inflores-
cences, these solitary to several in the axils of the sparsely leafy
4-5 mm. thick terete branchlets; leaves trifoliolate, the petioles
4-10 cm. long, the distinct lateral petiolules about half as long as
the terminal, the entire oblong-elliptic leaflets subequal, mostly
about 1 dm. long, 4-5 (8) cm. wide, broadly obtuse or rounded at
base, the lateral unequal, all shortly and obtusely acuminate, firm-
chartaceous or subcoriaceous with 6-8 lateral nerves anastomozing
before the margin, the laxly reticulate veins subprominent both sides,
epidermis lacking mucus; inflorescences 6-10 cm. long, the stiped
cincinni 5-9-flowered nearly to base, the pedicels 2.5 mm. long;
bracts and bractlets deltoid; flowers 3 mm. wide. — Type climbing to
25 meters, the trunk diameter 4.5 cm.
San Martin: Juanjui, Klug 4176 (det. Sand with, with query;
distributed as Connarus Patrisii). — Loreto: Between Iquitos and
Pongo de Manseriche (Tessmann ^^^^; 4462, type collections).
7. ALLOPHYLUS L.
Erect shrubs or small trees with normally trifoliate leaves, the
lateral leaflets more or less oblique, all often pellucid-punctate or
Flora of Peru 367
-lineolate. Inflorescence racemiform or loosely paniculate, axillary,
the small or minute flowers globose or nearly, and usually closed.
Sepals 4, opposite in pairs, the outer much smaller. Petals each
with a 2-lobed or bifid scale and 2-4 disk glands opposite them,
the disk unilateral. Stamens usually 8; anthers short-ellipsoid, the
pollen grains trigonous-placentiform. Ovary deeply bilobed, the
lobes joined by the bilobed style. Fruit an obovoid-globular in-
dehiscent coccus, the erect seed with a very short but fleshy aril.
— The monographer has relied primarily on the number of inflo-
rescence branches — or their lack — and the size of the flowers in
order to group the species. The first character, particularly, is not
entirely reliable or discernible, especially in undeveloped specimens.
Leaves 1-foliolate.
Younger branchlets reddish-hirtellous A. amazoniciLS.
Younger branchlets soon glabrous A. loretensis.
Leaves 3-foliolate.
Panicles (so far as known) consistently simple; leaves soon gla-
brous or essentially, except A. semidentatiLs, rarely A. edulis,
always pellucid-punctate.
Leaflets obovate, nearly or quite half as wide as long, pubescent
beneath A. semiderUatus.
Leaflets about elliptic-lanceolate, often a third as wide as long.
Leaves membranous-chartaceous, hypoderma none, greenish
in herbaria.
Leaflets (medial) canaliculately cuneate at base. .A. edulis.
Leaflets all petiolulate or at least medial A. punctattis.
Leaves coriaceous in type, reddish in herbaria, and upper
surface with hypodermal tissue A. peruvianus.
Panicles usually with one or more branches or rarely simple and
the leaves then (in Peru) densely pubescent beneath.
Leaves soon glabrous or essentially; inflorescence branches often
elongate.
Inflorescence branches often, at least some, with one or more
branchlets; leaflets usually broadly obovate-elliptic,
subabruptly acuminate, membranous. . . .A. panicxdatus.
Inflorescence branches simple; leaflets often about lanceolate
or oval, firm, usually subcoriaceous unless in A. leio-
phloeus.
368 Field Museum of Natural History— Botany, Vol. XIII
Rachis branches slender even in fruit, rarely 1 mm. thick,
soon glabrate A. leiophloeus, A. floribundus.
Rachis branches stout even in flower, tardily glabrate.
Leaflets lanceolate, 4-6 cm. wide A. scrohiculatus.
Leaflets oval, 2-3 cm. wide A. amentaceus.
Leaves obviously pubescent especially beneath, at least the
nerves with conspicuous trichomes.
Leaflets somewhat elliptic-obovate, sharply denticulate, 4-9
cm. wide; inflorescence branches elongate. .A. divaricatus.
Leaflets nearly elliptic, crenate or dentate, 2-5 cm. wide;
inflorescence branches usually short or none.
Leaves coriaceous; inflorescences usually short-branched.
A. coriaceus.
Leaves membranous; inflorescences rarely branched.
A. densiflorus.
AUophylus atnazonicus (Mart.) Radlk. Pflanzenfam. 3, Abt.
5: 312. 1895; 480. Schmidelia amazonica Mart, in Flora 22: Beibl.
1:6. 1839.
Shrub or small tree, the branchlets densely hirtellous, the older
glabrate brown branches pale lenticellate-punctate; leaves 1-foliate,
sometimes with stipelliform processes at juncture of the 1-2 cm. long
petiole and the obovate, elliptic-oblong or sublanceolate blade, this
about 1 to nearly 2 dm. long, 4-7 cm. wide, acute or subobtuse
at the sessile base, shortly and obtusely acuminate, crenate-serrate,
the lateral nerves about 10-15, the veins subclathrate, chartaceous,
glabrous, drying brownish-green, lustrous above, containing mucus,
minutely and obsoletely pellucid punctate; panicles half as long or
as long as leaves, sometimes exceeding; cincinni sessile or subsessile,
3-11-flowered, the ashy-puberulent typically white flowers 2 mm.
wide, the petal claw and scales barbate; fruit obovoid, subglabrous,
7 mm. long, 5 mm. thick. — The Peruvian plant, according to the
monographer, is var. angustifolia Benth., younger branches, more
slender panicles and petioles subglabrous instead of rufescent-
hirtellous; panicles subequaling the leaves, the cincinni sessile, but
this specimen, not seen, is probably A. loretensis Standley. A 7-
meter tree with yellow fruits (Spruce). F.M. Neg. 5651.
Loreto: Yurimaguas, (Spruce 3907 var., fide Radlk.). Brazil.
"Parana" (Brazil).
Flora of Peru 869
AUophylus amentaceus Radlk. Sitzungsb. Akad. Wiss. Mtiii-
chen 38: 215. 1909; 508.
Younger branches as the petioles (5-15 mm. long) and short
panicles fulvous tomentulose-puberulent, the older terete, rugulose,
glabra te, densely lenticellate; middle leaflet 3.5-7 cm. long with the
1-2 mm. long petiolule, all oval, about 2-3 cm. wide, shortly acumi-
nate, entire except for 1 or 2 teeth near the tips, fleshy rigid-coria-
ceous, glabrous above except for the rusty pubescent midnerve, also
beneath except fulvous barbate in the axils of the 6-10 lateral
nerves and a few setae on the prominently reticulate veins, brownish,
subopaque, with brown hypoderma above; panicles soon glabrate,
mostly geminate in the axils, 1-5 cm. long, to 3 cm. broad, the
younger amentaceous, densely multiflowered ; buds small, sub-
glabrous; otherwise unknown. — Like A. crassinervis Radlk. of the
West Indies it emits, moistened, the odor of Piperaceae. As in the
case of the Pav6n collection of A. Cominia (L.) Sw., 508, the type
may have come from Guayaquil. The latter is more tomentose,
the panicles rarely geminate, branched medially, often exceeding
the leaves. F.M. Neg. 23617.
Peru(?): Pav6n, tjije. Herb. Boiss.
AUophylus coriaceus Radlk. Sitzungsb. Akad. Wiss. MUnchen
38: 211. 1909; 488.
Branches 3-5 mm. thick, the terete younger as the short nearly
subsessile panicles fulvous with a dense spreading tomentum, the
older as the petioles (1.5-5 cm. long) and petiolules (3-6 mm. long)
sordidly so, finally glabrescent, obsoletely pale lenticellate; leaflets 3,
the middle one rhombic-lanceolate, cuneate at base, 4.5-13 cm. long,
3-5 cm. wide, the lateral a third smaller, obliquely ovate-lanceolate,
all definitely acuminate, petiolulate, above the lower third uncinate
or subcrenate-serrate, 8-14 lateral nerves, beneath densely and
finely, above laxly reticulate- veined, coriaceous, the nerves above
early, the veins also beneath ashy-tomentose, obsoletely punctate,
lineolate-pellucid; panicles 1-3 cm. long; pedicels very short; flowers
about 2 mm. wide, the sepals pilose-ciliate, sparsely appressed
puberulent at base; petal claw and scales ciliate, the blade of petal
sparsely; disk hirtellous; stamens except at tip densely pilose, the
rudimentary ovary densely setulose. F.M. Neg. 23620.
Cajamarca: Magdalena to Contumaza, 2,500 meters, Weber-
bauer 722S (det. in herb., A. stenodiclyus). Without data, Ruiz &
Pav6n, Herb. Boiss., type.
370 Field Museum of Natural History— Botany, Vol. XIII
Allophylus densiflorus Radlk. Sitzungsb. Akad. Wiss. Miin-
chen 38: 211. 1909; 489.
Similar to the allied A. coriaceus; younger branches as petioles
(1-3.5 cm. long) and peduncled panicles (1-5 cm. long) spreading
or subappressed puberulent, lenticellate-lineolate; middle leaflet
5-8 cm. long, 2-3 cm. wide, on petiolule 1-3 mm. long, all elliptic-
lanceolate, the upper one-third or one-half coarsely serrate, the
lateral little smaller, membranous, drying green or above fuscescent,
glabrous except for the sparsely puberulent nerves and veins and
beneath barbate in the axils; sepal cilia glandular; petal blade sub-
glabrous; disk glabrous. — Otherwise like the related species, and less
pubescent, perhaps a variant; the panicles are sometimes a little
branched; both species, with the Columbian A. stenodictyus Radlk.,
505, are very near A. mollis (HBK.) Radlk., 511, with somewhat
broader leaves in Colombia, the type locality. The extent of
branching accepted by the monographer to distinguish these two
similar shrubs certainly is not constant in development. Further-
more, these forms appear to be doubtfully distinct specifically from
the more northern widely ranging A. Cominia (L.) Swartz, 508, with
perhaps more serrulate leaflets. Radlkofer referred a specimen
from Guayaquil by Pavon (or Tafalla) to this species, apparently
the basis for his indication in his key, 462, of its occurrence in Peru;
cf. A. incanus Radlk., perhaps another variant, mentioned under
A. divaricatus Radlk. F.M. Neg. 23619.
Amazonas: Chachapoyas, Mathews 3208, type. — Piura: Frias,
Prov. Ayavaca, 1,400 meters, Weberhauer 64^1 7.
Allophylus divaricatus Radlk. in Mart. Fl. Bras. 13, pt. 3:
493. 1900; 512.
Subterete branches, terete petioles (1.5-7 cm. long) and basally
divaricately 3-6-branched panicles (7-14 cm. long) densely and
crisply pale-fulvous tomentose-strigillose, finally glabrate; middle
leaflet broadly elliptic-lanceolate and slightly obovate with 2-8 mm.
long petiolule, 8-19 cm. long, 4-9 cm. wide, the lateral a third
smaller, subsessile, oblique, all nearly to the base acutely, closely
and rather coarsely somewhat double serrate, herbaceous, drying
brownish-green above, lustrous and sparsely puberulent unless on
the nerves, pale green and densely ashy-hirtellous beneath, es-
pecially on the 12-20 approximate oblique nerves and the densely
clathrate veins, obsoletely and very minutely pellucid-punctate;
flowers white, many, crowded, only about 1 mm. broad; sepals
Flora of Peru 371
pilose and glandular-ciliate, the petal claw and scales barbate; disk
pubescent; style subglabrous. — The related A. incanus Radlk., 513,
of Ecuador has spreading panicle branches, merely subcrenate
leaves, the younger ashy subsericeous, all petiolulate, the middle
petiolule to 1 cm. long while the rather similar A. cinnamomeus
Radlk., 513, of Bolivia, with reddish-brown tomentum, subsessile
unequally and sharply serrulate leaflets, has yellowish flowers
notably about 3 mm. broad. Shrub or 8-meter tree (Spruce) with
straight trunk to 3 dm. in diameter; common in thickets on the
plain of Tarapoto (Williams). F.M. Neg. 5653.
San Martin: Near Rio Mayo, SpriLce j^89. Near Tarapoto,
Ule 6639; Williams 5969; 60SS; 6187.— i\mm: La Merced, Chan-
chamayo Valley, 1,000 meters, Weherhauer 1925; 282. — Loreto:
Rio Putumayo, Klug 2S6j^ (det. Standley). Brazil; Ecuador.
"Yurac-tortilla-caspi" (Williams).
Allophylus edulis (Camb.) Radlk. in Warming, Symb. pt. 37:
995. 1890 (Vid. Medd. Kjoeb. 1890; 244) 493. Schmidelia edulis
Camb. in St. Hil. Fl. Bras. 1: 381. 1827.
Usually nearly glabrous shrub or small tree, the rather slender
branches strict, the younger somewhat hirtellous ones densely
foliate, short, angulate, with many pallid lineolate lenticels, the
older ashy or reddish-brown, glabrate, the lenticels verruculose;
petioles mostly dilated apically, 3-5 cm. long, the middle leaflet
often 8-10 cm. long, 2.5-3 cm. wide, sometimes larger or smaller,
basally long-cuneate but subsessile, medially serrate, all conspicu-
ously acuminate, the lateral sessile, little smaller, ' more or less
oblique, lateral nerves 8-14, membranous-chartaceous, glabrous or
the younger especially on the nerves pubescent, often barbate in the
axils, lustrous and lucidulous above, at least early pellucid-punctate,
hypoderma none; panicles crowded at the base of the younger
branches, terminally and laterally, usually long-ped uncled, laxly
flowered, glabrous or minutely puberulent, the cincinni sessile or
stiped; flowers about 2 mm. broad, yellowish-white, the sepals
glabrous except the glandular-ciliolate margins, the petals on mar-
gins little, on scales densely barbate; disk puberulent; ovary glabrous
or rarely sparsely puberulent; fruits red, drying black, to 8 mm. long,
obovoid, glabrous, edible. — Illustrated, St. Hil., Juss., et Camb. PI.
Us. pi. 67 (1828, not 1824, fide Radlk. which makes the Camb.
citation I.e. above correct).
Rio Acre: Seringal San Francisco, Ule 9569. Brazil to Paraguay,
Bolivia and Guiana.
372 Field Museum of Natural History— Botany, Vol. XIII
Allophylus leiophloeus Radlk. Sitzungsb. Akad. Wiss. Miiii-
chen38:214. 1909; 503.
Slender tree, the strict terete lenticellate branches early pu-
berulent, finally glabrescent as also the petioles (3-6.5 cm. long),
equally long usually bi-branched panicles and leaves beneath; middle
leaflet obovate-lanceolate, cuneate at base to petiolule 2-3 mm.
long, the lateral a third smaller, elliptic-lanceolate, subsessile,
oblique at acutish base, all acutely acuminate, remotely and coarsely
serrulate above the middle, membranous, glabrous above except for
the 7-12 lateral nerves, densely and finely reticulate-veined, niti-
dulous both sides, obsoletely pellucid-punctate; pedicels nearly 2
mm. long; flowers about 2 mm. broad, white, the sepals nearly
glabrous except the pilose and glandular-ciliolate margins, the petal
scales and claw densely barbate, the petal blade sparsely ciliolate,
the scale lobes ligulate; disk subglabrous; fruit not known. — Ap-
parently a state of A. floribundus and all of the collections from San
Martin cited under that name probably belong here if A. leiophloeus
is valid. Allied by the author to A. petiolulatiis Radlk., 501, widely
distributed in Brazil and known from Bolivia, with leaflets typically
long-petiolulate, leaflets barbate in nerve axils beneath; there are
other minor differences but the Bolivian plants seem to be inter-
mediate and probably there is only a single species. These named
forms center around A. glahratus (HBK.) Radlk. of Colombia. Five
to 7 meters tall (Spruce).
San Martin: Tarapoto, Spruce W2, type. Juan Guerra, UU
6615 (det. Ra,dlk.).
Allophylus floribundus (Poepp. & Endl.) Radlk. Pflanzenfam.
3, Abt. 5: 312. 1895; 502. Schmidelia floribunda Poepp. & Endl.
Nov. Gen. & Sp. 3: 38. 1844.
Much branched, densely leafy tree, the older branches glabrate,
cinerescent, the younger as the panicles sparsely puberulent, the
latter crowded apically, and usually with 1-3 basal branches about
as long as the leaves, many-flowered, the pedicels glabrous and only
about 1.5 mm. long; petioles complanate above, 1-2 cm. long;
middle leaflet 8.5-15 cm. long, 2-4 (6) cm. wide, lanceolate as the
little smaller or oblique lateral ones, all shortly and mucronulately
subfalcate acuminate, shortly petiolulate, remotely and subrepandly
uncinulate-serrulate on the upper half, rigid chartaceous, somewhat
lustrous above, subopaque below, obsoletely punctate-lineolate
pellucid, glabrous except barbate in the 6-12 distant lateral nerve
Flora of Peru 373
axils beneath, where especially densely reticulate- veined ; flowers
about 1.5 mm. broad; sepals glabrous except the ciliolate margins;
petal claws and scales densely barbate; disk nearly glabrous; young
fruits sparsely puberulent, obovoid-globose. — The San Martin
material probably is referable to A. leiophloeus Radlk. if that is
separable. Loreto determinations by Standley. Small tree 4-11
meters tall with open crown, slender cylindrical trunk, the grayish-
brown bark with many small fissures (Williams). F.M. Neg. 31032.
Hudnuco: In woods at Cuchero, Poeppig 1^0, type. — Junin:
Chanchamayo Valley, 700 meters, Weberbauer 18S1; 19U7; 282.
La Merced, 52^9; Killip & Smith 2S50S; 2Jt062. San Ram6n,
KiUip & Smith 2U890. Colonia Peren^, KiUip & Smith 250W;
250U1- San Martin: Tarapoto, Ule 6592 (det. Radlkofer); Williams
5Jt5S; 6020; 6596; 6791 . Juanjui, Klug 3823 (det. Standley). Bella-
vista, Ferreyra U73U. San Roque, Williams 7001; 7007; 7239.
Chazuta, Klug 3879 (det. Standley). Lamas, Williams 6U05; 6U2.
— Loreto: Yurimaguas, WiUiam^ ^31. Pumayacu, Kliig 3155.
Balsapuerto, KItLg 3033; 3043. Florida, Rfo Putumayo, X% 2055;
2302. "Shitari-caspi" (Williams).
Allophylus loretensis Standley, sp. nov. in herb. A. avnazonicus
(Mart.) Radlk., var. angustijolius Benth. ex Radlk. Pflanzenreich
IV. 165: 481. 1932?
A. amuzonico arete affinis; insignis ramis junioribus thyrsis et
petiolis subglabris; petalis ad unguem et ad squamam leviter bar-
batis; cincinnis sessilibus. — Type Klug 3103. These differences,
slight indeed, seem to be constant; furthermore, the inflorescences
are very slender, nearly glabrous, sometimes almost as long as the
leaves, these usually drying gray-green, and the open flowers are
scarcely 1.5 mm. wide; fruits (Killip & Smith 26801) rather coarsely
venose, little narrowed to base, about 8 mm. long. It seems pro-
bable that Spruce 3907 included by Radlkofer in the variety of
Bentham (compare A. amnzonicus) belongs here if Standley 's 4-6
meter tree or shrub proves with more collections to be specifically
distinct. As usual the author has not indicated his own idea of
relationship and it may be questionable courtesy to adopt his
unpublished name (as for that matter often in other cases) but here
too the material has been generally distributed so that convenience
calls for the use of this name.
San Martin: Chazuta, Rio Huallaga, Klug 4001 (det. Standley).
— Junfn: Cahuapanas on Rio Pichis, Killip & Smith 26801 (Killip,
A. panicuiaius). — Loreto: Balsapuerto, Klu{i 3103, type. Brazil?
374 Field Museum of Natural History— Botany, Vol. XIII
AUophylus paniculatus (Poepp. & Endl.) Radlk. Pflanzenfam.
3, Abt. 5: 312. 1895; 514. Schmidelia paniculata Poepp. & Endl.
Nov. Gen. & Sp. 3: 38. 1844.
A rather tall subglabrous tree often 10-12 meters tall, the new
branches as the panicles sparsely ashy-puberulent, the former terete
in age, glabrous, the latter paniculate with many mostly branched
branches; petioles to 1.5 dm. long, semiterete, broadly complanate
above, dilated at tip, the middle petiolule to 2 cm. long, its obovate-
elliptic leaflet to 2.5 dm. long, 1 dm. wide; lateral leaflets a third
smaller, scarcely oblique, all well-acuminate, acutely petiolulate,
the upper half remotely and subrepandly serrulate, at maturity
chartaceous-membranous, drying brown, nitidulous, densely and
obsoletely pellucid-punctate, glabrous, early puberulent on the 10
distant arcuate-ascending nerves, the veins loosely clathrate and
densely reticulate; panicles subequaling the petioles, laxly many-
flowered, the cincinni sessile, the pedicels 1 mm. long; flowers about
2 mm. broad, usually white, the sepals glabrous except for pilose-
ciliate-glandular margins, the petal claw, scales and margins ciliate;
disk subglabrous; style glabrous. Flowers white or cream-colored
(Mexia). F.M. Neg. 31033.
Hudnuco: Cuchero and Pampayacu, Poeppig, type. Rocky
forested slope above Cayumba, Mexia 8321 (det. Standley, A.
peruvianus). Rio Cayumba, Mexia 8263 (det. Standley, A. peru-
vianus). — San Martin: Pongo de Cainarachi, Kliig 2731.
AUophylus peruvianus Radlk. in Mart. Fl. Bras. 13, pt. 3:
488. 1900; 497.
Like A. punctattis but (as to type) the stouter branches more
coarsely lenticellate, petioles 2-10 cm. long, the leaflets lanceolate,
cuspidate, coarsely serrate the upper two-thirds, in type rigid
coriaceous, glabrous, the younger nitidulous beneath, drying
reddish-brown, hypodermatic above; sepals (on fruits) glabrate,
ciliate. — Species of doubtful validity; Radlkofer himself. I.e. 497,
observed that there is in Herb. Boiss. a specimen by Pavon inter-
mediate between these named forms. The Madrid type is distinctly
punctate against a strong light. F.M. Neg. 5659.
Hudnuco: Chinchao, Sawada 89. Without locality, Ruiz &
Pavdn, type.
AUophylus punctatus (Poepp. & Endl.) Radlk. Pflanzenfam.
3, Abt. 5: 312. 1895; 496. Schmidelia punctata Poepp. & Endl. Nov.
Gen. & Sp. 3: 38. pi. 2U- 1844.
Flora of Peru 375
Small tree, the younger branches appressed yellowish puberulent,
soon glabrate, pale lenticellate, the older cinerescent; petioles sub-
terete, dilated at tip, 2.5-6.5 cm. long, the middle j)etiolule 3-8 mm.
long, the leaflets 8-20 cm. long or longer, 2.5-8 cm. wide, elliptic-
oblong or broadly lanceolate, the lateral similar or half as large,
somewhat oblique and often subfalcately incurved, all acuminate,
the upper half obsoletely or coarsely serrate (or subentire),
membranous-chartaceous, subopaque both sides, green or drying
brownish, more or less conspicuously pellucid-punctate and lineo-
late; hypoderma none, epidermis containing merely traces of
mucus, glabrous unless for barbate nerve axils beneath the lateral
nerves 8-15, venation subclathrate; panicles solitary or many,
mostly exceeding the petioles, laxly flowered, canescent or glabrous,
the cincinni subsessile, many-flowered, the pedicels 1-2 mm. long;
flowers whitish, 2 mm. broad, the sepals pilose and glandular-ciliate,
the scales and petals densely ciliate-barbate; disk glabrous (type)
or pubescent; stamens glabrous or pilose below; fruits to 9 mm. long,
8 mm. thick, orange or red. — This seems to be very near A. ediUis
(Camb.) Radlk.
Hudnuco: Tocache, Poeppig 1858, type; 2395. Monz6n, Weber-
bauer 8^85; 284.— San Martin: Tarapoto, Spruce U18; U60;
WiUiams 5556; 5716; 5781; 586U; 6108; 62U9; 6802 (det. Standley,
A. scrobiculatiLs) . Bellavista to Moyobamba, Ferreyra 1^825. —
Ayacucho: Rio Apurimac Valley near Kimpitiriki, Killip & Smith
22988.— horetxi: Yurimaguas, Killip & Smith 29086 (distrib. as A.
scrobiculatiLs) ; Williams W7. Florida, Rio Putumayo, Kliig 2081
(det. Standley). — Rio Acre: Near mouth of Rio Macauhdn, Krukoff
5286; 5If27. Bolivia; Ecuador; Brazil.
Allophylus scrobiculatus (Poepp. & Endl.) Radlk. Pflanzen-
fam. 3, Abt. 5: 312. 1895; 504. Schmidelia scrobiculata Poepp. &
Endl. Nov. Gen. & Sp. 3: 37. pi. 2U. 1844.
Tree typically with lenticellate branches, ashy-puberulent
especially at the nodes and ashy-puberulent mostly nodding panicles
usually with 1-4 branches and longer than the petioles, these sub-
terete, 3-5.5 cm. long; middle leaflet with exarate petiolule 3-5.5
mm. long, lanceolate, about 1-1.5 dm. long, 4-6.5 cm. wide, the
lateral little smaller, little oblique, all obtusish acuminate, acutely
and shortly petiolulate, remotely and repandly uncinate-denticulate
above the lower third, firm-chartaceous, drjring brown above, paler
brownish-green and subopaque beneath, minutely pellucid-punctu-
376 Field Museum of Natural History — Botany, Vol. XIII
late, glabrous unless barbate in the axils of the 8-12 arcuate-ascend-
ing nerves beneath where subclathrate and finely reticulate- veined;
pedicels short; sepals minutely pilose and glandular-ciliate (or gla-
brate); petal claw densely (as scales), blade sparsely, ciliate; disk
glabrous; fruits about 5 cm. long, 4 mm. thick, slightly ashy-puberu-
lent as the style. — Some of the material placed here has nearly
smooth branchlets, subglabrous sepals, while Woytkowski 507 is
further marked by almost hirsute branchlet tips. Type from a tall
tree with grayish bark and hard white wood. F.M. Neg. 5660.
Hudnuco: Fundo San Ricardo, Divisoria, 1,700 meters, Woytkow-
ski 507? — San Martin: Tarapoto, Williams 6712. — Loreto: In woods
at Yurimaguas, Poeppig 2Jf.51, type; 228^. Santa Rosa below Yuri-
maguas, Killip & Smith 2895^ (det. Standley) . Lower Rio Nanay,
Williams 5j^^; 567. Iquitos, Williams 7903 (det. Standley). Pebas,
Williams 1763. Florida, Rio Putumayo, Klug 2171. "Jimuiequir-
ey" (Huitoto, Klug), "shimbillo," "parinari," "quinilla colorada"
(all Williams).
Allophylus semidentatus (Miq.) Radlk. in Pflanzenfam. 3,
Abt. 5: 312. 1895; 486. Schmidelia semidentata Miq. Linnaea 22:
798. 1849.
Slender-stemmed shrub, the 2-3 mm. thick branches, panicles
and petioles, these 2.5-8 cm. long, early spreading pubescent with
soft trichomes, finally glabrate or sparsely persisting especially
beneath on the large trifoliate leaves; intermediate leaflets obovate,
8-20 cm. long, nearly half as wide on petiolules 1-5 mm. long, the
lateral little shorter, inequilaterally obovate-lanceolate, all notably
acuminate and above the middle coarsely and unequally serrate,
densely pellucid-punctate, containing mucus; panicles slender,
simple, 1.5-10 cm. long, the peduncles 0.5-3.5 cm. long; flowers
2 mm. wide, the puberulent sepals glabrous within equaled by the
white sparsely ciliolate petals with densely barbate scales, the disk
as ovary hirsute, the style subglabrous, the stamens pilosulous
below. — The typical form (known to me only from photograph) is to
be expected in the adjacent Department of Madre de Dios. Stand-
ley, without indication of relationship, has designated the Klug
collection as an undescribed species which it may prove to be but
as I hesitate to give him responsibility, since ex char, it seems too
near A. semidentatus, 1 propose it myself as merely a variant using,
however, the apt name under which it was distributed as a species:
var. pilosus Macbr., var. nov., ut videtur similis A. semidentata sed
Flora of Peru 377
differt foliolis papyraceo-membranaceis paullo pellucido-punctatis
et ramulis petiolisque conspicue pilosis. F.M. Neg. 21349.
San Martin : Juanjuf, 6-meter tree, KIilq S78S (type, var. pilosus).
— Rio Acre: Seringal Guanabara, Alto Xapuri, Ule 9570. To
eastern Brazil.
8. SAPINDUS [Plum.] L.
Trees with exstipulate mostly abruptly pinnate leaves and large
terminal divaricately branched panicles of small minutely bracted
and bracteolate obliquely sjmnmetric flowers, the sepals and petals
in the Peruvian species both 5, the stamens exserted, the fruits
smooth drupes.
Sapindus saponaria L. Sp. PI. 367. 1753; 639. S. peruvianus
Walp. Nov. Act. Acad. Leop.-Carol. 19: Suppl. 1: 312. 1843.
A small nearly glabrous tree with abruptly pinnate leaves, these
with 3-6 pairs of oblong-elliptic-lanceolate oblique subsessile acute
leaflets, the rachis and petiole more or less wing-margined, entire,
membranous, pale green, glabrous or softly pubescent; upper
petioles 2-7 cm. long, rachis segments 2-5 cm. long, the wings 2-6
mm. wide, petiolules 2-3 mm. long, the leaflets about 7-12 cm. long
or longer, 3-5 cm. wide; sepals glabrous except at the base and the
ciliolate margins; petals 3 mm. long, scarcely half as wide, ciliate,
sublobulate or squamate and villous above the claw; stamens ex-
serted, villous at base; disk complete, fleshy, cupulate, glabrous;
fruit usually with only one cell, yellowish, glabrous, globose as the
seed, 1.5-2 cm. in diameter. — According to Weberbauer it is found
mostly under 1,500 meters in the dry hot interandine valleys,
chiefly on the western slope. Illustrated, Radlkofer, I.e. page 645;
Ruiz & Pav6n, Fl. Peruv. 4: pi. SJ^ and many others.
The fruits with which the children play are called cholocos and
bolillos (Ruiz & Pav6n); the whitish pulp that surrounds the seed
contains saponin which foams or "suds" with water and is com-
monly used as soap, for instance to wash baize cloth (Ruiz & Pav6n) ;
the wood and roots, astringent, are said to be tonic.
Lima: Surco, Huara and Lima, Ruiz & Pav6n. Loma Zone,
Weberbauer, 148 and 150. ff Along streams, Weberbauer, 155; 162.
— Ancash :|Santa Valley near Caraz, Weberbauer, 172. Caracha,
1,200 meters, Weberbauer ^fi^.— Hudnuco: Roadside tree, S536.
San Martin: Riverbank, Juanjui, Klu^ ^^85. — Apurimac: Abancay,
Vargas. Rio Pachachaca bank, Goodspeed Exped. 1053^.— Cuzco:
378 Field Museum of Natural History— Botany, Vol. XIII
Prov. de Convencion {Hen era). Quillabamba, Soukup 135. — Tacna:
{Woitsehach). Warmer America; Africa; Oceanic Asia. "Sullucu,"
"jabonera," "cholocos," "jaboncillos" and "bolillos" (the fruits).
9. TOULICIA Aublet
Trees or tree-like shrubs with exstipulate mostly abruptly-
pinnate leaves and large subterminal sparsely branched panicles of
rather small nearly symmetric flowers, these in the Peruvian with
only 4 petals, deeply bifid scales, unilateral disk. Fruit about as in
Serjania or in Peru the cells subinflated. — In flower simulates
Talisia in part but the rounded tomentulose sepals are a dis-
tinguishing character.
Toulicia reticulata Radlk. Notizbl. Bot. Gart. Berlin 6. 154.
1914; 622.
Petioles (about 4 cm. long), sub terete rachises and leaflets
beneath as well as the laxly branched panicles even to the orbicular
sepals without softly rusty or yellowish tomentose; leaflets about
10 pairs, the lower short-ovate, about 6 cm. long, 4 cm. wide,
opposite, the upper with petiolule 5-10 mm. long, to 25 cm. long,
9 cm. wide, alternate, elliptic to broadly oblong, strongly oblique,
shortly obtuse-acuminate, acute at base, entire, subrevolute, rigid
coriaceous, glabrous and prominently reticulate above, the oblique
lateral nerves conspicuous beneath, epunctate, the epidermis lacking
mucus; flowers subsessile, 4 mm. across; disk glabrous; petals villous,
the scales filiform appendaged; fruit cells with wings 3-4 cm. long,
the former 12-14 mm. wide. — Related Amazonian species are T.
elliptica Radlk., 622, and T. bullata Radlk., 623, both with pubescent
disk, the former with glabrous alternate leaflets, those of the latter
subopposite, bullate, minutely pubescent beneath. To 30 meters
tall. F.M. Neg. 5668.
Loreto: Cachipuerto between Balsapuerto and Moyobamba,
Klug 3127 (det. Standley) .— Rio Acre: Seringal Auristella, Ule 9566,
type; 9567. Mouth of Rio Macauhdn, Krukoff 52J^7; 5560. Brazil.
10. POROCYSTIS Radlk.
Glabrous tree with subterete branches, exstipulate abruptly
pinnate leaves (petiolules and rachis obscurely sulcate) and terminal
or axillary panicles of white 4-petaled flowers borne on lateral
branchlets. Sepals 5, imbricate, concave, the 2 outer smaller.
Flora of Peru 379
Petals ovate, unguiculate, sericeous without with barbate bifid
scale within. Disk unilateral, tomentose. Stamens 8, filaments
exserted, villous. Ovary 3-celled, tomentose. Fruit consisting of
3 membranous inflated cocci cohering centrally and crowned by the
3-parted indurated style,
Porocystis toulicioides Radlk. Sitzungsb. Wiss. Miinchen
Akad. 8: 354. 1878; 630.
Small tree, the trunk diameter (known) to only 8 cm. ; flowering
branches to 1 cm. in diameter; petioles enlarged at base; leaves
3-4.5 dm. long with 7-12 alternate or subopposite oblong or elliptic
subequilateral leaflets 1.5-2.5 dm. long, 4.5-10 cm. wide with about
10 divaricate nerves anastomosing before the entire margins; panicles
equaling the leaves, on slender subflexuose branchlets, puberulent;
dichasia glomeruliform 5-11-flowered, the short pedicels articulate
below the middle, the pubescent bracts and bractlets small; flowers
scarcely wider than 3 mm.; sepals all appressed pubescent, the
petaloid inner with glabrous margins; fruits about 2 cm. high, 3-4
cm. wide, the subglobose seeds about 8 mm. thick. — Simulates
TotUicia guianensis Aublet with deeply sulcate branches and com-
pressed petioles. Since it is known in the area of Solimoes, Brazil,
it probably extends into Amazonian Peru. Illustrated, Radlkofer,
i.e. 628 (flowers, fruit, ex Pflanzenfam.) et Mart. Fl. Bras. 13, pt. 3:
pi. 108.
Peru (probably; cf. note above). Amazonian Brazil; British
Guiana.
11. TALISIA Aublet
Acladodea R. & P. Prodr. 133. pi. 29. 1794, ed. 2. 121. 1797.
Shrubs or trees, often sparsely branched and the leaves crowded
at the tips. Leaves exstipulate, abruptly pinnate, the petioles often
tumid at base, the rachises subterete or subtrigonous. Inflorescence
various, the flowers rather small with 5 sepals and petals, the disk
annulate or cucullate. Stamens 8 or 5. Fruit baccate, granulate,
ovoid or ellipsoid, rather large, mostly 1-celled and 1-seeded, the
ovary, however, 3-celled, attenuate into a subulate style. — Some
herbarium material may be confused with the genus Picramnia
of the Simaroubaceae.
Leaves pinnately trifoliolate T. peruviana.
Leaves pinnate, multifoliolate.
380 Field Museum of Natural History— Botany, Vol. XIII
Leaflets pubescent beneath, sublinear; calyx medially parted,
persisting T. pinnata.
Leaflets glabrous, suboblong or elliptic.
Calyx medially parted, persisting; stamens glabrous.
T. cupularis.
Calyx deeply parted, deciduous: stamens hirtellous.
T. cerasina.
Talisia cerasina (Benth.) Radlk. Sitzungsb. Akad. Wiss. Miin-
chen 8: 347. 1878; 845. Sapindus cerasinus Benth. in Hook. Joum.
Bot. Misc. 3: 197. 1851. S. ohlongus Benth. I.e. 198.
Small tree or shrub, glabrous except for the ample terminal
panicles, these more or less densely ashy-puberulent or white,
pyramidal with many erect or spreading branches and usually
longer than the leaves; branches stout, terete, lustrous; petioles
2-15 cm. long, the basally enlarged petiolules 2-10 mm. long, both
as the rachis subterete or early bisulcate; leaflets 3-8 pairs, opposite
or subaltemate, oblong, the lower suboblique, all abruptly short-
attenuate at the base, acutely acuminate, 4-24 cm. long, 1 longer,
1.5-8 cm. wide, subcoriaceous, lustrous at least above, drying green
or brownish, the 7-25 anastomosing lateral nerves prominent on
both sides; panicles to several dm. long and broad, the pedicels to
3 mm. long; flowers white or yellowish, 7-8 mm. long; calyx parted
nearly to base, more or less puberulent, glabrous within, the ovate
obtuse or rounded lobes ciliate, a third or a quarter as long as the
erect linear-oblong petals, these glabrous without except ciliate
below, and within as the scales densely fulvous villous, the latter
shortly bifid and as long; disk tumid, cupulate, shortly hirtellous
above; stamens 8, hirtellous, anthers linear-oblong, cordate at base;
fruit edible, cerasiform, minutely granulose, yellowish subsericeous,
apiculate. — Attains 5-10 meters (Ule). T. obovata A. C. Smith,
Brittonia 2: 154. 1936, of Bolivia and adjacent Brazil has only two
pairs of obovate leaflets 12-25 cm. long, 7-12 cm. wide, stamens 5,
the disk glabrous. Some of the flowering collections may be in-
correctly determined. The Burgos specimen from a tree to 30
meters tall, wood used for construction.
The leaves are said to furnish a black dye and a remedy for
gonorrhea.
Hudnuco: Tingo Maria, Burgos (or affine, Standley). — San
Martin: Juanjui, Klitg U303 (det. Standley). — Loreto: Yurimaguas,
Flora of Peru 881
Poeppig; Klug 2798 (det. Standley). Rio Mazdn, Williams 81 U-
Gamitanacocha, Rio Mazdn, Jos^ Schunke SOS. Mishuyacu near
Iquitos, Klug 2J^8; S89; 811; lOU; 1160; 1558; 2509 (this last
number det. Standley). Iquitos, Killip & Smith 2702U- — Rio Acre:
Seringal San Francisco, Ule 9517. Brazil. "Pitomba" (Spruce);
"juapina" (Burgos).
Talisia cupularis Radlk. Sitzungsb. Akad. Wiss. Munchen 8:
350. 1878; 852.
Tree, glabrous except for the large robust broadly pyramidal
panicles, these sometimes 0.5 meter long, 4 dm. wide, canescent
with a short velvety indument; branches terete, papillose-punctate,
4-8 mm. thick; petioles bulbous above the base, terete; leaflets
3-8 pairs, opposite or subaltemate, elliptic-oblong, somewhat
acuminate, attenuate at base into bulbously thickened petiolules
2-5 mm. long, the 7-12 lateral nerves anastomosing before the
margin and with the laxly reticulate veins rather prominent beneath,
coriaceous, drying brownish; bracts rigid; i>edicels 2-3 mm. long;
flowers white, 6-7 mm. long, the calyx parted to the middle, densely
puberulent within, the broadly ovate obtuse lobes closely ciliolate,
the twice as long oblong petals ciliate below, glabrous, eglandular,
equaled by the Ungulate entire erect scales, these densely villous
within; disk hirsute, cupulate, sinuate-pentagonous; stamens gla-
brous; fruit ovoid, yellowish-pubescent as the rather long style, the
persisting calyx appressed-cupulate. — The Krukoff specimen from
a tree 30 meters tall; vegetatively scarcely distinguishable from
T. cerasina. Illustrated, Mart. Fl. Bras. 13, pt. 3. pi. 113.
Rio Acre: Mouth of the Rio Macauhdn, Krukoff 5725; 56S0;
5787. Amazonian Brazil.
Talisia peruviana Standi. Field Mus. Bot. 11: 165. 1936.
Shrub or small tree 2.5-6 meters high, the slender subterete
branchlets minutely strigillose; leaves alternate, pinnately 3-foliolate,
slender-petioled, the leaflets shortly petiolulate (to 5 mm.), oblong-
elliptic, lanceolate-oblong, or obovate, 5.&-18 cm. long, 2-8 cm.
wide, subabrupt-acuminate or often long-cuspidate-acuminate, acute
or cuneate at base, glabrous, minutely and densely pellucid-punctate,
membranous; inflorescence racemiform, half as long as leaves, few-
flowered, pedicels 2-3 mm. long, densely strigillose, little elongate
in fruit, bracts minute; sepals 1.5 mm. long, obtuse, sericeous;
petals cuneate-obovate, 2 mm. long; disk thick, glabrous; ovary
382 Field Museum of Natural History— Botany, Vol. XIII
2-celled, strigillose; styles bifid to base, the branches short; fruit
broadly oval, to 2 cm. long, yellowish-green, rounded both ends or
somewhat narrowed at tip and obtusely tubercled, 2-celled; seeds
large, solitary in each cell. — Near to T. japurensis (C. DC.) Radlk.?
Generic position uncertain, ovary of Talisia said to be early 3-celled.
San Martin: Tarapoto, Williams 57^5, type; 6652; 6578; 6208.
Pongo de Cainarachi, Klug 2713. Loreto: Puerto Arturo, Williams
5218; 5118; 5303; Killip & Smith 27915; 2787 A. Yurimaguas,
Killip & Smith 276^5; 27673; 27636. Balsapuerto, Killip & Smith
2851^9; 28^8A; 28627. Santa Rosa, Killip & Smith 28857. "Sinca
zanango" (Klug).
Talisia pinnata (R. & P.) Radlk. Sitzungsb. Akad. Wiss. Miin-
chen 8: 351. 1878; 856. Acladodea pinnata R. & P. Syst. Veg. 262.
1798. Talisia acladodes Spreng. Syst. 2: 223. 1825.
A small unbranched tree, the stem crowned apically by the
crowded pinnate leaves and ample rusty tomentose panicles of
whitish flowers; petioles to more than 2 dm. long, tomentose, en-
larged above the base to 5 mm. thick, the multifoliolate leaves even
to 1 meter long, 2.5-4 dm. wide; petiolules 2-4 mm. long, the long
rachis like the petiole terete and striate, the leaflets alternate,
oblong-linear, obliquely subacute at base, acuminate, 1-2.5 dm.
long, 2.5-5 cm. wide, glabrous and lustrous above, bullate toward
the margins, membranous, drying brown, softly pubescent beneath,
especially on the prominent reticulate veins and many approximate
nerves; panicles about 3 dm. long, pyramidal, much branched, with
many minutely pinnate bractlike leaves; pedicels scarcely 2 mm.
long; flowers nearly 7 mm. long, the calyx medially divided, densely
reddish tomentose, the obtuse broadly ovate lobes ciliate, half as long
as the broadly ovate petals, reflexed, ciliate below, medially below
the tip fulvous tomentose, the tomentose erect shortly bifid scales
as long; disk cupulate, very hirsute above as the filaments, the
linear-oblong anthers cuspidate. F.M. Neg. 23636.
Hudnuco: In woods at Chacahuassi near Pillao, Ruiz & Pavon,
type.
12. DILODENDRON Radlk.
Tree or shrub allied to Cupania but the large leaves abruptly
bipinnate, the 3-4, rarely 5 petals (or lacking in male flowers)
esquamate, the sublobate concave disk glabrous. Seeds with short
basal cupulate aril. The shrub is leafless when in flower.
Flora of Peru 888
Dilodendron bipinnatum Radlk. Sitzungsb. Akad. Wiss.
MUnchen 8: 357. 1878; 1067.
Petioles and rachises obtusely triangular-sulcate, early hirtellous,
the 3-7 pairs of alternate or subopposite oblong pinnae reduced
toward the base, each with 4-9 pairs of oblong-lanceolate subsessile
acute more or less serrate leaflets 4-6 cm. long, 1-2 cm. wide, many-
nerved, smooth and glabrous above, minutely tuberculate-papillose,
hirtellous, stipitate glandular, opaque and glaucescent beneath,
minutely pellucid-punctate, containing mucus; panicles at tips
of defoliate branches or fasciculate on new branches above axillary
scales, flavescent-tomentulose, sometimes paniculate, 8-25 cm. long;
bracts and bractlets small; pedicels short, articulate at base; male
flowers with glabrous filaments, glabrate anthers, 3 mm. long, the
female 5 mm. long, the anthers glandular-setulose; capsule elliptic-
trigonous, 3-valved, 1.5 cm. long and broad, glabrous, within
hirsutulous. — Sometimes 10 meters tall or taller. Illustrated,
Radlkofer, I.e. 1066.
The seeds are said to provide an oil used for light and for food.
The rather soft wood has an unpleasant odor.
Cuzco: Prov. Convencion, Santa Ana, 1,100-1,300 meters,
Weberbauer 5020; 50ItS; SOU; 277; 316. Bolivia; Paraguay; Brazil.
13. CUPANIA [Plum.] L.
Trees or tall shrubs with terete more or less sulcate and lenticel-
late branches that are sparsely leafy with exstipulate actually but
not always clearly abruptly pinnate leaves that often exceed the
panicles or small regular flowers. Sepals free, 5, imbricate in 2 rows,
usually fleshy or subcoriaceous, mostly subequaling or a little
longer than the petals. Disk regular, annulate. Stamens of male
flowers exserted, 8 (-10). Fruit a capsule, trigonous to somewhat
3-lobed, dehiscing by 3 (2-^) lobes each provided with one ellipsoid
or obovoid seed more or less enclosed by the fleshy aril.
Capsules turbinate-triangular, shortly stiped; leaflets hirtellous or
tomentulose.
Leaflets papillose between the veins and hirtellous.
C. papillosa, C. latifolia.
Leaflets tomentose beneath and epapillose C. cinerea.
Capsules turbinate, more or less 3-comute, long (3-6 mm.) -stiped;
leaflets glabrescent C. scrobiculata.
384 Field Museum of Natural History — Botany, Vol. XIII
Cupania cinerea Poepp. & Endl. Nov. Gen. & Sp. 3: 38. 1844;
1031.
Younger branches puberulent; petioles 2-5 cm. long; leaflets
6-10, petiolulate, alternate, obovate-oblong, acute at base, obtuse
or subtruncate, 5-15 cm. long, 2-5 cm. wide, repand, serrate,
coriaceous, discolored, brownish above, more or less cinereous with
a minute and close puberulence beneath especially on nerves and
veins, epapillose, subeglandular, usually lustrous and with hypo-
derma above; panicles equaling or exceeding the leaves, minutely
tomentulose including the oval 2.5 mm. long sepals, disk and
turbinate- triangular capsule, this densely tomentose within, 1.5 cm.
long; cymules sessile, the bracts and bractlets 2-3 mm. long, the
pedicels 1-2 mm. long; petals white, brownish within (Woytkowski),
2 mm. long, nearly glabrous, the scales a third as long, laxly pu-
bescent; stamens pilose near base. — C. polyodonta Radlk., 1030, of
Ecuador has 4-6 hirtellous-puberulent leaflets. There are a number
of species known from Colombia and northern Brazil to Venezuela
that of course, may be found within Amazonian Peru. Determina-
tions mostly by Standley. F.M. Neg. 31044.
Often about 10 meters tall, beautiful (Poeppig), or nearly 20
meters with straight round chocolate-brown barked trunk 2 dm. or
more in diameter, the crown spreading (Williams).
San Martin: Tarapoto, Spruce W2; Williams 6023; Woytkowski
35078. San Roque, Williams 730^. — Loreto: Yurimaguas, in margin
of woods, Poeppig 3096; Diar. 2338, type; Klug 2795. Florida, Klug
2102; 2335. Rio Mazan, Jos^ Schunke 3^5. Bolivia; Colombia.
"Sama" (Herzog), "puca yacu," "huapina" (Williams), "fuapina"
(Woytkowski), "sama" (Bolivia, Herzog).
Cupania latifolia HBK. Nov. Gen. & Sp. 5: 126. 1821; 1030.
Younger branches terete, puberulent, soon glabrescent and
lenticellate; leaves ample, often 3-4 dm. long with 5-6 mostly
alternate leaflets, the upper obovate, the remaining obovate-oblong,
all emarginate, retuse, subcuneate at base, remotely and equally
serrate, more or less petiolulate, 8-20 cm. long, 4.5-5 cm. wide,
coriaceous, obscurely subclathrate-veined, glabrous above, minutely
asperulous and sparsely papillose beneath; panicles 2-3 dm. long,
laxly branched, tomentulose, bracts and bractlets subulate, 2-3 mm.
long; pedicels 1 mm. long; sepals ovate, nearly 3 mm. long, sub-
coriaceous tomentulose, the unguiculate petals about as long,
pubescent at base, the 2 scales a third as long, pubescent; disk
Flora of Peru 885
tomentulose; stamens 3 mm. long, the basal trichomes reddish; ovary
tomentose; capsule obovate-trigonous, shortly stiped, tomentose
within and without, 1.5 cm. long, the obovoid basally arillate seed
1 cm. long. — C. polyodorUa, 1030, of Ecuador is similar but the
leaves are said to be epapillose. F.M. Neg. 5689.
San Martin: Juanjui, Kltig ^266 (det. Standley, sp. nov.). —
Amazonas: Tactamal to Vilaya, Prov. Luya, 1,400 meters, Weber-
bauer 71JtS. — Cuzco: Macchu-picchu, Soukup 205; 865. To Panama,
Venezuela.
Cupania papillosa Radlk. Sitzungsb. Akad. Wiss. Miinchen 9:
520. 1879; 1029.
Small tree with rusty tomentulose flowering branchlets including
the large panicles, the older glabrescent and lenticellate; leaves
ample, often 2.5-3 dm. long, typically with 6-10, apparently often
fewer, alternate or subopposite oblong leaflets, obtuse or in Peru
somewhat retuse, shortly acute at base, in type 6-15 cm. long, 2-5
cm. wide, in Peru to 3 dm. long, half as wide (C. latifolia?), more or
less serrate-dentate, coriaceous, the nerves and veins beneath
hirtellous tomentulose, papillose between the prominently reticulate
veins, glabrous above; pedicels 1-2 mm. long; sepals ovate, tomentu-
lose, 2 mm. long scarcely equaled by the basally pubescent clawed
petals; disk tomentose; capsules turbinate-triangular, shortly stiped,
tomentose within and without, 1-1.5 cm. long, the subglobose
basally arillate seed nearly 1 cm. long. — Toward C. americana L.
with subglobose capsules and, especially C. latifolia HBK., 1030,
with flowers 3 mm. long and obovate subretuse leaflets and it seems
probable that at least as regards the Peruvian material, there is no
substantial distinction.
Peru (possibly; cf. note above). Colombia.
Cupania scrobiculata Rich. Act. Soc. Hist. Nat. Paris 1: 109.
1792; 1051.
Branches sulcate, early pulverulent; leaflets mostly 6-8, rarely
10-12, sometimes subopposite, oval or elliptic-oblong or oblong,
obtusish to acuminate, entire or repand-dentate, petiolulate or sub-
sessile, chartaceous-coriaceous obviously clathrate and reticulate-
veined beneath and mostly notably foveolate in the nerve axils,
glabrescent both sides, lustrous or sometimes opaque beneath,
obsoletely pellucid-punctate and lineolate. usually 5-15 cm. long,
2-2.5 cm. wide, panicles axillary or subterminal, often 2-3 dm. long,
386 Field Museum of Natural History— Botany, Vol. XIII
pulverulent tomentulose; bracts subulate-filiform, about 2 mm. long;
pedicels scarcely 1 mm. long, about 2 mm. long in fruit; disk glabrous;
sepals 1.5 mm. long, oval-oblong, chartaceous, tomentulose without,
nearly equaled by the acute or obtuse ciliolate petals; stamens 2 mm.
long, pubescent below the middle; capsules turbinate, more or less
tricornate, 1-2 cm. long contracted into stipe 3-6 mm. long, reddish
tomentose without, lanate within, the black ellipsoid seed two-thirds
arillate.— Illustrated, Mart. Fl. Bras. 13, pt. 3: pL 115.
Rio Acre: Mouth of Rio Macauhdn, Krukoff 5600; 5523; 552U.
To Panama and the Guianas.
14. MATAYBA Aublet
Shrubs or trees similar to Cupania except that the small calyx is
cupulate, opening early, 5-dentate-lobed, the segments subimbricate
only at base and, in the Peruvian species, the leaflets are entire.
Species mostly little known and the Peruvian collections are too
few or too incomplete for definite determination.
Leaflets 2-4 cm. long, 1-1.5 cm. wide M. peruviana.
Leaflets much larger.
Leaflets 3-5, 7-12 cm. wide, drying pale green M. purgans.
Leaflets 6 (2)-12, 1.5-5.5 cm. wide, drying dark.
Fruits glabrous within as without; leaflets 2-6. .M. arbor escens.
I^Yuits densely tomentose within; leaflets to 12. . .M. guianensis.
Matayba arborescens (Aublet) Radlk. Sitzungsb. Akad. Wiss.
Miinchen 9: 535. 1879; 1082. Sapindus arborescens Aublet, PL
Guian. 1: 357. pi. 139. 1775.
Glabrous, except the tips of the young branches, these yellowish-
tomentulose; leaflets usually 6 (3-8), alternate or subopposite,
elliptic-lanceolate or lanceolate, acuminate, entire, petiolulate or
subsessile, chartaceous, hypodermatous tissue lacking, densely
pellucid-punctate, 7-21 cm. long, 4-8 cm. wide, the 9-15 lateral
nerves rather prominent on both sides; petiolules 2-5 mm. long,
canaliculate above, bulbous at the base; panicles shortly tomentu-
lose or laxly puberulent, sometimes subequaling the leaves; bracts
and bractlets minute; cymules of male flowers subsessile, those of
the female more or less stiped; pedicels about 1.5 mm. long, 3-4 mm.
long in fruit, calyx scarcely exceeding 1 mm., deeply lobed, the acute
chartaceous-coriaceous lobes appressed puberulent; petals acute.
Flora of Peru 387
about 1 mm. long, pubescent, exceeded by the lanate scales; stamens
nearly 3 mm. long, the filiform filaments pubescent below, the
anthers puberulent; ovary appressed pilose; capsules 1.5-2 cm. long
and broad, the black lustrous seed with white aril. — The Peruvian
collections are all in fruit and the determination is certainly open
to question.
Loreto: Mishuyacu near Iquitos, Kliig 2508; 251S; 6S6. To the
Guianas.
Matayba guianensis Aublet, PI. Guian. 1: 331. pi. 128. 1775;
1097.
Much branched above, the subterete or lightly sulcate branches
glabrescent or puberulent; petioles somewhat complanate and pu-
bescent above; leaves 1.5-3.5 dm. long with 2-12 usually oblong-
ovate-lanceolate leaflets, sometimes with some smaller ones between
them, commonly rather acutely acuminate, more or less contracted
to the short or longish petiolules, 5-15 cm. long, 1.5-5.5 cm. wide,
few-nerved, 1-m'^y foveolate (or not at all), glabrous or lightly
pilose only beneath, subimpunctate or densely so and lineolate;
panicles ample or contracted, equaling or exceeding the leaves,
puberulent or tomentulose, the cjrmules shortly stiped, the white
flowers on pedicels 1-2 mm. long; calyx 1-nearly 2 mm. long, cori-
aceous, the lobes rounded, somewhat puberulent; petals 1-2 mm.
long, equaled by the reddish lanate scales; disk glabrous; capsule
trigonous-subglobose, shortly stiped, 1-2 cm. long, more or less
verruculose, glabrate without, densely tomentose within. — Variable.
The Killip & Smith species in fruit doubtful, the capsule long-stiped.
According to the author attains 20 meters.
San Martin: Zepelacio, Klug S658 (det. Standley). — Hudnuco(?):
Haenke (fide Radlk.).— Loreto: Santa Rosa, Killip & Smith 28782?
To the Guianas.
Matayba peruviana Radlk. Sitzungsb. Akad. Wiss. MUnchen
9: 536. 1879; 1080.
Branches terete, early tomentulose, with few leaves of 8-14
opposite or subaltemate shortly lanceolate obtusely acuminate
chartaceous-coriaceous many-nerved petiolulate leaflets 2-4 cm.
long, 1-1.5 cm. wide, acutish at base, glabrous above, appressed and
sparsely pubescent beneath except on midnerve, barbate on the
many round foveoli and with immersed digitiform glands, minutely
lineolate and punctate-pellucid; petioles and rachises tomentose.
388 Field Museum of Natural History— Botany, Vol. XIII
especially above; panicles axillary, solitary, subequaling the leaves,
tomentose, the white flowers shortly pedicelled; calyx nearly 1 mm.
long, coriaceous, appressed setulose pilose without; petals 1.5 mm.
long, the 2 lanate scales a third as long; stamens puberulent at base;
disk glabrous; style 2-3-lobed, equaling the setulose ovary. — A
branched tree about 4 meters tall (Spruce) . F.M. Negs. 6037; 23630.
San Martin: Guayrapurina near Tarapoto, Spruce j^619, type.
Loreto: Ule 6719. "Canela ucsha."
Matayba purgans (Poepp. & Endl.) Radlk. Sitzungsb. Akad.
Wiss. Miinchen 9: 536. 1879; 1094. Cupania purgans Poepp. &
Endl. Nov. Gen. & Sp. 3: 38. 1844.
Shrub or tree 1-6 meters high with subterete lightly sulcate
glabrescent branches and large leaves with 3-5 alternate or sub-
opposite leaflets, the glabrous common petioles 9-12 cm. long,
complanate above, convex beneath; leaflets petiolulate, broadly
elliptic or the lower subovate, acutish or subrounded at base, very
obtusely acuminate-apiculate, 15-30 cm. long, 7-12 cm. wide, char-
taceous, efoveolate, green both sides, microscopically glandular and
pilose, otherwise glabrous, obscurely pellucid-punctate, hypoderma
none, nerves only about 8; panicles thjnrsoid, scarcely half as long
as the petioles, mostly 2-3-fasciculate, rusty tomentulose, the
cymules sessile, the pedicels 1 mm. long, the small flowers white;
calyx and petals 1 mm. long, the former tomentulose, the 1 mm. long
scales lanate; capsule shortly stiped, subchartaceous, glabrate
without, the endocarp pilose, 18 mm. long, 16 mm. broad, the dark
seed enclosed by the bluish viscid-succulent aril. — The name refers
to Poeppig's observation as to the property of the seeds. Illustrated,
Mart. Fl. Bras. 13, pt. 3: pi. 120.
San Martin: Tarapoto, Ule 6U^; Spruce; Williams 6^93. —
Loreto: Balsapuerto, Klug 2950 (det. Standley); Killip & Smith
28528 (det. A. C. Smith). Amazonian Brazil. "Itua" (Poeppig).
15. LLAGUNOA R. & P.
Shrub or tree distinctive in Peru by the conspicuous green or
slightly purple-tinged flowers, 1-7 on rather long pedicels in the
axils of the simple or mostly simple leaves. Flowers oblique with no
petals but a prominent unilateral widely expanded disk, in the
Peruvian species unequally lobed. Stamens a little exserted in
the male flowers, the filaments glabrous or merely puberulent.
Â¥
Flora of Peru 389
Style subulate-filiform. Capsule subglobose-trilobed, pendulous,
3-valved, each valve with a solitary lustrous black globose seed
without aril.
Llagunoa nitida R. & P. Syst. Veg. 252. 1798; 1343. Amirola
nitida (R. & P.) Pers. Syn. PI. 2, 565. 1807.
Young branches, petioles (1-2 cm. long) and inflorescences
(2-3 cm. long) lightly pubescent, finally glabrous; leaves simple or
trifoliate, the lateral then small, the terminal sometimes lobate,
dentate, broadly ovate-elliptic, sometimes oblong-elliptic, acutish
or obtuse, rounded or narrowed at base, 6-11 cm. long, 2-5.5 cm.
wide, glabrous or even tomentose beneath (var. mollis (HBK.)
Radlk.), the younger densely glandular; dichasia usually 3-7-
flowered, the peduncles and 5-15 nmi. long pedicels glandular and
puberulent as the calyx within, this expanded 1-1.5 cm.; stamens
6 mm. long, typically glabrous with red anthers; fruit crustaceous,
15-18 nrni. across, the seeds 6 mm. in diameter. — Trunk said to
attain at least 1 dm. Illustrated, Hook. Icon. 2: pi. 132. F.M.
Neg. 36021.
Known, according to Ruiz and Pav6n, as the "Arbol de cuentas
de rosario," in reference to the use of the lustrous black round seeds
for the making of rosaries; the wood beautiful for cabinets, being
white with black markings (Stork & Horton).
Cajamarca: Prov. Ja^n, Weberbaiier 6179. — Amazonas: Chacha-
poyas, Mathews; Weberbauer 1^302; 191. Llata, 2283 (det. Johnston).
— Ancash: Valley of the Rio Puccha, Weberbauer 3736; 174.—
Hudnuco: Mufla, 3935; Ruiz & Pav&n, t5rpe. — Apurimac: Valley
Rio Pincos, Weberbav£r 5915 (robust and leaves oblong-elliptic).
— Ayacucho: Tambo, Weberbauer 55Jt9. — Huancavelica: Prov. Taya-
caja, hills and ravines, 2,000 meters, Stork & Horton 10JtO6. — Cuzco:
Valley Urubamba, Weberbauer 5069; 237. Torontoi, 2,400 meters,
Cook & Gilbert 1770. Prov. Andahuaylas, 2,600 meters, Vargas
8801.— Puno: Sandia, Weberbauer, 245. Ecuador. "Arbol de rosa-
rio."
16. DODONAEA L.
Reference: Sherff in Field Mus. Bot. 23: 269-317. 1947.
Resinous-viscid shrubs with subopposite exstipulate simple
leaves and small dioecious or hermaphrodite regular flowers, pedi-
celled in panicled racemes, corymbs or panicles. Sepals often 4,
valvate or narrowly imbricate, finally reflexed and deciduous.
390 Field Museum of Natural History— Botany, Vol. XIII
Petals none. Disk obsolete but in the female flowers a short carpo-
phore. Stamens 5-8, the filaments very short. Fruit a 2-5-angled
or winged capsule that simulates a samara; the cells 2- (or 1-)
seeded.
Dodonaea viscosa (L.) Jacq. Enum. Carib. 19. 1760; 1363.
Ptelea viscosa L. Sp. PL 118. 1753.
Glabrous viscous shrub-tree, the erect reddish-brown flowering
branches compressed, angled and with an elevated line below the
insertion of the more or less petioled leaves, these varying from
subobovate-cuneate to lanceolate or sublinear, acute or obtuse,
entire but the margins sometimes unequally subrepand, to 15 cm.
long, 2 cm. or so wide, subchartaceous, pale green, lustrous and
with many fine lateral nerves; panicles in flower about half as long
as the leaves, the 5-8 mm. long pedicels twice as long in fruit; flowers
greenish-white or reddish, 3 mm. long often hermaphrodite; sepals
usually 4, trinerved, puberulent marginally; capsule suborbicular,
excised apex and base or subcordate, mostly triquetrous, 3-celled,
each cell surrounded by a membranous radiately reticulate-veined
often roseate wing. †” Herrera in Contrib. Fl. Cuzco, ed. 1. 2: 118
gave two names to Raimondi collections, both without description,
which ought not to be cited even as synonyms. Sherff has thought
it worth while to reinterpret the many variations, those in Peru
being var. linearis (Harv. & Sond.) Sherff, f. angustifolia (Benth.)
Sherff (Stork & Horton 1005; Vargas 9781, leaves 5-11 cm. long);
var. vulgaris Benth., f. Burmanniana (Schum. & Thon.) Radlk.
(Stork & Horton 1078j^; lOUlS), leaves 7-11 by 1.5-2 cm., acute
or obtusish and f. Schiedeana (Schlecht.) Radlk., leaves to 16
cm. long, 2 cm. wide, gradually acuminate both ends; var. arbor escens
(Cunn.) Sherff, f. spatulata (Sm.) Sherff (Chachapoyas, Williams
7564), leaves often more or less sinuate-dentate.
Common, especially in the interandean valleys between 1,000
and 3,000 meters and known everywhere by the native name
"Chamana" (Weberbauer). Used for fuel; crushed leaves applied
effectively in poultices for sprains (Ruiz & Pavon). Leaves mixed
with coca if latter too strong and with branches used for mattresses,
the gum sticking them together firmly (Stork & Horton).
Cajamarca: Socota, Stork & Horton 10095. Above Santa Cruz,
Weberbauer, 189. — Amazonas: Chachapoyas, Williams 7564- —
Ancash: Caraz, 2,200 meters, Weberbauer 3010; 173. Huantar,
Puccha Valley, Weberbauer, 174. Grass steppes, Rio de Chiquian,
Flora of Peru 391
Weberbauer, 111. — Hudnuco: Near Hudnuco, 2052; Kanehira 6J^,
Mito, SS35. Below Ambo, dominant on dry rocky eastern slopes,
2^16. Uspachaca, ISIS. San Rafael, Sawada Pi 20. Valley of the
Marafl6n, Weberbauer, 190. Valley of Utcubamba, Weberbauer,
191.— Junfn: Tarma, Killip & Smith 21818; Ruiz & Pav6n. Tarma
Valley, 2,700 meters, Weberbauer 2S87; 176.— Ayacucho: Near Rio de
Lomas, Weberbauer 575S. — Huancavelica: Stork & Horton lOUlS.
— Apurimac: Stork & Horton 1078 j^; Vargas 9781. — Cuzco: Prov. de
Paruro, Raimondi. Pomachaca, Urubamba, Weberbau£r 50U9; 211.
Ollantaitambo, Cook & Gilbert 7S7. — Puno: Sandia, Weberbauer
5^7; 238. All warm regions. "Chamana," "chamisa" and "cha-
massa" (Ruiz & Pav6n).
RHAMNACEAE. Buckthorn Family
Commonly shrubs (Gou^nia, Ampelozizyphus, Sageretia, scan-
dent) or small trees, rarely herbs, often thorny or the stipules
spiniform, these otherwise small or obsolete, always simple-leaved
and the flowers regular, usually 5- (or 4-) merous and in little
umbels that frequently are racemose or panicled. Petals, if present,
ordinarily cucullate or involute, and inserted with the stamens into
the edge of the thin or fleshy disk which often lines the short or
long calyx-tube and sometimes unites it to the 2-5-celled (rarely
incompletely 2-celled) ovary, the cells usually 1-ovuled. Fruit
a drupe with 1-3 pyrenes or a capsule with 1 erect seed in each cell,
or less often consisting of 3 cocci that may be winged.
Family known for several products, especially for "cascara
sagrada," obtained from Rhamnu^ Purshiana DC. of western North
America, and R. cathartica L. of the Old World. The fruit "jujube"
(Zizyphu^ Jujuba Mill.) of the Mediterranean region and the similar
Z. mauritiana Lam. may be cultivated in Peru; unlike the Peruvian
species these shrubs have fruits about the size of olives, reddish or
yellowish with a sweet edible pulp, and the latter species is tomentose.
For an excellent account of these and other cultivated Rhamnaceae
in Argentina, some of which of course are probably grown in Peru
for ornament or for hedges, see Marzocca y Marthi, Ministr.
Agric. y Ganad. 7, fasc. 120: 1-48. 1951.
The key has been devised of course as an aid in the determination
of Peruvian components of the family and not to suggest possible
relationships or to give technical characters.
392 Field Museum of Natural History— Botany, Vol. XIII
Shrubs, rarely scandent but then without tendrils and lateral nerves
not parallel.
Leaves multinerved, the nerves from midnerve subparallel; fruit
capsular 1. Alzatea.
Leaves medially 3-nerved from at or near the base.
Erect shrubs; fruit drupiform 2. Zizyphus.
Liana, long-leaved; fruit capsular 7. Ampelozizyphus.
Leaves pinnately nerved, sometimes lacking except in vegetative
(growing) periods.
Plants sparsely leafy, the small leaves often absent except in
growing season; branchlets, unless younger, often thorny;
fruit drupiform, or tardily capsular.
Leaves as branchlets opposite or nearly; ovary 2-4-celled.
Flowers sessile or nearly; fruits drupiform 6. Scutia.
Flowers pendent; fruits finally capsular 10. Colletia.
Leaves as branchlets alternate; ovary incompletely 2-celled.
3. Condalia.
Plants abundantly foliose; spines if present small, stipular.
Leaves opposite or subopposite (Peru).
Flowers sessile in open inflorescence; upper stems often
scandent; drupe with 3 nutlets 4. Sageretia.
Flowers pedicellate or subsessile but crowded; erect shrubs
or trees.
Caljrx tube persisting but not adhering to baccate fruit;
leaves without basal glands 5. Rhamnidium.
Calyx tube adhering basally to capsular fruit; leaves
(Peru) biglandular 8. Colubrina.
Leaves alternate; fruit baccate or drupiform.
Shrubs never spiny; fruit somewhat fleshy, indehiscent.
9. Rhamnus.
Shrubs often spiny; fruit baccate, the carpels finally
dehiscing 8. Colubrina.
Liana or clambering, tendrils usually present, the leaves with sub-
parallel lateral nerves 11. Gouania.
1. ALZATEA R. & P.
Generic character that of the single species. — Seems affine
MayteniLS but apetalous (DeCandolle) ; some of its characters
Flora of Peru 393
suggest affinity to Icacinaceae; but according to Loesener, Pflanzen-
familien, ed. 2, 20b, it may be a part of Rhamnaceae. The leaves
simulate those of some Guttiferae.
Alzatea verticillata R. & P. Fl. Peruv. 3: 20, pi. 2^1. 1802.
Glabrous stout-trunked tree with purplish verticillate branches,
opposite and verticillate petioles, ample coriaceous entire oblong-
obovate leaves, lustrous above, and many-flowered terminal corymbs
of lutescent apetalous flowers; calyx campanulate, colored, 5-parted;
stamens 5, apparently inserted on disk, filaments short, anthers
erect, cordate; style short, stigma obtuse; ovules affixed centrally
base to apex, the sessile ovary obcordate and 2-celled as the bivalved
several-seeded capsule; seeds winged, aril none. — Illustrated, Ruiz
& Pav6n, Prodr. pi 7. F.M. Neg. 29349.
Hudnuco: In woods near Mesapata, Chinchao, Ruiz & Pav6n,
type. Bolivia (fide Rusby).
2. ZIZYPHUS [Miller] L.
Commonly conspicuously thorny, the alternate subdistichous
coriaceous leaves 3-plinerved. Flowers in short axillary cjones.
Petals 5 (rarely none), cucuUate, deflexed. Calyx 5-parted with
broadly obconic tube persisting beneath in fruit, the ovate acute
spreading lobes carinate within and 3-angled. Disk plane, margin-
ally pentagonous, free. Ovary superior or semisuperior, 2- (rarely
3-4-) celled. Drupes somewhat fleshy, 1-3-seeded. — Originally
Ziziphus, derived from the Arabian or Greek native name.
Leaves subrotimd, villous on nerves beneath Z. piurensis.
Leaves oblongish, glabrous or puberulent.
Leaves puberulent, 1-1.5 cm. wide Z. Weherhaueri.
Leaves glabrous, 3-6 cm. wide Z. cinnamomum.
Zizyphus cinnamomum Tr. & PI. Ann. Sci. Nat. s^r. 5. 16:
380. 1872.
Cinereous-barked branchlets with numerous oblong lenticels;
leaves oblong-elliptic, obtuse, to 1.5 dm. long, half as wide, glabrous,
coriaceous, glaucescent, very finely and closely reticulate- veined,
the 3 primary nerves prominent; peduncles short, arcuate-reflexed,
lenticellate; pedicels and calyces tomentulose (authors). F.M. Neg.
4703.
394 Field Museum of Natural History— Botany, Vol. XIII
Loreto: Pongo de Manseriche {Tessmann ^703, det. Mansfeld).
Colombia.
Zizyphus piurensis Pilger, Bot. Jahrb. 54: Beibl. 117: 46. 1916.
Younger branchlets, petioles, these to 1 cm. long, and axillary
and terminal cymules densely puberulent; leaves broadly ovate,
usually subrotund, rounded both ends, distinctly crenate or obtusely
serrate-crenate, 5- (8) cm. long, 3-5 cm. wide, chartaceous, glabrous
above, villous beneath, especially toward the base of the 3 principal
nerves, the dense reticulate venation faint; sepals obviously carinate
within, 1.5 mm. long, equaled by the petals. — Allied to Z. thyrsiflora
Benth. of northern Ecuador with much less pubescent leaf-veins
and cymules; that species also has more remotely crenulate leaves
that are firmer and lustrous above. However, more material may
show the Peruvian tree, known to attain 8 meters, only a variant.
There is also a resemblance to Z. undulata Reiss. of Brazil, glabrate,
the leaves acutish.
Piura: Morropon to Salitral, 150 meters, Weberhauer 5962, type.
"Palo negro," "evano" (Ruiz & Pavon for Z. thyrsiflora).
Zizyphus Weberbaueri Pilger, Bot. Jahrb. 54: Beibl. 117: 46.
1916.
Branchlets divaricate, geminately short-spinose at some of the
upper nodes, the spines mostly 5-10 (14) mm. long; petioles 4-5 mm.
long, canescent puberulent as the young branchlets and small
axillary and terminal cjntnes including the scarcely 1.5 mm. long
sepals; leaves rather rigid, nearly concolorous, sparsely puberulent,
beneath on the 3 principal nerves especially toward the base, densely
reticulate-veined, minutely serrulate or subentire, ovate-oblong-
lanceolate or rarely long-oval, 1.5-3.5 cm. long, 8-16 mm. wide;
petals narrowly cochleate-spatulate, 1.5 mm. long; disk rather thick,
lightly crenulate; fruit reddish, globose-ellipsoid, 1 cm. long. — To
6 meters tall. Related to Z. mistol Griseb. of Argentina and to Z.
ohlongifolius Sp. Moore of Brazil, the last with different shaped
leaves, shorter spines, the first with leaves more glaucous beneath
and less pubescent cymes, smaller fruit (Pilger). The species of
Spencer Moore seems to be nearest but that is glabrous; the validity
of these species depends on the variability of these characters, as yet
unknown.
Apurlmac: In savana below Curahuasi, 2,100 meters, Weber-
hauer 5920, type.
Flora of Peru 395
3. CONDALIA Cav.
Much branched, the often spreading branchlets usually ter-
minating in a rigid spine. Leaves alternate, entire, with minute
stipules. Flowers axillary, solitary or usually in small umbels with
several flowers, sometimes panicled. Calyx deeply 5-lobed; petals
often none. Ovary free from both calyx and disk, incompletely
2-celled with 1 or 2 ovules; style base persisting on the drupe. — The
euphonious name commemorates a Spanish physician, A. Condal,
companion of Loefling on his voyage to the Rio Orinoco.
Gondalia Weberbaueri Perk. Bot. Jahrb. 45: 463. 1911.
Shrub with numerous short apically subulate-spinescent branch-
lets, these early sparsely gray-pilose as the 1.5-3 mm. long petioles;
leaves obovate or obovate-oblong, long-attenuate to base, acutish,
mucronate, mostly 1-2.5 cm. long, 3.5-9 mm. wide, entire, glabrous
above, grayish-pilose or on the 6-7 lateral nerves tomentose beneath;
stipules ovate, persisting; flowers axillary, solitary or fasciculate,
3.5 mm. long, the glabrous filiform pedicels 3-6 nmi. long; calyx
membranous, lightly pilose without, the ovate acuminate lobes
spreading; petals none; ovary glabrous, 2-celled(?), the ovules
soUtary; drupes ovoid, 7-8 mm. long, 5 mm. thick, ligneous, the style
5 nrni. long. — Easily distinguished from the related C. huxijolia
Reiss. by the form of the pubescent leaves and the pubescent calyx
(Perkins); my collection from a shrub to 1 meter high, fruits pur-
pUsh. F.M. Neg. 5840.
Junin: Tarma, Ruiz & Pav&n; Weberbauer 172J!f, type; 1076. —
Hudnuco: Hudnuco and Pillao, Ruiz & Pavdn. "Tanacancha"
(Ruiz & Pav6n). Bolivia.
4. SAGERETIA Brongniart
Resembles Scuiia but the minute flowers disposed in panicles
or glomerulate in the axils of the opposite or suboppK)site often
decussate branchlets. Disk cupulate, marginally 5-lobed. Ovary
3-celled. Drupes globose with 3 indehiscent pyrenes.
Sageretia elegans (HBK.) Brongn. Ann. Sci. Nat. s^r. 1, 10:
359. 1827. Rhamnus elegans HBK. Nov. C^n. & Sp. 7: 53, pi. 619.
1824.
Branches more or less spinescent, the slender virgate upper ones
often somewhat scandent, usually densely canescent puberulent
396 Field Museum of Natural History— Botany, Vol. XIII
as often the younger leaves and the leafy panicles of sessile greenish-
white flowers; leaves lanceolate to ovate-elliptic, glabrous (or nearly)
and lustrous at maturity, subcoriaceous, rounded or subcordate
at base, acute or acuminate, usually 4-9 cm. long; calyx laxly
tomentulose, 1-1.5 mm. long; fruit black or dark, subglobose,
6-8 mm. in diameter.
San Martin: Lamas, Williams 6U17. Tarapoto, Williams 61 3 U;
6251; 6503; Spruce U16. To Mexico.
5. RHAMNIDIUM Reissek
Shrub or small slender tree much like Rhamnus except that the
leaves are opposite or subopposite and the fruit finally baccate,
only 1-2-celled.
Rhamnidiutn elaeocarpum Reiss. in Mart. Fl. Bras. 11,
pt. 1: 94. 1861.
Branchlets opposite, abundantly lenticellate, the tips as petioles
(4-12 mm. long), leaves beneath and cjmies more or less puberulent;
stipules interpetiolar, oblong-acuminate, deciduous; leaves elliptic-
oblong, acute or obtusish, often subcordate at base, usually 7-10
cm. long, 2.5-5 cm. wide, with 8-12 lateral nerves; peduncles 4-8
mm. long, the many-flowered cymes simply dichotomous, finely
pubescent; calyx 4-5 mm. long, subequaling the pedicels, tube
broadly obconic, lobes erect, acute, strongly tubercled within,
carinate; petals bilobed, involute; fruit ellipsoid, 10-12 mm. long,
calyx cup and often stamens persisting. — Forming undergrowth
in dense forest to about 6 meters, the bark coarsely fissured (Wil-
liams, Field Mus. Bot. 15: 299. 1936; see also for wood anatomy).
Illustrated, Reissek, I.e., pi. 31.
San Martin: Near Tarapoto, Williams 6887; 6888 (det. R.
Gross). To Paraguay and Brazil.
6. SCUTIA Comm.
Scypharia Miers, Contr. Bot. 1: 299, pi ^2. 1861, fide Weber-
bauer, Field Mus. Bot. 8: 83. 1930.
Spiny or rarely unarmed shrubs, the branchlets often angled,
the coriaceous leaves opposite or nearly and pinnately nerved, the
5-merous flowers fasciculate or congested in small axillary umbels.
Calyx-tube hemispheric or turbinate, the acute lobes thickened
apically. Petals plane or cucullate, nearly equaled by the stamens.
Flora of Peru 897
Disk with undulate margin. Ovary free, 2-4-celled. Fruit obovoid
or subglobose, dry or scarcely fleshy, with 2-4 pyrenes and basally
enclosed in the calyx-tube. — S. gtiayaquilensis (HBK.) Weberbauer,
while similar to the known Peruvian species, is distinguishable by
the form of the leaves (Weberbauer) ; they are elliptic, rounded but
mucronate at tip, decurrent to short petioles, softly canescent
pubescent beneath.
Flowers sessile S. spicata.
Flowers shortly pedicellate S. paiLciJlora.
Scutia pauciflora (Hook, f.) Weberb. Field Mus. Bot. 8: 84.
1930. Discaria pauciflora Hook. f. Trans. Linn. Soc. 20: 229. 1851.
Scypharia (parviflora) pauciflora (Hook, f.) Miers, Contr. Bot. 1:
301, pi. It2. 1861. D. parviflora Hook. f. ex Miers, I.e.
Terete branches and branchlets spinescent; leaves caducous, few,
oblong-obovate or oblong, mucronate, shortly petioled; flowers
solitary or binate, subsessile; petals very broadly spatulate, bifid;
ovary 2-celled. — Distinguished by the very small flowers and the
bifid petals (Hooker f.).
Peru: (possibly). Galapagos; Ecuador.
Scutia spicata (Willd.) Weberb. Field Mus. Bot. 8: 83. 1930.
CoUelia spicata Willd. ex Roem. & Schult. Syst. 5: 513. 1819. Rham-
ntts senticosa HBK. Nov. Gren. & Sp. 7: 54. 1824. Sageretia senticosa
(HBK.) Brongn. Ann. Sci. Nat. s6r. 1, 10: 360. 1827. Scypharia
senticosa (HBK.) Miers, Contr. Bot. 1: 301, pi. U2. 1861. Scutia
maritima Perk. Bot. Jahrb. 45: 464. 1911.
Glabrous, the many terete or sub-4-gonous green spiny branchlets
subopposite the opposite subulate spreading spines 2.5-6 cm. long;
stipules minute, ovate-subulate; leaves few, solitary at base of spines
(petioles canaliculate, articulate to base, 2 mm. long), ovate, obtuse,
rounded at base, sub-5-plinerved, subcoriaceous, glaucous-green,
about 2.5 cm. long; flowers minute, 1-2 or 5-6-fasciculate-glomerate,
sessile; calyx subhemispheric, medially 5-parted, the ovate acute
spreading segments valvate before an thesis; p)etals 5, scale-like,
emarginate-bilobed, included, subcucullate; anthers 2-celled, affixed
dorsally above base; ovary depressed-globose, 3-celled, the cells
1-ovuled; style very short, the 3 stigmas obtuse; disk thin, seeming
obsolete; fruit globose, 1-3-seeded, the seeds lenticular. — After
HBK. This, as the related S. arenicola (Casar.) Reiss., has thinner
disk than in S. Imxifolia Reiss. (Weberbauer). The stout green
398 Field Museum of Natural History— Botany, Vol. XIII
thorns, often 6 cm. long, make passage through thickets of these
shrubs impossible; the small red fleshy fruits are sour but very-
palatable, according to Svenson, who gives a habital photograph,
Amer. Joum. Bot. 33: 396. 1946. Attains about 3 meters often
in stands alone, and characteristic for the dry coast land from
southern Ecuador to region of Chala, southern Peru, and reaching
at most 1,600 (1,800, Mexia) meters, this in central Peru (Weber-
bauer). Illustrated, Miers, Contr. Bot. 1: pi. U2 C. F.M. Neg.
9489.
Tumbez: Coastal plain, Weberhauer 771^. — Piura: Quebrada
Mogollon, Amotape Hills, abundant (Haught & Svenson 115^0).
Contumasay, Prov. Truxillo, Bonpland (type, R. senticosa). Ni-
gritos, Haught 68. — Libertad: La Goldina, Prov. Trujillo, Worth
9056 (det. Johnston). — Ancash: Yautan, 2561. — Lima: Chosica to
Matucana, Mexia 0U076. Above San Bartolom^, Weberhauer 5207.
Prov. Chancay, Goodspeed 1735 Jt. Prov. Huarochiri, Goodspeed
Exped. 30219 (det. Killip). Rio de Lomas, Weberhauer 5739. "Mo-
lono" (Bonpland), "Hpe," "muchilco" (both Weberhauer). Gala-
pagos; Ecuador.
7. AMPELOZIZYPHUS Ducke
Robust liana with alternate entire 5-nerved leaves, the 2 outer
nerves slender or obscure, and hermaphrodite flowers in axillary
cymes mostly on the uppermost leafless branches forming elongate
racemes or in ample panicles. Stipules caducous. Calyx-tube
shortly turbinate, the five 1-nerved apically callosed lobes spreading,
subequaled by the long-clawed petals, these inserted with the
shorter stamens at the margins of the adnate plane entire disk.
Ovary 3-celled, completely connate with calyx-tube and disk, the
solitary ovules erect. Capsules stiped by the stout torus the more
or less refiexed calyx-lobes persisting, 3-seeded, the seeds exal-
buminous, thus distinct from Coluhrina the capsules at maturity
dehiscent elastically (Ducke). Immature fruits suggest drupes,
perhaps evidence of the tenuous character of the taxonomy within
the family.
Ampelozizyphus amazonicus Ducke, Arch. Inst. Biol. Veg.
2: 158. 1935.
Glabrous except reddish-pubescent growing parts including
inflorescences; reddish bark of branchlets laminulately deciduous;
petioles to 2.5 cm. long; leaves ovate-elliptic, rounded or obtuse
Flora of Peru 899
at base, shortly acuminate or obtuse, coriaceous, 1.5-2.5 dm. long,
7-12 cm. wide, or usually much smaller on fertile branchlets;
peduncles to 2 cm. long; pedicels 2-3 mm. long; calyx 2 mm. long,
the lobes tomentulose within; fruits glabrous, obovoid. — Inflores-
cence and leaves suggest certain Menispermaceae; outer bark has
odor of methyl salicylate, as that of Pourouma, some Polygalas and
Parkia oppositifolia (Ducke). Capsules depressed-trigonous,
strongly callose-carinate, to 2 cm. high, 3 cm. wide; endocarp
crustaceous (Ducke, I.e. 4: 47. 1938). Illustrated, Ducke, I.e.
pis. 1, 2, opposite 172 and I.e. 4: 47. pi. S (fruit).
Loreto: Mishuyacu near Iquitos, Klug J^IO; 566; 589. Amazon-
ian Brazil.
"Saracura-mira" (Ducke).
8. COLUBRINA Richard
Cormonema Reissek in Mart. Fl. Bras. 11, pt. 1: 96. 1861. Caesia
Veil. Fl. Flum. 3: pi. 23, Text 107. 1825.
Trees or shrubs, sometimes spinescent, the alternate or opposite,
entire or dentate, pinnate-nerved leaves sometimes biglandular at
petiole apex or base of blade. Flowers 5-merous, in axillary fascicles
or umbelliform cymes. Cal5rx-tube hemispheric, the spreading lobes
often 3-angled and carinate within. Persisting petals clawed, in-
serted below the fleshy angulate or lobate disk which lines tube and
surrounds the free globose 3-celled ovary. Drupe finally capsular-
baccate, the epicarp thin, the carpels within crustaceous or with
a hyaline membrane; seeds obovoid. — Reissek separated as a distinct
genus species with leaf-glands and, especially, the carpels with a
hyaline membrane within; it is questionable if the characters are
concomitant and in any case it is practical taxonomy to treat them
as indicating sectional division under one name.
Mention may be made of C. Spriicei (Suesseng.) Cowan, Brit-
tonia 7: 405. 1952 {Cormonema Spriicei Suesseng. Bot. Archiv
[Konigsberg, etc.l 39: 387. 1938) from Rio Solim5es, Brazil, which
according to the author is similar to Cormonema ovalifolia Donn.
Sm. of Central America, differing in having darker lustrous leaves
with more acute tips. It has been most agreeable to find a student
supporting my union of two genera, in this instance in manuscript.
Leaves opposite or subaltemate, the glands at base of blade.
C. glandulosa.
Leaves alternate, the glands usually at apex of petiole. .C. VeUozoi.
400 Field Museum of Natural History — Botany, Vol. XIII
Colubrina glandulosa Perk. Bot. Jahrb. 45: 465. 1911.
Unarmed tree, the younger branches and leaves beneath rufo-
tomentose, glabrescent or finally glabrous except for the tomentose-
pilose cymes and calyces; petioles 1-1.5 cm. long, leaves opposite
or sometimes subaltemate, oblong-ovate or oblong, rounded at the
biglandular (rarely uniglandular) base, obtusely acuminate, 7.5-17.5
cm. long, 3-8 cm. wide, entire, chartaceous, lateral veins 4-6, these
often with a few minute trichomes beneath; flowers 4 mm. long;
pedicels 1-2 mm. long; calyx-lobes acuminate, spreading, callose-
tipped; fruit subglobose, 6 mm. in diameter, the 3 seeds obovoid.
— Allied by the author to C. rufa Reiss. of Brazil but distinct by the
glabrous glandular-based leaves. To 10 meters high (Klug). F.M.
Neg. 5846.
San Martin: Juanjui, Klug JtSJfO (det. Standley). — Huanuco:
Between Monzon and Rio Huallaga, 600 meters, Weherhauer 362S,
type.— Rio Acre: Ule 9633 (det. Pilger).
Colubrina Vellozoi Cowan, Brittonia 7: 405. 1952. Cormo-
nema spinosum (Veil.) Reiss. in Mart. Fl. Bras. 11, pt. 1: 96. 1861,
not Colubrina spinosa Donn.-Sm., 1897. Caesia spinosa Veil. Fl.
Flum. 3: pi. 23, Text 107. 1825.
Much branched, often armed, the tips of the short or long
branchlets typically pubescent; petioles short or to 1 cm. long;
leaves lanceolate to oblong- or ovate-elliptic, puberulent on nerves,
finally glabrous, biglandular at apex of petioles or very base of
blade, usually chartaceous, often several cm. wide and about twice
as long, obtuse to acuminate; fascicles ordinarily many-flowered,
the pedicels at last 4 mm. long; calyx spreading, 3 mm. broad, the
lobes callose- tipped ; fruit spherical, the crustaceous 3 carpels or
cocci 1-seeded and with a very thin subhyaline inner membrane.
— The var. peruviana Macbr. ex Cowan, I.e. (Cormonema spinosum
var. peruvianum Macbr. Field Mus. Bot. 8: 124. 1930) has cupuliform
instead of patuliform leaf-glands, spines when present mostly about
2 cm. long, leaves subcoriaceous. To 7 meters tall. The species may
not be specifically distinct from C. heteronema (Griseb.) Standley,
known from Panama to Mexico.
San Martin: Chazuta, Klug ItlU2; ^l^S (det. Standley).— Junln:
La Merced, 5368 (type, var.); 526 J^; Killip & Smith 23530. Brazil.
9. RHAMNUS L.
Unarmed with usually alternate pinnately nerved leaves, small
deciduous stipules and axillary racemose or fasciculately cymose
Flora of Peru 401
4-5-merous flowers. Calyx tube after anthesis medially circum-
scissile the lower portion urceolate, persisting but free about the
base of the baccate drupe, the 3-angled lobes carinate within, erect
or spreading. Petals rarely none, sometimes plane. Disk plane,
the subsessile stamens inserted below its thin margin. Ovary free,
3-4-celled, the fruit with usually 2-4 scarcely or not dehiscent
pyrenes.
Branchlets granulately lenticellate; leaves soon glabrous, subentire.
R. granulosus.
Branchlets smooth, sparsely if at all lenticellate.
Leaves serrulate, firm R. Jelskii.
Leaves crenulate, membranous R. riojae.
Rhamnus granulosus (R. & P.) Weberb. Pflanzenfam. 3,
Abt. 5: 410. 1895. Ceanothus granulosus R. & P. Fl. Peruv. 3: 5,
pi. 228. 1802.
Densely leafy shrub or small tree, the many branchlets con-
spicuously lenticellate, the tips, younger petioles (1-1.5 cm. long)
and inflorescences early rusty villous-hispidulous except the last
soon glabrous; leaves oblong-elliptic or slightly obovate, shortly
acute at base, shortly acuminate, usually about 9 cm. long, 5 cm.
wide, or to 15 cm. long, 7 cm. wide, subentire or above the middle
obscurely and repandly mucro-denticulate, lustrous especially above,
glabrous or early more or less pubescent beneath along the nerves
or in their axils, soon coriaceous; peduncles to about 1.5 cm. long;
pedicels in fruit to 5 or 6 mm. long, umbellately several; calyx lobes
scarcely 2 mm. long, ovate, acutish; petals obovate, clawed, included;
disk obscure; ovary subrotund-turbinate, 3-celled; fruits sub-
spheroid, 5 mm. thick, sparsely appressed puberulent-hispidulous,
the seeds obovoid. — Tjrpe collections from Chinchao, Cuchero and
Vitoc, Ruiz & Pavdn. To 8 meters tall (Weberbauer).
Hudnuco: Pampayacu, Kanehira. Cuchero, Poeppig 12SJ^.
Chinchao, Weberbauer 6802 (det. Pilger).
Rhamnus Jelskii Szyszyl. Dissert. Classis Math.-Phys. Acad.
Litt. Cracov. 29: 224. 1895. R. pubescens (R. & P.) Tr. & PI. Ann.
Sci. Nat. s^r. 5, 16: 379. 1872, not Poir. nor Sibth. & Sm. Ceano-
thus pubescens R. & P. Fl. Peruv. 3: 6, pi. 228. 1802.
Shrub-tree, the numerous branchlets only at tips, younger
petioles, these 1-1.5 (2) cm. long and flowering cymes more or less
402 Field Museum of Natural History— Botany, Vol. XIII
canescently rufo-hispidulous or sub villous; leaves oblong-lanceolate
to -elliptic or somewhat ovate, acute to rounded at base, acuminate,
3-12 cm. long, 2-6 cm. wide, glandular-serrulate usually nearly
from base to apex, subcoriaceous in age, lightly pubescent to gla-
brate above and beneath or the subparallel (evenly spaced) nerves
more villous, as sometimes also the fine veins; peduncles about 1
cm. long or much shorter, pedicels 3-7 mm. long, fulvous villosulous
even in fruit; calyx 2.5-3 mm. long, greenish-yellow, the ovate-
lanceolate acute lobes erect; drupes depressed ovoid or subglobular,
lightly pubescent to glabrous, reddish-black, finally 6-8 mm. thick
and nearly as high. — Sometimes 5-8 meters tall and furnishing
a dye ( Weberbauer) . The closely related R. chrysophyllus (Reiss.)
Weberb. (or R. pubescens var. chrysophyllus (Reiss.) Ktze.) has the
leaves densely and apparently fulvous lanate beneath; R. Jelskii,
ex char, seems to be the most pubescent form of the Ruiz and Pavon
species within Peru, and R. chrysophyllus if not specifically distinct
would be of course the earlier name. F.M. Neg. 23297 {R. pu-
bescens).
Piura: Ayavaca, 2,900 meters, Weberbauer 6372. — Cajamarca:
Tambillo {Jelski 32^, type). Cutervo, Raimondi. Chota, Weber-
bauer U220. — Amazonas: Chachapoyas, Williams 7556 (det. Gross);
Mathews 767 (Mitten Herb.). Pariahuanca, Weberbauer 6593. —
Junin: Chacahuasi, Ruiz & Pavdn (type, R. pubescens). Panti,
Weberbauer 6593. Vitoc, Ruiz & Pavdn (det. Mansfeld). — Hudnuco:
Cani near Mito, 2,800 meters, 3J^37 (det. Standley). Chinchao,
Weberbauer 6820. — Huancavelica: Mantaro Valley, Weberbauer 6502.
Surcubamba, 2,600 meters, in bushwood. Stork & Horton 10352.
"Aravisa" (Raimondi).
Rhamnus riojae Perk. Bot. Jahrb. 45: 465. 1911.
Branchlet tips, petioles (5-10 mm. long) and cymes including
the calyces fulvous- tomentulose, the former soon glabrescent;
leaves oblong or obovate-oblong, cuneate at base, acuminate,
papyraceous or chartaceous, very sparsely fulvo-pilose on nerves
above, manifestly on the arcuate unevenly spaced nerves and veins
beneath, minutely glandular-serrulate, often 6.5-14 cm. long, 2-4
cm. wide; peduncles 1-10 mm. long, cymes 1-1.5 cm. long; flowers
3.5 mm. long, greenish-white; pedicels 5-8 mm. long; calyx-lobes
acute; ovary 3-celled, glabrous; drupes depressed globose, about
6 mm. in diameter. — In savana woods, to 2 meters tall; perhaps
not consistently distinguishable from R. Jelskii. F.M. Neg. 4697.
Flora of Peru 403
San Martin: Mayo near Tarapoto, Spruce 1^877 (det. Reissek,
n. sp. ined.)- — Loreto: Rioja, west of Moyobamba, Weherhauer
A697, type. Pumayacu, Balsapuerto to Moyobamba, Kltig SI 81
(det. Standley, R. pnbescens).
10. COLLETIA Comm.
Often leafless shrubs, the decussately opposite branches spinose,
notably compressed and sometimes enlarged at the nodes, these not
articulate. Flowers 4-6-merous (petals may be lacking), fascicled or
solitary below the spines, the 1-flowered pedicels nutant. Calyx
membranous, the tube produced above the adnate disk. Ovary
3-celled. Drupes finally capsular, coriaceous with 3 crustaceous
bivalved cocci. — For a discussion of the morphology of this and
related genera and a taxonomic review of Colletia see Miers, Contr.
Bot. 1: 230-304. 1851-61. A French botanist, D. Collet of about
1700, is remembered by the name.
Ck>lletia spinosissima Gmelin, Syst. Nat. ed. 13, 2: 408. 1791.
C. spinosa Lam. Illus. 2: 91, pi. 129. 1797. C. horrida Willd. Sp.
PI. 1: 1113. 1798. C. polyacantha Willd. ex Roem. & Schult. Syst.
5: 513. 1819. C. WeddeUiana, C. adculata, and C. Kunthiana Miers,
Contr. Bot. 1: 257, 263. 1851-1861. C. Ephedra Vent. Choix 11:
pi. 16. 1803. C. adculata Miers, Ann. & Mag. Nat. Hist. ser. 3,
5: 214. 1860. C. obcordata Vent. Jard. Cels pi. 92. 1800.
Much branched, to 1 meter tall, nearly leafless, the branches
and branchlets decussately opposite, pale green, hirtellous, apically
spinescent with subulate homy point; leaves opposite, subsessile,
obovate-lanceolate or spatulate, obtuse, concave, carinate beneath,
glabrous; stipules 2, ovate, caducous; peduncles 1-2 (3^) from
axillary tubercles, 1-flowered; flowers pendent, yellowish, only 1 in
3 or 4 fertile; calyx campanulate, 10-nerved, membranous, glabrous,
the acute oblong-lanceolate segments reflexing; disk fleshy, entire;
petals none; stamens exserted, anthers reniform, dorsiflexed; ovary
short-ovoid, sessile, 3-celled; style little exceeding the stamens, the
stigma subcapitate; capsules subglobose, glabrous, peduncles 4 mm.
long, cocci rounded, chartaceous, seeds subellipsoid, lustrous, 4 mm.
long. — After HBK. and thus Miers' C. Kunthiana. Calyx greenish-
yellow; anthers black (Stork & Horton). Miers, Contr. Bot. 1:
253. 1851-1861, considers the plant of Commerson from Buenos
Ayres as the type, which however was figured from a specimen by
Jos. de Jussieu from Peru; Lamarck considered them the same.
404 Field Museum of Natural History— Botany, Vol. XIII
Miers furthermore proposed C. aciculata for the Peruvian plant,
and also several other names; types not seen but from accumulated
materials and descriptions it is not evident that they represent
more than vegetative variations, the characters apparently being
relative and scarcely concomitant or constant; the problem however
is for a student of the group.
Contains colletin that seems to possess tonic properties; bark
and young branches serve as an efficacious aid for soap, and entire
plant is an excellent fuel for baking ovens (Herrera). Illustrated,
Minist. Agric. y Ganad. Argent. 7, fasc. 120, fig. 9. F.M. Negs. 9490
(C. horrida); 35969 (C. Weddelliana).
Piura: Huancabamba, on the paramo (Bonpland, type, C.
horrida) . Pampano above Pisco, Weberbauer 5370. — Ancash : Casma,
Raimondi. Huardz, gravelly river bluff, 2528. Llata, shores of
Rio Maraiion, 2280. — Libertad: Prov. Pacasmayo, Raimondi. — â–
Lima: Above San Bartolom^, Weberbauer 5207. Above Lima,
Raimondi. — Junin: Tarma to Palca, Weberbauer 1727 (det. Perkins);
176; Ruiz & Pavdn; Dombey; Martinet. Near Huancayo, Killip &
Smith 22030. — Arequipa: Misti Volcano, Weberbauer 4^835. — Huan-
cavelica: Gravelly hills. Pampas, Stork & Horton 10238. Lake
Titicaca (Weddell If391, type, C. Weddelliana). — Apurlmac: Trail to
Abancay, West 37 UU (det. Johnston, C. Weddelliana). — Cuzco:
Valle del Huatanay, 3,200-3,600 meters, Herrera 6^,0. Near Cuzco,
Weberbauer U893. Ollantaytambo, Cook & Gilbert 416; 1936. Chas-
pyoc, Huarocondo Rio, Edmund Heller 2170. — Puno: Salcedo,
Soukup ^66. "Ccacara" (West), "Hague" (Ruiz & Pavon), "zarza
de moyse" (Bonpland), "naqui" (Dombey), "yaquil" (Dombey),
"roqque." To Chile, Uruguay, Argentina.
11. GOUANIA L.
Cirriferous often high-climbing shrubs with alternate leaves and
polygamous 5-merous flowers in terminal or axillary spikes or
racemes, the branches or branchlets often produced as a tendril.
Disk filling short-obconic calyx-tube, 5-gonous or 5-comute. Ovary
inferior, 5-celled, style 3-branched. Fruit coriaceous, crowned by
the persisting calyx, usually 3-winged the 3 subligneous cocci in-
dehiscent, separating in age from the axis.
At least one species, G. lupuloides, is well known in many places
as a dentifrice, while an infusion of the stems is said to furnish a
pleasant bitter in the manner of hops and thus the common English
name of "chew-stick."
Flora of Peru 405
The need for revision of the genus is great; the Peruvian plants
may be referable to one or more of the Brazilian species if Reissek's
work in Mart. Fl. Bras. 11, pt. 1: 101-111. 1861 is sound; on the
other hand the characters seem intangible that define these from
the older widely distributed forms. Without a modem revision
available, expediency has determined the key characters which may
or may not prove to indicate specific values when the plants are
completely known.
Leaves soon glabrous beneath unless on the nerves.
Leaf glands of teeth lacking or minute G. lupuloides.
Leaf glands obvious.
Glands unless in age closed and tipped with trichomes.
G. trichodonta.
Glands glabrous, mostly open, patelliform or cupulate.
G. adenophora.
Leaves pubescent beneath even when mature.
Fruits winged; callus of leaf-teeth closed or nearly. .G. polygama.
Fruits wingless; callus of leaf-teeth more or less patelliform.
G. aptera.
Gouania adenophora Pilger, Notizbl. Bot. Gart. Berlin 6:
314. 1915.
Scandent, sparsely cirriferous, glabrescent except new branchlets,
axils of the spiciform inflorescences, these to 18 cm. long, and the
slightly villous leaf-nerves on both sides; petioles to 1.5 cm. long;
leaves ovate- or oval-elliptic, rounded and often a little inequilateral
at base, shortly acuminate, acute, to 8 cm. long, 4-4.5 cm. wide,
coarsely and irregularly serrate-dentate or undulate, the teeth bear-
ing large cupulate glands, 6-7 nerves prominent only beneath; flowers
canescent villous, 5-merous; sepals 1 mm. long, about equaled by the
cucullate, scarcely clawed petals; disk glabrous. — Outstanding by
the large glands of the leaf-margins, comparable otherwise to G.
trichodonta (Pilger). Quite possibly a part of G. lupuloides. G.
acreana Pilger, I.e., to which Tessman 3805 was referred, seems also
to be a variant, the many spiciform racemes forming an efoliate
panicle. For that matter all of the following material has been
determined as G. lupuloides or G. domingensis var. pubescens (Poir.)
Ktze. or var. heterocarpa Ktze. F.M. Negs 5853; 5852 (G. acreana).
San Martin: Juanjui, Klu^ ItSll. — Junin: Puerto Yessup, Killip
& Smith 2622U; 26272.— Loreto: Yurimaguas, Williams U981;
406 Field Museum of Natural History— Botany, Vol. XIII
Killip & Smith 28059. Iquitos, Ule 16 pt., type; Killip & Smith
27107. Mishuyacu, Klug 163; 36 U; 52U; 600; 716; 1392; 1517.
Pebas, Williams 16 UO. Pucallpa, Soukup 3067.
Gouania aptera DC. Prodr. 2: 39. 1825. G. alnifolia Reiss. in
Mart. Fl. Bras. 11, pt. 1: 106. 1861? G. aptera Poepp. mss. ex DC.
fide Reissek.
Younger branchlets apically subtomentulose; tendrils at tip
or at base of spikes; petioles 6-10 mm. long; leaves subcordate or
ovate-elliptic to elliptic-subrotund, subacuminate or acute, glandu-
lar-crenate with large patelliform glands, 5-7.5 cm. long, 2.5-5 cm.
wide, early densely pubescent and tomentulose especially on the
6-7 lateral nerves, in age ferrugineous-puberulous on both sides;
spikes interrupted, slender, subracemose; calyx tomentose-fur-
furaceous, 3 mm. wide, longer than pedicels; disk glabrous, the lobes
little shorter than calyx teeth; style puberulent; petals cucullate.
— Scarcely distinguishable from G. polygama, sens. lat. unless the
fruits (unknown) are actually not alate; however, G. lupuloides has
a variety with fruits obscurely or not alate, as also G. adenophora.
Illustrated, Reissek, I.e. pi. 26, jig. 7 (leaf). F.M. Negs. 7015; 5854
(ined. name).
San Martin: Tocache, Poeppig 1976, type. Vitoc, Ruiz &
Pav6n. Chazuta, Klug Jf.129 (det. Standley, G. lupuloides). — Junin:
La Merced, Killip & Smith 23It98; 2JtOA6. Warm America.
Gouania lupuloides (L.) Urban, Symb. Ant. 4: 378. 1910.
Banisteria lupuloides L. Sp. PI. 427. 1753. Rhamnus domingensis
Jacq. Enum. PI. Carib. 17. 1760. G. domingensis (Jacq.) L. Sp.
PL ed. 2. 1663. 1763.
Sprawling or trailing over shrubs or trees, typically glabrous or
glabrescent except the minutely pubescent inflorescence; petioles
much shorter than the ovate to elliptic leaf-blades, these usually
shortly and obtusely acuminate, rounded or subcordate at base,
more or less crenate-serrate, often obscurely and distantly, the
marginal glands lacking or minute; flowers yellowish-green, the
pubescent calyx 1.5-2 mm. long including the 1 mm. long lobes
which are equaled by the petals; capsule wings 8-9 mm. broad, the
axis about 5 mm. high, the seed 2.5-3 mm. long. — After Fawcett
and Rendle. Perhaps not in Peru in typical form but understanding
of the group to which it belongs needs monographic work with
modern methods. G. domingensis var. heterocarpa Ktze. with fruits
Flora of Peru 407
obscurely if at all alate seems to be represented by Williams 80Jt2.
Schunke 27U may be G. Ulei Pilger if that is distinct. Illustrated,
Fawcett & Rendle, Fl. Jamaica 5: 73.
Loreto: Florida, Rfo Putumayo, Klug 2361 (det. Standley).
Mouth of Rio Santiago, Tessmann J^53S; U682. Iquitos, Williams
80Jt2; Mexia 6S92 (det. Standley). Rio Mazdn, Josi Schunke 27 j^.
"Chirapasacha" (Schunke). To Mexico; West Indies.
Gouania polygama (Jacq.) Urban, Symb. Ant. 4: 378. 1910.
Rhamnus polygamus Jacq. Enum. PI. Carib. 17. 1760. G. tomentosa
Jacq. Sel. Stirp. Amer. Hist. 263. 1763.
Upper scandent or sprawling branches, leaves beneath and
calyces more or less tomentose, the leaves sometimes sparsely or
pilose; petioles usually 1-1.5 cm. long or the lower to 2.5 cm.;
leaves ovate, the upper rounded or subtruncate or cordulate at
base, mostly 5-8 cm. long, 3.5-6 cm. wide, the lower broadly ovate,
to 1.5 dm. long, two- thirds as wide, cordate at base, all shortly
acuminate, the tip itself obtusish, mucronate; serrations uneven,
often serrate and rather obscure, the glands or calluses ordinarily
closed ; racemes in the upper axils and terminal often forming ample
panicles; caljrx 1.5 mm. long, the pedicels in fruit scarcely as long;
fruits to 14 mm. broad, the axis 3-4 mm. high, glabrate or the body
pubescent, the hard wings subrotund. — It is possible that one or
more of the four herbarium names proposed for some of the fol-
lowing specimens will be found to indicate at least variants but
Urban in herb, referred the two by Reissek and one of the two by
Ruiz & Pav6n to G. tomentosa as a variant. F.M. Neg. 5857 (ined.
name).
Hudnuco: Pampayacu, Rio Chinchao, 5121. Pozuzo, j^655;
Ruiz & Pav6n.—Junin: La Merced, 5233; 5236. To Mexico; West
Indies.
Gouania trichodonta Reiss. in Mart. Fl. Bras. 11, pt. 1:
108. 1861.
Habit of G. aptera branchlets ferrugineous pubescent especi-
ally on the angles; leaves broadly elliptic or subovate-elliptic,
acuminate, dentate-crenate, 5-7 cm. long, 3-5 cm. wide, the glandu-
lar teeth minutely fasciculate-pubescent, early pubescent on both
sides, finally glabrous above, lateral nerves 6, veins obscure; spikes
densely flowered; calyx appressed pubescent, 2 mm. long, little
exceeding pedicel; disk elevated, the lobes nearly two times shorter
408 Field Museum of Natural History — Botany, Vol. XIII
than calyx segments. — Kuntze referred this to G. domingensis var.
pubescens (Poir.) Ktze., that is to G. lupuloides, and indeed there
seem to be specimens intermediate in character. Illustrated, Reissek
I.e., pi 26, jig. 3 (leaf). F.M. Negs. 32601; 23291 (as G. alnifolia).
Loreto: Yurimaguas, Poeppig 3078, type. Lower Rio Nanay,
Williams 358 (det. R. Gross). Mishuyacu near Iquitos, Klug 13 5 U
(det. Standley). "Granadilla" (Williams). Bolivia.
VITACEAE. Grape Family
Reference: Planchon in DC. Monogr. Phan. 5. 1887.
Scandent (Peru) mostly by tendrils, the scabrous nodose or
articulate branchlets soon or tardily ligneous, the sap watery.
Leaves alternate (unless the lower), entire to variously lobed or
foliolate, usually palmately or pinnately compound, pellucid-
punctate dots frequently present, always with petiole articulate
at base and often dilated with 2 free stipules. Flowers small, com-
monly 4-5-merous, hermaphrodite or unisexual, variously borne but
often in a compound thyrse, the peduncles ordinarily cirriferous and
opposite the leaves. Calyx cupuliform, sometimes entire. Stamens
opposite the caducous free or calyptrately united petals, included
at the base of the disk, this various in form, mostly intrastaminal.
Ovary cells 2-6, 1-2-ovuled; style short, slender or none. Fruit
baccate, sometimes only 1-2-celled and more or less juicy-pulpous
with 1-4 seeds.
Besides "uva" the grape, and the beneficent beverages and other
products it yields, the family is well known especially in northern
climes for ornamental vines, as Virginia Creeper (Parthenocissus
tricuspidata (Sieb. & Zucc.) Planchon, and is often referred to as
the Vine Family.
The Peruvian species are here regarded as belonging to a genus
distinct from Vitis, the latter principally in temperate regions and
having the petals united and calyptrately deciduous.
CISSUS L.
Character of the family but the cymose corymbose flowers
always 4-parted, hermaphrodite, the petals free or early lightly
cohering. Disk cupulate, 4-lobate, adnate to the base of the 2-celled
ovary. Fruit 1-2-seeded, not edible.
Leaves all simple C. sicyoides.
Leaves compound unless the uppermost.
Flora of Peru 409
Leaflets entire or serrulate.
Leaves in part 5-foliolate, the leaflets equally cuneate to base.
C. granulosa.
Leaves 3-foliolate, the lateral obliquely rounded at base.
Stems sometimes alulate but not at all crenulately.
Leaflets sessile or the terminal cuneately petiolulate,
obovate-lanceolate, usually obtuse or acute . . C. erosa.
Leaflets often all well-petiolulate, acuminate, the terminal
rhombic *. C. rhombifolia.
Stems strongly crenulate-alate C. vlmifolia.
Leaflets irregularly lobed or lobate C gongylodes.
Cissus erosa Richard, Act. Soc. Hist. Nat. Paris 1: 106. 1792;
548. C. salutaris HBK. Nov. Gen. & Sp. 5: 225. 1822. C. quadria-
alata HBK. I.e. Vitis salutaris (HBK.) Baker in Mart. Fl. Bras. 14,
pt. 2: 211, pi 52. 1871. V. erosa (Richard) Baker, I.e. 210.
Typically glabrous except the somewhat strigose usually long-
ped uncled inflorescences, the slender stems acutely tetragonous-
sulcate or narrowly alate; leaflets 3, subsessile or the terminal
cuneately petiolulate, the lateral oblique at base, all oblong- or
ovate-lanceolate, obtuse or acutish or even shortly acuminate,
mucronulately crenate-serrulate or subentire, drying firm-charta-
ceous, lustrous, somewhat rufescent, the rather few nerves and many
reticulate veins prominent; inflorescences including the peduncles
crimson, or the flowers rarely white; fruits globose-ovoid, about 6
mm. long. — The var. salutaris (HBK.) Planchon, I.e. is more or less
hirtellous, especially the leaf-nerves, and apparently often with
stouter stems; the leaflets are obtuse as in the type by Leblond from
French Guiana, or acute. The interpretation of the species may be
open to question. Illustrated, Mart. Fl. Bras. 14, pt. 2: pi. 52.
F.M. Neg. 23806.
San Martin: Juanjuf, Klu^ ^258 (det. Standley, C. rhombifolia).
Tocache, Poeppig (det. Herb. Wien, C quadrialata) . — Hudnuco:
Below Tingo Maria, Stork & Horton 951tO (det. Standley, C. salutaris).
— Junin: Satipo, Soukup 28Jt9. — Loreto: Florida, Rio Putumayo,
Klug 2092. Rio Mazdn, JosS Schunke 6U (var., det. Standley). —
Cuzco; Gay (det. Planchon). — Ayacucho: Near Rio Apurlmac,
Weherhauer 56SU. "Navarria" (Schunke). To Mexico and the
West Indies.
410 Field Museum of Natural History — Botany, Vol. XIII
Cissus gongylodes (Baker) Burchell ex Planchon in DC.
Monogr. Phan. 5: 550. 1887. Vitis pterophora Baker in Mart. Fl.
Bras. 14, pt. 2: 213. 1871. V. gongylodes Baker, I.e. 209.
Tetragonous lower stems extraordinarily crenulate-alate or the
angles of the wings deeply crisped; leaflets 3 with one or more angled
lobes, the subsessile intermediate often trilobed, all rhomboid,
puberulent-pilose on the nerves, the lateral sessile or petiolulate
(Peru); cymes many-flowered, short-corymbiform; corolla depressed
globose, glabrous, th^ reddish-brown petals finally expanding. — The
Peruvian plant referred here by Baker was given an herbarium name
by Poeppig and seems to be at least a variant, since the lateral
leaflets are long- (1.5 cm.) petiolulate; it may become var. lobata
[Poeppig] Macbr., var. nov., foliolis lateralibus longe petiolulatis.
C. spinosa Camb. 549, related but lightly lobate, might be repre-
sented by my ^706 from Pozuzo but the incomplete material shows
only spine bases; the obovoid fruits are 12 mm. long, nearly 10 mm.
thick at apex. Illustrated, Pflanzenfam. ed. 2. 20D: 274. fig. 80.
Loreto: Yurimaguas, Poeppig 2273 (type, var. lobata, herb.
Vienna). Brazil.
Cissus granulosa R. & P. Fl. Peruv. 1: 64, pi 101. 1798; 555.
Among Peruvian species well-marked by the mostly 5-foliolate
leaves and the usually conspicuously granulate-tuberculate older
branches, the smooth younger obscurely 4-margined; leaflets sessile
or subsessile, cuneate-obovate, apiculate at the rounded or barely
acute tip, minutely serrulate, fleshy, 4-7 cm. long, 2-2.5 cm. wide,
finely reticulate- veined beneath; cymes dichotomously divided,
sometimes ample, often shorter than the leaves; calyx repandly
lobed; petals ovate-oblong, finally spreading, about 3 mm. long;
fruit globose, to nearly 10 mm. in diameter. — Completely glabrous
unless for an obscure puberulence on the slender pedicels. My
collections on low sunny shrubs or trailing on slide rock. F.M. Negs.
9780; 18235.
Hudnuco: Yanano, 3726; 3781. — Junin: Huasahuasi, Ruiz &
Pavdn, type. — Ayacucho: Yanamonte, 2,700 meters, Weberhauer
56J^6. — Loreto: Yurimaguas, Poeppig 2j^7.
Cissus rhombifolia Vahl, Eclog. Amer. 2: 10. 1798; 544. C.
obliqua R. & P. Fl. Peruv. 1: 65, pi. 101. 1798. Vitis rhombifolia
(Vahl) Baker in Mart. Fl. Bras. 14, pt. 2: 207. 1871.
Flora of Peru 411
Much like C. erosa but the stems striate-sulcate, not alate, and,
especially the leaflets often all abruptly long-petiolulate, or maybe
typically the lateral subsessile, more or less acutely acuminate, often
reddish hirtellous especially on the rather prominent nerves be-
neath, the reticulate venation much less marked, sometimes gla-
brescent; flowers umbellulately and densely congested, typically
pubescent including the red petals. — C. microcarpa Vahl, 546, to be
expected, seems to be scarcely distinguishable but is glabrous and
said to have narrowly 4-alate branches, these angled above, the
terminal leaflet long-, the lateral short-petiolulate. Type of C.
obliqua with all leaflets long-petiolulate, maybe a variety, glabrous
except for a few long appressed trichomes, according to Ruiz &
Pavon found at Hudnuco, Tarma, Huariaca, Rondos, Chablan and
Chaucha. F.M. Neg. 18238 (C. obliqua).
Tumbez: East of Hacienda Chicama, Weberbauer 7663. — Caja-
marca: On stone walls, Chota, Stork & Horton 10039 (det. Standley).
— San Martin: Chazuta, Klug Jt053 (det. Standley). — Hudnuco:
Chicoplaya, Ruiz & Pav&n. — ^Junfn: Near Tarma, Ruiz & Pavdn,
type. Near Palca, Stork 10973. — Loreto: Near Iquitos, Klug 11 U6.
Rio Mazdn, Josk Schunke 3JtO (det. Standley). — Cuzco: Prov.
Quispicanchis, Vargas 7762. Machupicchu, Vargas 67UU. "Sapo-
huasco" (Schunke). To Mexico and the West Indies.
Cissus sicyoides L. Syst. Nat. ed. 10: 897. 1759; 521. Vitia
iicyoides Morales in Poey, Repert. Fis.-Nat. Cuba 1: 206. 1866.
C. umbrosa HBK. Nov. Gen. & Sp. 5: 223. 1822. C. canescens Lam.,
lUustr. 1: 331. 1791. C. compressicaulis R. & P. Fl. Peruv. 1: 64,
pi. 100. 1798.
Scandent or creeping, the branches terete or compressed, tuber-
culate or smooth, striate, the leaves cordate-ovate or oblong, some-
times abruptly and cuneately contracted at base, sometimes sub-
hastate, acutely acuminate or obtusish, somewhat denticulate,
rarely incised-lobate, thick-membranous, glabrous or especially
beneath more or less pubescent; cymes corymbiform, shortly
peduncled, umbellately and dichotomously divided, the small
flowers greenish-yellow, white or purplish, the obovoid-globose
fruits 1-seeded. — After Planchon, who interpreted the name as
applicable to a single species highly variable in leaf-serration,
-indument and -form as well as in size of flowers; he has listed
a dozen or so of the scarcely recognizable variations as forms and
among those that are cited as occurring in Peru are formas morifolia
412 Field Museum of Natural History— Botany, Vol. XIII
Planchon, 525, umhrosa (HBK.) Planchon, 530, canescens (Lam.)
Planchon, 531, and compressicaulis (R. & P.) Planchon, 531; the
first is said to have terete glabrous branchlets and short petioles, the
second tetragonous hirtellous branchlets, the third and fourth more
or less canescent pubescent, the latter with more prominent ser-
ration but the character is certainly intangible. However, Svenson,
Amer. Joum. Bot. 33: 463. 1946, observed two distinct forms in
the vicinity of Guayaquil, Ecuador, one apparently forma com-
pressicaulis with thin petals 1.5 mm. long and rotund anthers only
0.5 mm. long, the other quite glabrous, with thickened petals 2 mm.
long and elongate anthers 1 mm. long. Therefore there may be
some genetic characters that will require a revision of the present
understanding of this plant. However, in at least one herbarium
specimen, glabrous or glabrate, the anthers have appeared to be at
least subrotund. Determinations mostly by Standley, only a few
of the many collections cited. As presently understood the plants
vary also in habit and size from vigorous lianas to slender creepers,
depending in part at least on age and habitat, which ranges from
sandy beaches to shrubby thickets and tall forests, the stems how-
ever apparently always flexible and therefore serving as cords and
for baskets. When high-climbing, bundles of long cord-like fibers
are developed and reach finally to the ground where they often
take root; indeed any part of the plant may survive when cut. The
leaves supply a soap-like lather if rubbed in water. — F.M. Negs.
23808 (C. compressicaulis) ; 35989 (C. umhrosa) .
Tumbez: Southeast of Hacienda La Choza, Weherhauer 7703
(f. compressicaulis). — San Martin: San Roque, Williams 7J^73.
Lamas, Williams 6339. Tarapoto, Williams 5U0; 5853; 6172.
Juan Guerra, Williams 6839. — Ancash: Santa, Stork & Horton 9155
(det. Johnston). — Hudnuco: Chacahuasi, Vitoc, Ruiz & Pav6n
(type, f. compressicaulis). Churuplaya, Mexia 8253 (f. compressi-
caulis). Tingo Maria, Stork & Horton 9Jt65; Allard 20373 (det.
L. Smith). Pampayacu, Poeppig 2126. Pillao, Ruiz & Pavdn
(forma). — Lima: Near Lima, Dombey; Jussieu; Gaudichaud (f.
canescens). Chancay, Ruiz & Pavdn (type, C. compressicaulis).
Callao, Soukup 213^ (f. compressicaulis). — Junin: La Merced, 5255;
5560; Killip & Smith 23694- (f. morifolia). — Lore to: Lower Rio
Nanay, Williams 256; 289; 458. Iquitos, Williams 8001; 3531;
7906; 1393; Klug 1513; 151 A. Rio Paranapura, Klug 3938. Pu-
mayacu, Klug 3229. Creek Carapisa above Pongo de Manseriche,
Mexia 6256 (f . umhrosa) ; 61 78; 61 70. Near Yurimaguas, Klug 281 7;
Flora of Peru 418
WUliaitis 5155; It995; U76; U08. Florida, Klug 2066; 2S52. Ca-
ballo-Cocha, Williams 2291; 2Jt28. Leticia, Williams S150.—
Lambayeque: Chiclayo, Stork llltSl; West 3579 (det. Johnston).
— Ayacucho: Near Kimpitiriki, Apurimac Valley, Killip & Smith
2S02^; 230SS. —Cuzco: Valle del Urubamba, 2,200 meters, Herrera
SS12. To Mexico and the West Indies. "Ampato-huasca" (Wil-
liams) ; "zapo-huasca" (Mexia) ; "yedra" (Ruiz & Pav6n) ; "paja de
la culebra" (West).
Cissus ultnifolia (Baker) Planchon in DC. Monogr. Phan. 5:
552. 1887. Vitis ulmifolia Baker in Mart. Fl. Bras. 14, pt. 2: 213.
1871.
Glabrous or essentially except the puberulent peduncles and
pedicels; stems terete, multistriate and with 4 strongly crisped
wings; leaflets 3, coriaceous, lustrous above, reticulate-veined, acute
or obtuse, oblong-elliptic or the terminal subobovate, mostly 8-10
cm. long, about half as wide, minutely serrulate, all well-petiolulate,
rarely sessile; corolla glabrous, — Climbing tangle along bank and
open places, the flowers crimson, mature fruit black (Mexia). F.M.
Neg. 32627.
Loreto: Yurimaguas, Poeppig Addenda 22, type. Mishuyacu,
KliLQ 929. Above Pongo de Manseriche, Mexia 6317 (det. Standley).
"Sapohuasco Colorado" (Mexia).
TILIACEAE. Linden Family
References: Burret, Notizbl. Bot. Gart. Berlin 9: 592-880. 1926;
Schumann in Mart. Fl. Bras. 12, pt. 3: 119-200. 1886.
Trees, shrubs or infrequently more or less sufTrutescent herbs
with alternate entire, serrate or rarely lobed often stellate pubescent
leaves. Stipules conspicuous in Vallea. Flowers hermaphrodite or
hermaphrodite and pistillate, sometimes involucrate. Sepals
usually 4 or 5, valvate, the usually imbricate petals as many or
rarely reduced or wanting. Stamens numerous (exceptionally 100),
free or shortly connate, the anthers 2-celled, at least finally opening
longitudinally or apically (Sloanea). Ovary free (rarely inferior),
2-many-celled (rarely 1-celled by abortion), each cell with 2-many
ovules. Style simple or more or less divided at tip; stigma some-
times sessile. Fruit dry, often rough or even spinose, dehiscent or
indehiscent. — R. Weibel, Candollea 10: 155-177. 1945, in his careful
study of the placentation in this family has verified the existence
414 Field Museum of Natural History — Botany, Vol. XIII
of the parietal and concluded that Mollia and Goethalsia are genera
properly placed here.
For convenience the family Elaeocarpaceae, by many students
treated as distinct, is included here; C. Earle Smith, Jr., Contr.
Gray Herb. 175: 3. 1954, summed up the relationship as follows:
the family is a somewhat arbitrary assemblage of genera, mostly
removed from the Tiliaceae because they lack the mucilage ducts
common to the members of that family; some of the characters
suggest affinities with the StercuUaceae and the Malvaceae. Smith
found that about a third of the unidentified specimens of "Sloanea"
examined in herbaria belonged to the Flacourtiaceae; the latter
however have petaliferous flowers and rather thin-walled, usually
many-seeded fruits.
Interesting family as the source of hemp (jute) and other valu-
able fibers (Triumfetta, Cor chorus).
Perennials, often suffrutescent below; fruit a linear capsule; flowers
small, yellow 1. Corchorus.
Shrubs, the stems and branches slender, virgate; fruit a prickly bur;
flowers yellow 2. Triumfetta.
Trees or more or less arborescent shrubs; fruit sometimes armed
but not as above.
Leaves 3 (-7) -nerved from base (including midnerve), sometimes
pinnately nerved above base; anther dehiscence longitudinal
or early by apical chink.
Stipules wanting or small or caducous (subpersisting in Apeiba
tibourbou).
Leaves cordate-ovate, angulate-lobulate; flowers many, small
(except T. calycina) ; fruit finely bristly.
Fruit a bristly bur; flowers 1-2 (3), bracteolate.
2. Triumfetta.
Fruit disciform, bristle fringed; flowers in 3's (2's), often
ebracteolate 3. Heliocarpus.
Leaves entire or serrate; flowers few, medium size; fruits
various, never quite as above.
Flowers not involucrate; capsules alate, tubercled or bristly
(unknown for Neotessmannia).
Leaves slightly if at all oblique; ovary superior, 1-many-
celled.
Flora of Peru 415
Capsules not alate, opening by a central hole or at top
with woody teeth; anthers appendaged . 4. Apeiha.
Capsules alate apically, semi-bivalved; anthers not
appendaged 5. Mollia.
Leaves strongly oblique; ovary inferior, multicelled
below 8. Neotessmannia.
Flowers (each) early enclosed in a parted or calyciform
involucre; capsules ligneous, not alate apically, smooth,
5-valved.
Involucre divided; anthers connate, opening longi-
tudinally 6. Luekea.
Involucre cupulate; anthers divaricate above, early
opening by a chink 7. Lueheopsis.
Stipules conspicuous, persisting (sometimes absent in Vallea on
fertile branches; Apeiha tibourbou has tardily deciduous
stipules).
Fruit dry; stipules subrotund 9. Vallea.
Fruit fleshy; stipules linear 10. Muntingia.
Leaves pinnately nerved from base; anthers often opening by an
apical chink 11. Sloanea.
1. CORCHORUS [Toum.l L.
More or less suffrutescent herbs frequently hirsute with simple
trichomes, rarely stellulate, the leaves serrate, the shortly peduncled
almost subsessile 1-few-flowered bracteate inflorescences opposite
them or in the axils. Sepals and yellow petals 5 (4). Stamens
usually many, all antheriferous. Ovary 2-5-celled, many-ovuled;
style short the dilated stigma crenulate. Capsules elongate-linear
and smooth or rarely subglobose and muricate, 2-5-ovuled, some-
times septate between the numerous pendulous or horizontal seeds.
Weibel, Candollea 10: 173. 1945, confirmed the observation of Payer
that the ovary is 1-celled in bud.
Two similar Asian species (C. clitoritis L. and C. capsularis L.)
are the source of the important fiber, jute.
Sepals at most 4 mm. long; capsules narrowly 3-winged, with 3
finally horizontal apical horns C. aestuans.
Sepals longer than 4 mm.; capsules beaked or 4-apiculate.
Capsules acuminate-beaked; herbs or ligneous toward base.
416 Field Museum of Natural History— Botany, Vol. XIII
Capsules and usually stems hirsute, the trichomes spreading.
C. hirtus.
Capsules and often stems glabrate, the trichomes appressed or
obscure C. orinocensis.
Capsules obtuse but minutely 4-apiculate; shrubby species.
C. siliquosus.
Corchorus aestuans L. Syst. Nat. ed. 10, 1079. 1759. C.
acutangulus Lam. Encycl. 2: 104. 1786.
Annual or perennial herb, the stem sometimes woody at base,
glabrate or pilose; leaves ovate to rounded, more or less acute,
crenate, the 2 lowest serrations now and then bristle-tipped; sepals
3-4 mm. long, cucullate at tip, equaled by the obovate petals;
capsules glabrous, 3-celled, 6-angled with 2 or 3 of the angles winged,
the beak with 3 entire or bifid horns.
Peru (probably). Widely distributed in tropical regions.
Corchorus hirtus L. Sp. PI. ed. 2, 747. 1762. C. pilolohus Link,
Enum. Hort. Berol. 2: 72. 1822.
Similar to C. orinocensis but more or less hirsute with spreading
trichomes this indument extending to the capsules these often
curved near the base and compressed; leaves ovate to lanceolate-
oblong, rarely 5 cm. long, obtuse to acuminate; sepals pilose, 6 mm.
long, the petals about as long. — Many Peruvian specimens referred
here seem to belong rather to the related species which perhaps
should be treated as a variety, as by Schumann.
Lima: Huara, Ruiz & Pav6n (det. Burret). — Loreto: Rio Paran-
apura, Klug 3960, in part. Yurimaguas, Poeppig 2098. American
tropics.
Corchorus orinocensis HBK. Nov. Gen. & Sp. 5: 337. 1823.
C. hirtus L. var. orinocensis (HBK.) Schuman in Mart. Fl. Bras. 12,
pt. 3: 127. 1886. C. mompoxensis HBK. I.e. 339, fide Dahlem Herb.
Usually erect, sometimes 2.5 meters tall, often woody below and
with 1-2 or few branches, sometimes a marsh herb, generally gla-
brate but the stems early lineately puberulent; leaves ovate to
lanceolate, or narrowly elliptic, 3-10 cm. long, acute to acuminate;
stipules filiform; sepals 6 mm. long or longer; ovary 3-celled; capsules
straight or nearly, sparsely appressed puberulent or almost glabrous,
with erect beak and transverse partitions. — Illustrated, Fawcett &
Rendle, Fl. Jam. 5: 86. F.M. Negs. 9493; 35423.
Flora of Peru 417
Piura: Serrdn, Weberbauer 5991. — San Martfn: Waste land,
Tarapoto, Woytkowski S5050 (det. Cuatrecasas) ; WiUiams 5582
(det. C. pilohulus, Standley); Spruce ^89. Pongo de Cainarachi,
Kltig 27S6 (det. Standley). — Libertad: Raimondi (det. Burret). —
Junin: La Merced, 5225. — Loreto: Balsapuerto, Klug 28^1 (det.
Standley, C. pilobulus). Rio Paranapura, Klug S960, part. Yuri-
maguas, WiUiaim U166; U6S; 5011; 50U1. To Texas and the West
Indies. "Espada pichana" (Williams).
Corchorus siliquosus L. Sp. PI. 529. 1753.
A more or less shrubby herb sometimes a meter or two high,
marked in fruit by 2-celled capsules obtuse but minutely apiculate
with 4 teeth; leaves only 1-4 cm. long, puberulent or glabrate, ovate
or oblong-lanceolate, acute or obtuse; sepals linear, about equaled
by the petals, these 5-6 mm. long; capsules glabrous but lineately
puberulent at the edges of the valves, not transversely septate. —
A specimen from near Lima by Ruiz & Pavon referred here by the
collectors was determined at Madrid by Burret as C. hirtus; how-
ever, it is without the fruit, diagnostic, and the species is probably
within Peru.
Peru (see note above). South America to Mexico, Florida and
the West Indies.
2. TRIUMFETTA [Plum.] L.
Reference: Ko Ko Lay, Ann. Mo. Bot. Gard. 37: 315-395. 1950.
Suffrutescent stellate-pubescent (at least some trichomes) tall
herbs or slender shrubs, the usually small yellow or brownish-yellow
flowers axillary and few or fasciculate or sometimes disposed in an
elongate cylindrical inflorescence with or without bract-like leaves.
Sepals 5, as petals, or these rarely reduced or wanting, glandular or
foveolate at base. Stamens ordinarily many above the commonly
5-glandular elevated torus or gonophore that supports the 2-3 (5)-
celled spinulose ovary, each cell with 2 anatropous collateral pendu-
lous ovules; style filiform, the stigma entire or shortly 2-3-parted.
Capsules subgolobose, echinate-spinose, indehiscent or separating
into two or three 2-seeded cocci or finally falsely 5-10-celled, each
cell 1-seeded, exceptionally 1-celled, 1-seeded by abortion. — Honors
John Baptiste Triumfetti of Bologne, physician and director of the
Rome Botanic Garden.
Several species supply a fiber similar to hemp. My indebtness
to the exactingly executed and intelligent revision of Ko Ko Lay
418 Field Museum of Natural History — Botany, Vol. XIII
will be apparent, but it seems to me that when the evidence is
all in fewer entities, probably with more variations, will be rec-
ognized.
Flowers large, distinctly longer than 2 cm.; fruit spines 1-2 mm.
long, minutely pubescent T. calycina.
Flowers about 1 cm. long (rarely 2 cm.), usually shorter; fruit spines
retrorsely hispidulous to glabrous, 3-6 mm. long.
Leaves green, usually soon glabrescent, the trichomes often mostly
simple.
Indument of leaves scattered, stellate; flowers 14-18 mm. long.
T. grandiflora.
Indument sparse but rather even, mostly simple; flowers 8-13
mm. long T. bogotensis.
Leaves more or less canescent, at least beneath, with stellate or
mostly stellate trichomes.
Buds about 1 cm. long (8-11 mm.), appendage 1 nam. long;
leaves with some simple trichomes T. abutiloides.
Buds shorter than 1 cm. (5-8 mm.), appendages obsolete or
often 2 mm. long or longer; trichomes all stellate.
Leaves obviously 4-glandular at or near sinus; fruit body
6-8 mm. thick; spines many T. althaeoides.
Leaves eglandular or not obviously 4-glandular; fruit body
3-5 mm. thick; spines 75 or fewer except T. Bartramia.
Petals as gonophore developed, the latter at least 0.5 mm.
long.
Sepals deeply cucullate; gonophore short; tomentose
fruit body 3-4 mm. thick; spines glabrous or
essentially T. Bartramia.
Sepals appendaged or not deeply cucullate; gonophore
obvious, 5-glandular; spines hispidulous.
Sepal appendages to 2 mm. long; fruit body pubes-
cent, 3-5 mm. thick, spines 75 T. semitriloba.
Sepal appendages 3 mm. long; fruit body tomentose,
2-3 mm. thick; spines 25-40 T. oligacantha.
Petals as gonophore absent or obsolete; fruit body about
3 mm. thick, lightly stellate; spines hispidulous,
about 50 T. Lappula.
Flora of Peru 419
Triumfetta abutiloides St. Hil. Fl. Bras. Mend. 1: 287. 1827;
365.
Allied and in general similar to T. hogotensis but the indument
at least in part tomentose, the simple trichomes often gland-tipped,
the stellate approximate, the sepal appendages only about 1 mm.
long, and, especially, the mature fruit with six 1-seeded cells, the
septa false, the spines 100 or more; terminal leaf-lobe long-acuminate,
lateral lobes usually obtuse; petals 7-10 mm. long; stamens 20, the
filaments retrorsely serrulate; fruit body somewhat pubescent, in
Peru often nearly glabrous, 3-5 mm. thick. — Determinations mostly
by Lay, as elsewhere. I am not entirely convinced that the species
is in Peru. F.M. Neg. 35401.
Lima: Chosica, ^97 (distr. as T. semitriloha) . — Hu^nuco: Near
Hudnuco, 205S; S521 (det. Macbride, T. semitriloha) ; Ruiz & Pav6n.
Puente Durand, Stork & Horton 9572 (det. Standley, T. semitriloha).
Cuchero, Poeppig IW. — Loreto: Mishuyacu, Killip & Smith 2989 J^.
— Cuzco: At 700 meters. Biles. Marcapata, Vargas 3079. Torontoi,
Cook & Gilbert 819. Santa Ana, Cook &Gilhert 1^37. To Colombia,
Haiti, Argentina and Brazil. "Rata-rata" (Biies).
Triumfetta althaeoides Lam. Encycl. 3: 420. 1791; 371. T.
semitriloha Jacq. f. althaeoides (Lam.) Uittien in Pulle, Fl. Surinam
3: 56. 1932.
Openly growing bush sometimes a couple of meters high more or
less ferrugineous-tomentose the trichomes all stellate, the leaves
obscurely if at all 3-lobed and with 4 conspicuous glands at the
shallowy cordate basal sinus the rather regular serrations also there
usually glandular; petioles 4-6 cm. long; leaves ordinarily at least
1 dm. long, nearly as wide; cymes of 3-4 cjmiules or terminal
branches, peduncles 3-5 nrni. long; pedicels 1-2 mm. long; flowers
hermaphrodite, the buds about 6 mm. long, apical appendages 1 mm.
long or nearly obsolete; sepals 6-7 mm. long; petals linear, 3-4 mm.
long; gonophore stout, about 0.75 mm. long, the glands small;
urceolus less than 0.5 mm, high, 5-lobed; stamens 20, filaments
6-serrate; fruit body 6-8 mm. in diameter, sparsely stellate, 4-5-
celled, later falsely 8-celled, not more than 6 seeds maturing; spines
about 200, to 3 mm. long, retrorsely pilosulous; seeds p3rriform,
about 2 mm. long, 2.5 mm. broad. — Illustrated, Ko Ko Lay, I.e. 372.
Loreto: Near Iquitos, Williams 1401; 7950; 7958; Tessmann 3592
(det. Burret, T. semitriloha). Mishuyacu, Klug 1086; 1289. Balsa-
puerto, Klug 305 j^. Purubana, Williams 1316; 132U. Rio Nanay,
420 Field Museum of Natural History — Botany, Vol. XIII
Williams W. Rio Itaya, Williams 138. Near Caballo-Cocha,
Williams 2062; 2068; 206It. To Trinidad. "Caballosa" (Tessmann).
Triumfetta Bartramia L. Syst. Nat. ed. 10: 1044. 1759; 382.
T. indica Lam. Encycl. 3: 420. 1791? T. rhomhoidea Jacq. Enum. PI.
Carib. 22. 1760, fide Fawcett & Rendle. Bartramia indica L. Sp. PI.
389. 1753.
Shrub, a meter tall or taller, the older branches glabrous and
conspicuously white-lenticellate, the younger densely and shortly
stellate- tomentose as the slender petioles, these 3-5 cm. long;
leaves broadly ovate to rhombic-ovate, obscurely to deeply 3-5-
lobed, usually 4-5 cm. long and nearly as wide, sometimes 7-8 cm.
long, 6-7 cm. wide, obtuse or rounded at base where rarely glandular,
abruptly acuminate, nearly glabrate or with scattered coarse short
trichomes; cymes of 3-5 cymules axillary, peduncles 1-2 mm. long,
pedicels 1 mm. long; flowers hermaphrodite, the buds most expanded
at apex; sepals deeply cucullate, 5-7 mm. long, lightly stellate or
glabrate; petals broadly obovate, about 5 mm. long; gonophore very
short, glands small, urceolus deeply many-lobed; stamens 10-15,
filaments with 4-6 retrorse serrations; fruits in dense nodose clusters,
the densely tomentose body about 3-4 mm. in diameter, (2) 3-celled,
the cells 1-2-ovuled, 1-seeded, the 75-100 spines 1-1.5 mm. long,
nearly or quite glabrous. — Following the classification specialists
have established in the Malvaceae this is a generic type, since the
fruit is not naturally dehiscent. Illustrated, Mart. Fl. Bras. 12,
pt. 3: pi. 27.
San Martin: Tarapoto, Williams 5925. — Loreto: Iquitos, Killip
& Smith 27229. Puruchana, Williams 1314. New and Old World
tropics.
Triumfetta bogotensis DC. Prodr. 1: 506. 1824; 366. T.
Jelskii Szyszyl. Diss. Math. Nat. Acad. Litt. Cracoy [Elrakow],
29: 224. 1895?
A meter or two high, the indument mostly stellate and long single
trichomes mixed, the latter predominant or alone on the long
branches, in the inflorescence axils and on the sepals without;
petioles tomentose, 4-6 cm. long; leaves broadly ovate, usually
3-lobed, to about 1 dm. long, 8 cm. wide, rounded to subcordate at
base, acuminate, irregularly serrate, the simple trichomes of the
pubescence appressed; cymes axillary, peduncles 4-7 mm. long,
pedicels 3^ mm. long; flowers hermaphrodite, buds oblongoid,
constricted below the slender apical appendages, these 2-3 mm.
Flora of Peru 421
long; sepals oblong, 8-13 mm. long; petals broadly obovate, 8-9
mm. long, 3-4 mm. wide, claw densely plumose; gonophore slender,
1 mm. long, equaled by the glands, the short unlobed urceolus
distinctly ciliate; stamens 25-30, the filaments not serrated; fruit
body at maturity 3^ mm. in diameter, 3-celled, the slender re-
trorsely pilosulous spines about 3 mm. long; seeds 6, lenticular, 2-3
mm. long, about 2 mm. wide. — The identity of T. Jelskii, allied by
the author to T. caudcUa Tr. & PI. of Colombia, incompletely known,
the type not seen by Ko Ko Lay, is not certain ex char, but except
for the "subsessile flowers" which may be due to undeveloped
condition, the description seems to fit this species, the "laxly hirsute
leaves" especially suggesting it.
Cajamarca: Cascas and Cutervo, Raimondi ("vel affine," Burret).
Tambillo (Jelski 280, type, T. Jelskii).— Cuzco: San Miguel, Cook
& Gilbert 891 . Machupicchu, Vargas 795. Tropical America.
Trlumfetta calycina Turcz. Bull. Soc. Nat. Mosc. 36: 574.
1863. T. midtilocularis Hochr. Ann. Cons. Jard. Bot. Gendve 18:
104. 1914?; 356.
Spreading shrub often forming clump or a small tree 2-A meters
high, densely stellate-tomentose, the more or less spreading trichomes
rather ferrugineous; petioles 4-6 cm. long; leaves broadly ovate,
narrowly acuminate, irregularly serrate, usually 3-lobed, to about
1 dm. long, 6-7 cm. wide; cjmies axillary usually with only
1 cymule, peduncles 8-12 mm. long, pedicels about 6 mm. long;
flowers hermaphrodite, the buds 22-25 mm. long, appendages 2-3
mm. long; sepals about 27 mm. with appendages; petals 24 mm.
long, 4-6 mm. wide, the 4 mm. claw densely plumose; gonophore
to 2 mm. long, the glands to 1.5 mm.; urceolus 10-lobed; stamens
25-30 (or sometimes 16?), the filaments retrorsely 2-serrulate; fruit
body 6-8 mm. in diameter, lightly and deciduously stellate, 3-5-
celled becoming 8-9-celled and -seeded, the many puberulent spines
only 1-2 mm. long; seeds ovoid, about 2 mm. long and broad. — It
may be an error to use Turczaninow's name, based on Mathews 889
without data for this species, but, as Ko Ko Lay points out, the
description agrees except that the author noted the stamens as 15
or 16; this is probably a slip of the pen or the printer since it seems
unlikely that this distinctive plant, compared with T. mollissima
HBK. of Colombia in herbaria, but that with smaller flowers,
tomentose fruits, has not been found again with so few stamens.
P.M. Neg. 23820 (T. muUilocularis) .
422 Field Museum of Natural History— Botany, Vol. XIII
Ancash: Higaris, 2^52. — Hudnuco: Yanano, 3809. Acomayo,
Woytkowski 3U013 (det. Cuatrecasas). Ambo, 3150. Pampayacu,
Kanehira 112. Near Hudnuco, Sawada P72; Ruiz & Pav&n;
Dombey. Mito, 1571. — Junin: Carpapata, Killip & Smith 2^61;
Stork 10970; Ochoa 287. Huacapistana, Rose 18550; Sandeman W9.
Ayacucho: Near Huanta, Killip & Smith 22309. — ^Apurimac:
Abancay, Goodspeed Exped. 10552. Marcapata, Vargas 9685. San
Miguel, Cook & Gilbert 1126; 1163. Machupicchu, Vargas 796;
West 6U73; Soukup 143. — Puno: Soukup 521. Sandia Weberbauer
507 (as T. macrantha). Valle del Urubamba, H err era 962; 2653.
Prov. del. Cercado, Weberbauer (as T. macrantha). To Colombia.
"Ratan," "r'ata-r'ata" (Vargas).
Triumfetta grandiflora Vahl, Eclog. Amer. 2: 34. 1798; 363.
T. Schunkei Macbr. Candollea 5: 381. 1934?, fide Ko Ko Lay.
Shrub 2-4 meters high, the branches stellate-hirtellous or gla-
brate; leaves long-petioled, ovate or broadly ovate, large, cuspidate-
or caudate-acuminate, rounded or subcordate at base, glandular-
serrate, almost glabrous in age, early sparingly pubescent with
small stiff stellate trichomes; sepals glabrate, 15-18 mm. long,
about equaled by the linear-oblanceolate petals; capsules 1-1.5 cm.
in diameter, nearly or quite glabrous including the many slender
prickles. — After Standley and Steyermark, Fl. Guatemala, Fieldiana:
Bot. 24, No. 6: 322. 1949. Perhaps should be interpreted to include
T. mollissima. My plant was proposed primarily on the character of
style entire, apparently found to be variable by Lay. The sepals
and petals of this species were described as only about 12 mm. long,
but they may be nearly 15 mm. long when moistened. F.M. Neg.
32620 (Poeppig).
Junin: Chanchamayo Valley, Schunke 1597 (t5npe, T. Schunkei).
— Loreto: Yurimaguas, Poeppig 2053. To southern Mexico; West
Indies.
Triumfetta Lappula L. Sp. PI. 444. 1753; 378. T. quinqueloba
Turcz. Bull. Soc. Nat. Mosc. 36, pt. 1, 1: 574. 1863, fide Lay. T.
heterophylla Lam. Encycl. 3: 420. 1791. T. Hostmanni Miq. Linnaea
22: 466. 1849.
Slender, tall, frutescent, the often lax branches coarsely fer-
rugineous-tomentose, none of the trichomes simple; petioles elon-
gate, sometimes nearly 1 dm. long; leaves broadly ovate, commonly
somewhat pandurately 3-5-lobed, the unequal serrations usually
Flora of Peru 423
glandular, obtuse to rounded at base, the lobe-tip, at least the
terminal, acuminate, often less than 1 dm. long, nearly as wide,
sparsely pubescent above, more tomentose beneath; cymules usually
only 2 in each axillary cyme, the flowering peduncles 2-3 mm. long,
the pedicels shorter; flowers hermaphrodite, the medially constricted
buds 3-5 mm. long, the very short appendages rarely wanting;
sepals densely tomentose, 4-6 mm. long; petals undeveloped as
gonophore and glands, the urceolus nearly obsolete; stamens 10
(rarely 5 or 15), the filaments smooth; fruit body about 3 mm. in
diameter, sparsely stellate, 3- or sometimes 2-celled by abortion,
the slender retrorsely pilosulous spines 2-3 mm. long. — The plant
of Lamarck with body of fruits 2-2.5 mm. thick may be a variant,
apparently the common form in Peru; it is considered distinct by
Fawcett & Rendle, Fl. Jamaica 5: 84. 1926. Illustrated, Ko Ko Lay,
I.e., 378; Mart. Fl. Bras. 12, pt. 3: pi. 27 (T. keterophylla) . F.M.
Neg. 23817 (T. Hostmanni).
San Martin: Chazuta, Kliig U150 (det. Standley). — Junin: La
Merced, KiUip & Smith 23^19 (det. Standley).— Amazonas(?):
(Mathews 1625, type, T. quinqueloba) . — Loreto: Balsapuerto, Klug
3093 (det. Standley, T. semitriloba). Iquitos, Killip & Smith 2723 J^
(det. Macbride, T. rhomboidea). Bolivia to the West Indies; western
Africa.
Triumfetta oligacantha Hochr. Ann. Cons. Jard. Bot. Geneve
18: 106. 1914; 377.
Closely resembles T. semitriloba but the leaves not lobed, caudate-
acuminate, rather regularly serrate; peduncles 8-10 mm. long,
pedicels 3-5 mm. long, sepal appendages about 3 mm. long, stamens
about 30, the filaments smooth, and, especially, fruits with 3 cells
each 1-seeded and spines only 25-40. — It seems probable that the
type, as many labeled simply Pavdn, was actually collected by
Tafalla, particularly since the species is otherwise known from the
region of Guayaquil.
Peru(?): Without data, Pavdn, type. Adjacent Ecuador.
Triumfetta semitriloba Jacq. Enum. PI. Carib. 22. 1760; Sel.
Stirp. Amer. Hist. 147. 1763; 373.
Suffrutescent, the older indument including that on the leaves
rather scabrous, short and entirely of stellate trichomes, early often
abundant and more or less tomentose; petioles 3-6 cm. long, the
trichomes in separate tufts; leaves usually broadly ovate but vari-
424 Field Museum of Natural History — Botany, Vol. XIII
able, often more or less 3-lobate and only the terminal lobe acumi-
nate, sometimes also the lateral, ordinarily to about 8 cm. long,
6 cm. wide; serrations commonly glandular, always very unequal;
cymes rarely terminal, with 2-3 cymules, peduncles and pedicels 2-3
mm. long; flowers hermaphrodite, 5-8 mm. long, appendages 1-2 mm.
long; sepals greenish, 6-10 mm. long, subequaled by the petals; gono-
phore less than 1 mm. long, urceolus scarcely half as long; stamens
15-25, the filaments 4-serrulate; fruit body 3-5 mm. in diameter,
lightly to rather densely stellate, 3-celled, each cell 2-seeded but
rarely 6-seeded, the 50-75 slender retrorsely hispidulous spines 2-3
mm. long. — Illustrated, Ko Ko Lay, I.e. 374.
San Martin: San Roque, Williams 7160 (det. Standley, T.
Lappula). — Junin: Chanchamayo Valley, Raimondi (det. Burret).
— Loreto: Yurimaguas, Williams 3950; Ji360 (det. Standley).
Widely distributed in the tropics.
3. HELIOCARPUS L.
Reference: Ko Ko Lay, Ann. Mo. Bot. Gard. 36: 507-541. 1949.
Stellate pubescent trees or shrubs with serrate usually somewhat
3-lobed leaves and small flowers in terminal or rarely axillary
panicled cymes, these of hermaphrodite or pistillate flowers, the
latter apetalous. Sepals free, 4 as the basally foveolate petals,
these about the base of a more or less elevated torus, this 4-glandular
below the many distinct stamens. Ovary 2-celled (falsely 4-celled
at base), the cells pseudo-septate between the 2 ovules; style filiform,
the stigma bidentate or biparted. Carpels small, compressed, with
2 series of plumose bristles, each cell with 1 pendulous seed, the
endosperm oily.
Bark produces a strong durable fiber and the wood of the rapidly
growing Peruvian species, according to Ruiz and Pavon, was
favored for rafts. Seibert observed that it yields a latex of possible
value.
Heliocarpus popayanensis HBK. Nov. Gen. & Sp. 5: 341.
1823; 532. H. americaniis L. var. popayanensis (HBK.) Schum.
in Mart. Fl. Bras. 12, pt. 3: 142. 1886. H. stipulatus Hochr. Ann.
Cons. Jard. Bot. Geneve 18: 121. 1914, fide Ko Ko Lay.
Older branches sparsely lenticellate, glabrate, the younger
somewhat tomentose with stellate and simple trichomes, the long
petioles densely; leaves more or less distinctly 3-lobed, often 1.5-2
Flora of Peru 425
dm. long, 14-18 cm. wide, finally deeply cordate, lightly stellate
above, glabrate in age, usually densely stellate beneath the pu-
bescence, on the nerves mostly simple; inflorescences commonly
terminal, the hermaphrodite about a dm. long and slightly wider,
the pistillate usually about 1.5 dm. long, 2 dm. wide; flowering
peduncles 3-radiate; buds without appendages at sepal tip; sepals
5 mm. long, petals 4 mm. long, in pistillate flowers the former 3-4
mm. long, the latter wanting; stamens about 12; style shortly bifid,
each stigma with 3 acute lobes; fruits ellipsoid to ovoid, slightly
tomentulose or glabrate. — Leaves are frequently without lobes at the
southern and northern limits of the range and present great diversity
of shape and size, according to Ko Ko Lay, upon whose careful
well-considered revision I have freely drawn with appreciation.
Sometimes 30 meters; flowers white, greenish-red or roseate.
Wood used for rafts (Ruiz & Pav6n). — Determinations, except as
noted, by (or verified by) Lay. Illustrated, Ko Ko Lay, I.e., 533.
F.M. Neg. 35443.
Cajamarca: Querocotillo, 2,000 meters, Weberbauer 712U. Nan-
cho, Raimondi. — San Martin: Moyobamba, 890 meters, KliLg S627;
W&ytkowski S5S17. Tarapoto, Spruce j^58. Juanjui, Kliig JtSBS. —
Junin: Vitoc, Ruiz & Pavdn. Near La Merced, KiUip & Smith
23926 (det. Standley). Pichis Trail, KiUip & Smith 2586^. Chan-
chamayo Valley, Schunke 290; 293; 1599. — Hudnuco: Near Hudnuco,
Sawada 101. Tocache, Poeppig 189U; 3102 (tjrpe, H. stipulatus
Hochr). Cuchero, Dombey; Ruiz & Pavdn. Monz6n, WeberbaiLer
3432. Muila, 4065. Sunny brushy slope, near Pozuzo, 4765. —
Loreto: Yurimaguas, KiUip & Smith 28166; 27831; 27853; Williams
4297. Balsapuerto, Klu^ 3076. — Cuzco: Prov. Convencion, Soukup
790. San Miguel, Cook & Gilbert 1072.— Madre de Dios: Iberia,
Seibert 2029 (det. Killip).— Rio Acre: Mouth of Rio Macauhdn,
Krukoff 5261. Western South America. "Palo de balsa" or "huam-
po" (Ruiz & Pav6n); "llausa-quiro" (Williams); "llaosa-pancho"
(Cook & Gilbert); "yausu-quiru" (Woytkowski) ; "huampo bianco"
(Sawada).
4. APEIBA Aublet
Reference: Uittien, Recueil. Trav. Bot. N^rl. 32: 244-251. 1935.
Trees or tree-like shrubs with ample 3-5-nerved leaves and
greenish-yellow flowers disposed in 2-3-dichotomous cymes that are
terminal or opposite the relatively short petioles. Sepals free,
usually, as the short petals, 5, these smooth to base. Stamens
426 Field Museum of Natural History— Botany, Vol. XIII
indefinite; filaments short; anthers erect, linear with a membranous
connective produced apically. Ovary 10 (8) -many-celled, the cells
many-ovuled; style simple. Fruit large, disk-like or depressed-
globose, tubercled or coarsely bristly, coriaceous, dehiscing centrally,
the numerous compressed seeds in a lustrous pulp. — Uittien in
Pulle, Fl. Surinam 3, pt. 1 : 438-440. 1941, revised his work in accord
with the observations of Ducke, Archiv. Inst. Biol. Veg. Rio Jan.
4: 51-52. 1938. Neither of these able botanists, however, have
succeeded in including exactly in their classifications the Peruvian
examples of these interesting trees, suggesting that there are more
species than they describe or fewer and these more variable.
Fruit covered with long hirsute or hispid bristles; branchlet tips
and petioles long-villous; leaves denticulate.
Leaves distinctly cordate at base; fruit subglobose. .A. tibourhou.
Leaves obscurely or not cordate; fruit depressed . .A. Schomhurgkii.
Fruit sharply echinate; branchlets as petioles glabrous or puberulent;
leaves entire A. memhranacea.
Apeiba membranacea Spruce ex Schumann in Mart. Fl. Bras.
12, pt. 3: 145. 1886.
Slender apparently glabrous tree but the leaves especially on the
nerves very minutely and evanescently puberulent and sometimes
glabrous, sometimes with tufts of trichomes in the axils of the lateral
nerves; leaves oblong-elliptic or elliptic, rounded or obscurely
cordulate at base, tip acuminate rarely minutely serrulate, the
nerves 6-9; flowers about 1.5 cm. wide; fruits depressed-globose,
about 6 cm. across, densely echinate with narrowly conic-based
short spines.— As here interpreted there is considerable variation
in the degree of development of the trichome tufts in the nerve
axils of the leaves beneath. Most of the Peruvian collections except
as noted were referred to A. aspera Aublet, Brazil to Guianas,
included by Uittien in A. glabra Aublet with bristly fruits and
4-5-nerved leaves and A. echinata Gaertn., same range, with nearly
the fruits of A. membranacea but the leaves grayish-tomentulose
beneath. However, the Peruvian tree seems to be the same as
A. intermedia Uittien of Surinam but the fruit of that species is
unknown. Attains 25 meters, the trunk 3 dm. in diameter (Tess-
mann) ; flowers golden or ochre yellow. F.M. Neg. 23809.
Hudnuco: Chicoplaya, Ruiz & Pav6n. — Loreto: Yurimaguas,
Poeppig 21 U; Williams 4.012; Killip & Smith 29075. Rio Mazan,
Flora of Peru 427
Josi Schunke 95. Rio Itaya, Williams SSI 5. Florida, Klug 2S4S
(det. Standley). Mouth of Rio Santiago, Tessmann U622 (det.
Biuret).— Rio Acre: Ule 9585; 9598. Mouth of Rio Macuahdn,
Krukoff 5S0Jt. Colombia; Amazonian Brazil. "Maqui-supa"
(Williams); "maqui-zapanaccha" (Schunke).
Apeiba Schomburgkii Szyszyl. Diss. Math. Nat. Acad. Litt.
Cracov. 27: 140. 1894. A. tihcurhou Aubl. var. membranacea Lockh.
ex Griseb. Fl. Brit. W. Ind. 708. 1864, fide Uittien.
Branchlets and petioles villous with long trichomes; stipules
more or less promptly deciduous; leaves rounded or obscurely
cordate at base, membranaceous, smooth above but early minutely
stellulate especially on the nerves and with a few deciduous long
simple trichomes, lightly stellate- tomentose beneath, 7-9- (12)
nerved; flowers white; fruit depressed, nearly disk-shaped, opening
with 3-7 rather large woody teeth. — After Uittien; in the Peruvian
specimen that seems to belong here the leaves are 9-12-nerved and
the fruit bristles are hispid with appressed trichomes. F.M. Neg.
9251.
Loreto: Puerto Arturo, Williams 500 J^ (det. Standley, A. tibour-
bou). To Colombia and the Guianas. "Maqui-sapa-fiaccha"
(Williams).
Apeiba tibourbou Aublet, PI. Guian. 1: 538, pi. 21S. 1775.
A. hirsuta Lam. Encycl. 1: 208. 1783. A. tibourbou Aublet var.
rugosa Szyszyl. Diss. Math. Nat. Acad. Litt. Cracov. 27: 140. 1894.
A. albiflora Ducke, Archiv. Jard. Bot. Rio Jan. 3: 209, pi. 20. 1922,
fide Ducke.
Branches, petioles, these 1-3 cm. long and prominent ovate-
lanceolate rather tardily deciduous stipules hirsute with spreading
trichomes; leaves firm-chartaceous, typically densely stellate-
tomentose beneath, more or less rugose and sparsely stellulate above,
distinctly cordate at base, 9-15-nerved; flowers yellowish or white,
about 2.5 cm. wide, the petals 1-1.5 cm. long; fruit nearly globose,
slightly depressed, opening with minute woody teeth, the flexible
bristles abundantly spreading hirsute. — After Uittien. The Peru-
vian specimens vary in having scarcely to very rugose leaves often
lightly tomentose beneath; Tessmann recorded one tree 30 meters
tall the trunk 1 dm. in diameter but Williams noted the tree as
attaining 12 meters with often inclined small buttressed trunks
about 3 dm. in diameter, the wood spongy. Determinations mostly
428 Field Museum of Natural History— Botany, Vol. XIII
by Standley. Illustrated, Mart. Fl. Bras. 13, pt. 3: pi 29 (except
fruit). F.M. Neg. 9249.
San Martin: Chazuta, Klug WA- Juanjui, Klug ^21 3. Tara-
poto, Williams 56H; 6727. — Junin: La Merced, 5569. — Loreto:
Mishuyacu, Klug 1087. Yarina Cocha, Tessmann 3^03 (det.
Burret). Yurimaguas, Poeppig 2097. — Rio Acre: Mouth of Rio
Macauhdn, Krukoff 5291. To Central America, the Guianas and
the West Indies. "Maqui-sapa-naccha," "maqui-sapa" (Williams) ;
"tibourbou."
5. MOLLIA Mart.
Reference: Baehni, Candollea 5: 403-425. 1934.
Usually pubescent often slender branched trees with entire or
only apically serrate leaves and rather showy white flowers solitary
or fasciculate in the axils, sometimes pedunculate. Sepals free,
5 as the petals. Stamens indefinite shortly coalescent in clusters
of 10, the 5 outer longer alternate to the petals; anthers narrowly
sagittate. Ovary 2-celled, the cells many-ovuled; style filiform.
Capsules 2-celled, ligneous, shortly alate apically, the cells semi-
2-valved, spuriously septate between the many compressed emar-
ginate seeds. — Leaves perhaps always barbate beneath in the nerve-
axils. My indebtedness to the distinguished Director of the Con-
servatory of Botany, Geneva, Switzerland, for his stimulating
association and thought-provoking work over many years may here
be given this merited recognition. Besides, the following mention
may be made of Williams 963 from Nanay especially to note the
native name "Uchu-huayo"; it has ovoid fruits barely pointed
apically.
Leaves densely lepidote only beneath, serrulate M. gracilis.
Leaves equallj^ lepidote both sides, entire M. Williamsii.
MoUia gracilis Spruce, Joum. Linn. Soc. Bot. 5: Suppl. 2: 59.
1861; 413, 424.
Slender branchlets, petioles, these 5-6 mm. long, and leaves
both sides lightly and minutely lepidote, the sessile or shortly
pedunculate 1-few-fiowered inflorescences cinereous lepidote; leaves
chartaceous, obscurely serrulate toward obtusish tips, rounded at
base, barbate in the nerve axils beneath, elliptic or oblong-elliptic,
to 10 cm. long, 6 cm. wide, often smaller; pedicels slender, 2-3 cm.
Flora of Peru 429
long; calyces tjrpically about 2 cm. long; fruits unknown. — 8-meter
tree (Tessmann). Weibel, Candollea 10: 169, 172. 1945, described
the placenta tion, showing the ovary to be 1-celled in bud. F.M.
Neg. 32618.
San Martfn: Tarapoto, Spruce It9It9. — Loreto: Santiago, Teas-
mann U198. Adjacent Colombia and Brazil.
Mollia Williamsii Baehni, Candollea 7: 136. 1936.
Younger branchlets sparsely, petioles, these 12-13 mm. long,
peduncles and leaves beneath except the nerves densely lepidote
pubescent; leaves entire, ovate, rounded or subacute at base, long-
acuminate, 11.5-13.5 cm. long, 4.5-nearly 5 cm. wide, coriaceous,
lightly squamose above, not barbate in the axils of the 3 nerves
beneath; inflorescences shortly ped uncled, 2-4-flowered, fruiting
pedicels 2-2.5 cm. long; capsules opaque, scabrellous, sparsely
lepidote basally, densely apically, distinctly mucronate, ovoid,
complanate, narrowly alate, depressed laterally at septum, 11-13
mm. long, 15-17 mm. wide, 11-12 mm. thick. — Fruits recall those
of M. lucens Baehni of the Rio Negro but those are lustrous and
scarcely bilobed, and the short peduncles and larger long-acuminate
leaves mark M. Williamsii easily. Type from a straight cylindric-
trunked tree about 7 meters tall.
Loreto: Rio Nanay, Williams 1160, type. "Uchu-mullaca."
6. LUEHEA Willd.
Reference: Burret, Notizbl. Bot. Gart. Beriin 9: 822-837. 1926.
Trees or tall shrubs, the usually dentate leaves stellate-tomentose
beneath, the showy white or rose-colored flowers disposed in multi-
bracteolate axillary cymes or terminal panicles. Sepals and petals
5, the latter glandular-thickened within at base. Stamens many
obscurely or very shortly coalescent (in one species completely con-
nate) in phalanges of 5 or 10, the filiform exterior without anthers,
basally lanate or pubescent; anthers of the inner sagittate. Ovary
5-celled, many-ovuled; style simple. Capsules subligneous, semi-
5-valved, the numerous seeds wing-margined above. — The bractlets
may be somewhat coalescent but are never jjermanently united; the
anthers are parallel-connate to apex, and dehisce at the same time
by long chinks (Burret). The nerve-axils beneath are usually if not
always without the tufts of trichomes found in MoUia.
430 Field Museum of Natural History— Botany, Vol. XIII
Leaves oblong-elliptic or elliptic but mostly rather more than twice
as long as wide; sepals 1-2 cm. long; capsules at most 3 cm.
long, obovoid.
Petals oblanceolate; leaves acuminate; capsules nearly shaggy
pubescent below L. Tessmannii.
Petals obovate; leaves shortly acute; capsules rather appressed
pubescent L. paniculata.
Leaves broadly elliptic, mostly less than twice as long as wide;
sepals about 3 cm. long; capsules 3 cm. long or longer.
Capsules cylindrical, 3.5 cm. long or longer; staminodia filaments
shorter than tube L. grandiflora.
Capsules sphaeroid, 3 cm. long; staminodia filaments longer than
tube L. tarapotina.
Luehea grandiflora Mart. Nov. Gen. & Sp. 1: 99, pi. 61. 1826;
832. L. densiflora St. Hil. and L. laxiflora St. Hil. Fl. Bras. Merid.
1: 293-294. 1827, fide Burret, ex char.
Branches and branchlets slender, glabrous; petioles stout, 5-10
mm. long; leaves elliptic, very shortly acuminate or apiculate,
slightly oblique at the rounded-subcordate base, commonly less than
twice as long as wide, denticulate, usually closely, glabrous or
nearly above, stellulate and more or less arachnoid tomentulose
beneath, the nerves and veins finally glabrate; flowers usually
several in dense or open inflorescences, the early conspicuous sepa-
rating narrow bractlets about as long as the sepals, these at lieast
about 3 cm. long, the oblong or elongate-obovate petals gradually
attenuate to the narrower or scarcely narrower claw; free part of
the staminodia short, fringe-like; capsules to 2.5 cm. long or longer,
less than half as thick, densely ferrugineous with stellate- tomentum.
— After Burret in part, who points out that there are floral differences
between this species and the more northern L. speciosa Willd. with
which it has been confused; apparently his conclusions are sound
but the characters relied upon may prove to be variable. The
Peruvian material seen is meager but seemingly it belongs here.
Illustrated, Mart. Fl. Bras. 12, pt. 3: pi. SI (as L. speciosa, fide
Burret). F.M. Negs. 35411; 35413 (L. densiflora, L. laxiflora).
Loreto: Cachipuerto, Klug 3131 (det. Standley, L. tarapotina).
Yurimaguas, Poeppig 2191, Addenda 62 (det. L. speciosa in herb.).
To southeastern Brazil.
Flora of Peru 481
Luehea paniculata Mart. Nov. Gen. & Sp. 1: 100, pi. 62. 1826;
834. L. parvifolia Huber, Bol. Mus. Paraense 2; 507. 1898, fide
Ducke.
Beautiful tree, the slender branchlets, petioles (5-7 mm. long),
leaves beneath and many-flowered terminal panicles more or less
ferrugineous tomentulose; stipules promptly caducous, 4-5 mm.
long; leaves broadly oval or elliptic, truncate or subcordate at base,
shortly acuminate, 7-12 cm. long, 4.5-7 cm. wide, more or less
inequilateral, coriaceous, glabrous above at maturity, unequally
dentate; bracts 4-5 mm. long; peduncles and pedicels 5-10 mm.
long; involucrate bracts 9, linear, acute, 8-9 mm. long; sepals
lanceolate, 11-12 mm. long; petals irregularly crenulate, about 16
mm. long, 13 mm. wide, white changing to rose, puberulent within
above the entire basal gland; stamens in 3-4 clusters, 4-7 mm. long;
ovary orange-tomentose; capsule about 2 cm. long, 12-14 mm. thick
medially, reddish tomentose, the seed wings rounded apically. —
Most of the Peruvian collections fide Burret and not seen by me
at least during the final preparation of this compilation. F.M.
Neg. 19678.
San Martin: Tarapoto, {Spruce U880). Morales near Tarapoto,
Williams 5699. — Hudnuco: Chinchao, Ruiz & Pavdn; Raimondi
(det. Burret); Rivero. Grass steppe, Prov. Huamalies, 1,000 meters
(Weberbauer S492).—Junin: Peren^, 700 meters (Weberbauer S29).
— Cuzco: Sta. Anna, Prov. Convenci6n {Weberbauer 5021; 50It7).
To southeastern Brazil. "Inchato" {Weberbauer); "calzoncillo-
panga" (Williams).
Luehea tarapotina Macbr. CandoIIea 5: 382. 1934.
Younger branchlets early reddish furfuraceous with short
stellulate indument; petioles 6-7 (12) mm. long; leaves broadly
and subobliquely elliptic, lightly cordate at base, subabruptly and
broadly acuminate, to 14 cm. long, 8 cm. wide, irregularly dentate,
chartaceous, green, slightly lustrous and very sparsely stellate
puberulent above, rusty-cinereous tomentulose beneath the nerves
and veins reticulate; fruiting pedicels about 2 cm. long, furfuraceous-
puberulent; capsules pentagonous-sphaeroid, obscurely angled,
about 3 cm. long, 2 cm. thick, densely and persistently rusty pu-
bescent; seeds lustrous, 7 mm. long. — The permanently pubescent
fruits and smaller seeds seem to distinguish this from L. apeciosa
Willd. The Klug specimen seen shows imperfect flowers; apparently
however the sepals attain 3 cm., the petals are obovate, rather
432 Field Museum of Natural History — Botany, Vol. XIII
abruptly broad-clawed, and the filamentous portion of the stamino-
dial phalanges is slightly longer than the united part or tube; the
significance of this character may be open to question. However,
the capsules are distinctive among the Peruvian species; but as
noted by Burret in his careful revision the fruits of several species
(Bolivian) are unknown.
San Martin: Juanjui, Klug J^3J^7 (det. Standley). Morales near
Tarapoto, Williams 570^, type. "Calzoncillo."
Luehea Tessmannii Burret, Notizbl. Bot. Gart. Berlin 9: 836.
1926.
Younger branches terete, dark red, glabrous, longitudinally
rugulose in drying, the divaricate branchlets above as the petioles
(4-8 mm. long) and inflorescences (5-8 cm. long) yellowish-stellu-
late; leaves elliptic or slightly elliptic-obovate, rather abruptly and
shortly but conspicuously acuminate, obliquely rounded at base,
evidently denticulate toward the tip, often 5.5-9.5 cm. long, 2.5-4.5
(6) cm. wide, subcoriaceous, soon glabrous above, very minutely
and closely puberulent with a cinereous or in herb, ochraceous
indument, clearly stellulate only if at all on the more prominent
nerves and the conspicuous clathrate-reticulate veins; bractlets
deciduous before anthesis, much shorter than the oblong sepals, these
yellowish strigose-pilose without, 12 mm. long; petals narrowly
oblanceolate, 14 mm. long, basally pilose; ovary yellowish-villous;
capsules 1.5-2 cm. long, densely villous especially toward the
narrowed base, somewhat obovoid. — PYuit unknown in type. Species
nearly L. cymulosa Spruce of the Upper Amazon with oblong-
lanceolate gradually acuminate leaves, less dense indument on the
lower surface (Burret) . These distinctions appear varietal but more
material is needed of the Spruce species from the region of the type.
Type 8 meters, trunk diameter 2 dm.; attains 15 meters (Schunke).
F.M. Neg. 9247.
Loreto: Flood free wood, Rio Itaya, Tessmann SSSIt, type. Sole-
dad, Tessmann 5180. Near Iquitos, Klug lJf72; Tessmann 5122.
Rio Mazdn, Jose Schunke 132. Caballo-Cocha, Williams 2Jtl9.
"Museg-gui-ey" (Klug, Huitoto); "boleyna" (Schunke).
7. LUEHEOPSIS Burret
Reference: Burret, Notizbl. Bot. Gart. Beriin 9: 83&-843. 1926.
As the name implies there are no obvious vegetative characters
to distinguish these trees from Luehea, but the bractlets are com-
Flora of Peru 433
pletely connate, and not separating in age; the petal bases are less
thickened, the stamens are always united below into a tube and
esp)ecially the anthers are free above the middle, somewhat divergent
and before anthesis with triangular subcochleariform apical opening,
this chink finally extending to base. — The flowers in some species
are precocious (Uittien). These observations have added to an
understanding of the species-relationships but from a standpoint
of floristic, which is to say practical, taxonomy they ought to
indicate a subgroup within Ldiehea from which except in flower they
are not distinguishable. As the thoughtful author has detailed in his
remarks the characters of bractlets and stamen-connation are found
in varying degree within the more narrowly defined Luehea.
Lueheopsis Juliani Macbr., sp. nov.
Arbor, 10 m. alta; foliis ut videtur non evolutis jam florens;
ramulis junioribus petiolisque (11 mm. longis) striatis et minute
stellulato-puberulentis; vetustioribus glabris; foliis late ellipticis
vel elliptico-obovatis basi valde inaequaliter rotundatis apice
breviter abrupteque acuminatis 11-16 cm. longis, circa 7 cm. latis
prope apicem grosse repando-dentatis coriaceo-chartaceis, supra
glabris nitidis subtus aequaliter et brevissime adpresseque cinereo-
pubescentibus, nervis principalibus 3 et venis transversalibus subtus
prominentibus subclathratis vix vel obscure reticulatis; ramulis
inflorescentiorum plerumque 1-3 cm. longis ut floribus stellulato-
pulverulentis; pedunculis circa 5 mm. longis; cupula 5 mm. alta,
dentibus ovato-acutis 1-2 mm. longis vel demum profundius fissis;
sepalis 14 mm. longis intus sericeo-strigosis; petalis vix 14 mm.
longis extus ad basin distincte adpresseque pilosiusculis; antheris
1.5 mm. longis. — Evidently near to and possibly a variant of L.
Hoehnei Burret, I.e. 841, of Mato Grosso, Brazil, to which species
Standley referred it, but the much larger leaves strongly oblique at
base and the pubescent petals suggest that it is specifically estab-
lished. The t)rpe consists only of a leafy branchlet and a separate
leafless inflorescence; it is probable that the honest collector Klug got
them from the same tree and while the size of the leaves may be due
to their being on a shoot their form is different from those of Burret's
tree and the petals of the latter are described as glabrous without,
the flowers too, apparently accompanied by leaves.
In naming this tree for the energetic and enthusiastic Curator
of the Department of Botany, Chicago Natural History Museum,
Julian Steyermark, I am sincerely pleased because my sense of
434 Field Museum of Natural History— Botany, Vol. XIII
indebtedness for his cooperation in selection of materials for the
preparation of this work is great.
Loreto: Mishuyacu near Iquitos, Klug 2528, type.
8. NEOTESSMANNIA Burret
Tree with stellate and some simple indument, simple alternate
leaves, regular hermaphroditic 4-5-merous flowers with valvate
distinct sepals, many free stamens, all equal, and with 2-celled
introrse linear anthers dehiscing by a short longitudinal nearly
apical chink, the pollen grains smooth, in tetrads pyramidally
connate. Ovary inferior, in lower part multicelled, in upper part
1-celled with long prominent lateral membranes between which,
above and below, are affixed densely the small anatropous ovules.
Style simple with clavate tip, the stigma decurrent. — Without
fruit position uncertain but seems to require a separate section
(Neotessmannioideae Burret) characterized by the unique (for
the family) inferior ovary, multicelled below, numerous ovules
between partial or complete walls, and pollen grains cohering in
tetrads (Burret); the author suggests the available data recall the
genera Apeiha and Muntingia.
The name meritoriously commemorates the famous ethnographer
Gunther Tessmann, who made exceptionally documented botanical
collections in Amazonian Peru and in Africa.
Neotessmannia uniflora Burret, Notizbl. Bot. Gart. Berlin
9: 125. 1924.
Reddish virgate branchlets at tip, leaves, especially beneath on
the nerves and petioles (12-18 mm. long) and finally supra-axillary
1-flowered peduncles more or less densely tomentulose with yellowish
slender stellate trichomes, some glanduliferous; stipules caducous,
the scars oblique; leaves very obliquely cordate at base, obtuse at
apex, broadly oblong, to 28 cm. long, 11 cm. wide, obscurely repand-
denticulate with 5-6 basal nerves and but 6 lateral, reticulate
venation prominent and dense beneath; sepals oblong, obtuse,
tomentose on both sides, to 18 mm. long; petals yellow, obovate-
oblong, 3-3.5 cm. long, membranous, lightly tomentose near base;
filaments subulate, 4 mm. long, little shorter than basally cordate
anthers; style 4 mm. thick at base; ovary semiglobose, tomentose,
with about 25 thin more or less complete divisions. — F.M. Neg.
9248.
Loreto: Flooded banks of Rio Aguaytia, Tessmann 3160, type.
Flora of Peru 435
9. VALLEA Mutis
Reference: F. Ballard, Bot. Mag. 157: pi. 9S65. July 2, 1934.
Slender branched shrubs or trees, at least some branches with
conspicuous reniform stipules, especially on sterile shoots, and
usually ovate, frequently cordate leaves. Flowers on axillary or
terminal peduncles, the 5 (rarely 4) sepals valvate, the 3-lobed
petals imbricate. Stamens indefinite, in 2 series, the disk biannulate,
the linear erect anthers apically dehiscing by 2 oblique pores.
Ovary 3-5-celled, the cells medially 2-ovuled; style subulate, the
3-5 filiform stigmas uneven. Fruits finally subligneous, bluntly and
camosely tubercled, tardily dehiscing, imperfectly 3-5-valved. —
Commemorates Felix Valle, author of "Florula Corsicae" (1762).
Vallea stipularis L.f. Suppl. 266. 1781. V. cordifolia R. & P.
Syst. Veg. 132. 1798. V. pyrifolia Turcz. Bull. Soc. Nat. Mosc. 31,
pt. 1: 237. 1858. V. pubescens HBK., Nov. Gen. & Sp. 5: 350. 1823.
V. glabra Miers, Contr. Bot. 2: 184. 1860-1869.
Attractive small tree, several to 6 meters or so tall, bearing
abundant leaves, bright green above, glaucous beneath, pretty
roseate to dark or bright red flowers gracefully borne in loose cymes
near or at the tips of the many brown-purple branchlets, and early
dark green somewhat berry-like fruits covered with soft processes,
often all on the same branches at the same time; indument, if present,
an unevenly deciduous tomentum of brownish trichomes mostly on
the peduncles and pedicels and leaves beneath or now and then
persisting in tufts in the nerve axils; stipules obsolete to conspicuous,
especially on young sterile shoots, reniform; leaves ovate-lanceolate
to broadly ovate, more or less acuminate or acute, rarely reniform
or rounded, cordate to subtruncate, usually 6-7 cm. long, 3-^ cm.
wide, but highly variable, the veins prominent; petals 3-lobed,
usually medially, 9-11 mm. long; stamens densely white pubescent,
the somewhat glandular filaments curved and filiform below the
greenish or yellowish anthers, both subequal; fruits subglobose,
about 1 cm. across, often dehiscing on the tree. — Bark of old trunks
fissured; leaves yellowish or reddish in the dry season, those of
young shoots sometimes crenate-lobulate. Var. pyrifolia (Turcz.)
Ballard, I.e., was proposed for glabrous specimens (except nerve-
axil pubescence); more striking at least is the variant designated
parvifolia by me, Candollea 5: 381. 1934, the leaves rotund-reniform,
rounded at apex, mostly 3-4 cm. wide, only 2.5-3.5 cm. long; how-
ever, as Diels has remarked, the conclusion of Ballard that the
436 Field Museum of Natural History — Botany, Vol. XIII
species is highly variable but not divisible, as unstable, is sound.
Illustrated, Bot. Mag. 9365; HBK., Nov. Gen. & Sp. 5: pi. A89;
Miers, Contr. Bot. 2: pi. 81 (V. pubescens).
Wood reported as valuable by Grisard and Vanden-Berghe, Rev.
Sci. Nat. Appl. 39: 587. 1892; West found a tea made from the leaves
used medicinally. Common, at least formerly, at middle altitudes,
especially from the central valleys, southward.
Piura, Cajamarca, Ancash: Raimondi. — San Martin: West of
Moyobamba, Weberbauer 266. — Hudnuco: Near Pampayacu, Sa-
wada; Kanehira. Near Huanuco, 2130; Weberbauer 1766. Pillao,
Woytkowski 3If091. Mito, lJl^79; i 7^7.— Junin: Near Tarma, Ruiz
& Pav6n (type, V. cordifolia); Woytkowski 35Jf.65. Huariaca, 3093;
Poeppig. Huacapistana, Killip & Smith 2^208 (type, var. parvi-
folia); Weberbauer 1766. Carpapata, Soukup 3Jt30 (va. parvifolia).
— Huancavelica: Salcabamba, Stork & Horton 10288. — Ayacucho:
Near Ocros, West 3672 (det. Johnson, V. cordifolia) . Near Huanta,
Killip & Smith 2331 If. — Apurimac: Abancay, Balls 6896. — Cuzco:
Wenner Gren Ruins, Metcalf 3076 Jf (det. Killip). Galea, Vargas
171. Ollantaytambo, Herrera 331,3; 3U0; 3U1; Cook & Gilbert
387; 631. Urubamba, Soukup 26; Weberbauer, 182.— Puno: Near
Limbani, Vargas 9638. Near Puno, Soukup ^^^. Oconeque,
Metcalf 30585. Tabina, Lechler 2083. Sandia, Weberbauer, 237.
Without locality (Mathews 30^8, det. Miers, V. pubescens); (Mathews
892, det. Miers, V. cordifolia). Bolivia; Ecuador and Colombia.
"Cugur," "cunhur" (Ruiz & Pavon), "gellccoy" (Stork & Horton),
"chchicllur" or "chchicllurmy" (Herrera), "tchillumay" (Cook &
Gilbert), "chillunmay" (Vargas), "sacha-capuli" (Spruce), "olla-
olla," "quellccoy" (both, Raimondi).
10. MUNTINGIA L.
Stellulate pubescent little tree with dentate leaves and white
flowers peduncled in the axils and usually 2 or 3 together, sometimes
solitary. Sepals free, commonly 5, early 6 or 7 as the broad smooth
petals, these approximate with the indefinite stamens about the
annulate disk. Ovary glandular-pilose, 5-7-celled, multi-ovulate;
stigma sessile, lobulate. PYuit indehiscent, globose, irregularly
many-celled, the numerous minute seeds pulp-imbedded. — See Lil-
loa 3: 31. and plates, Descole and O'Donell.
Bark furnishes a tough fiber suitable for basketry.
Flora of Peru 487
Muntingia Calabura L. Sp. PI. 509. 1753.
Shrub or sometimes a slender tree attaining 12 meters, with
many basally very oblique oblong-lanceolate leaves these densely
cinereous or brownish stellate beneath, puberulent or soon green
and glabrate above, commonly 5-12 cm. long, 2-3 cm. wide; petioles
2-6 mm. long, exceeded by the linear stipules; pedicels 1-3 cm. long;
sepals 8-12 mm. long, with nearly filiform acumen about equaling
the rarely rose-tinted broadly obovate petals; berry globose, red or
yellow, about 1 cm. in diameter, edible. — Branches divaricate, early
hirsute, in age glabrous and reddish brown. Illustrated, Mart. Fl.
Bras. 12, pt. 3: pi. SJ^.
Piura: Cafia Dulce, Haught 108; 197; Weberbauer, 150.— San
Martin: Tarapoto, Woytkowski 35162. Juanjui, Klug 3762. Vitoc,
Ruiz & Pavdn. — Amazonas: Along streams, Weberbauer, 155. —
Junin: La Merced, 5232. Colonia Peren^, Killip & Smith 25067;
25178. — Loreto: Rio Nanay, Williams 1^57. Lower Huallaga,
Williams ^931. Rio Pachitea, Sandemun 3308. Iquitos, Williams
8025; Mexia 6510, part. PongodeManseriche, Mexia 6169. Puerto
Arturo, Killip & Smith 27929. — Cuzco: Valle de Santa Ana, Herrera
S212; 3283. Santa Rosa, Soukup. Valle de Urubamba, Herrera
S220. To southern Mexico and the West Indies. "Bolina," "iuma-
nasa," "mullaca-huayo" (all, Williams), "yumanaza," "ccoillor-
ppanchu" (Herrera), "guinda yumanasa" (Mexia).
11. SLOANEA [Plum.] L.
Reference: C. Earle Smith, Jr., Contr. Gray Herb. 175: 3-114.
1954.
Trunks often buttressed. Leaves alternate or opposite, pin-
nately veined; stipules sometimes present only in bud. Inflores-
cences various but usually axillary and racemose or paniculate.
Sepals 4-11, ordinarily free. Petals none. Stamens about 50 to
more than a hundred, always with connective extended as a knob
or an awn. Pistil 3-6-celled, ovules 8-10 in 2 rows per cell, anatro-
pous, pendent, placentation axial. Fruit a loculicidally dehiscent
capsule with 3-6 rigid often ligneous valves, smooth or armed with
persisting or easily detached (and irritant) spines. Seeds 1-2 (3),
medially or nearly completely covered by a firmly attached aril
(except one species). — After Smith, who has given a detailed and
obviously well-considered and clearly presented monograph, from
which the following key and descriptions are compiled. The genus
is developed largely to the east and north of Peru but of course
438 Field Museum of Natural History— Botany, Vol. XIII
several species besides the following may be found within its north-
eastern boundary.
Sepals 4-11, unequal, not covering stamens and pistil in bud;
flowers in lateral racemes (subgenus Sloanea C. E, Smith).
Stipules caducous; flowers often crowded; capsules to 3.5 cm. long
(section Brevispicae C. E. Smith).
Leaves clustered apically.
Stamen awn elongate-filiform; capsules to 2 cm. long, spines
to 5 mm. thick S. rufa.
Stamen awn short, thick.
Capsules merely granulose; sepals 4 or 5 S. granulosa.
Capsules spinose; sepals usually 5 S. spathulata.
Leaves distributed evenly S. guianensis.
Stipules at least somewhat persistent; flowers borne laxly (section
Sloanea C. E. Smith) S. fragrans.
Sepals 4 (5), equal, covering stamens and pistil in bud; flowers (Peru)
not racemose (subgenus Quadrisepala C. E. Smith).
Flowers paniculate; capsule spines to 1 cm. long S. laxiflora.
Flowers 1-3, subumbellate; capsule spines short (2 nmi.).
S. temiflora.
Sloanea fragrans Rusby, Mem. N. Y. Bot. Gard. 7: 294.
1927; 68.
Stout twigs, rigid coriaceous obovate leaves and long (13-27 cm.)
inflorescences minutely puberulent to glabrate; stipules 2.5-13 cm.
long, obovate, obtuse, unevenly dentate; petioles terete, 5-20 cm.
long, alternate; leaves 4.5-6.5 dm. long or longer, 2-3 dm. wide,
acute at base, obtuse, undulate-margined; peduncles 1 dm. long,
pedicels to 2.5 cm. long, bracts ovate, 4-10 nun. long, unevenly to
few-dentate; flowers 1-1.5 cm. long, sepals 7-9, ovate to lanceolate,
obtuse, 4-6 mm. long, puberulent-sericeous on both sides; stamens
4-8 mm. long, anther awn flattened, to 1 mm. long, the awns in bud
imbricate, forming a globose mass; style not parted; capsule (young)
densely covered with curved spines. — A large tree.
Loreto: Pongo de Manseriche, mouth of the Rfo Santiago,
Tessmann Jt609. Bolivia to Colombia.
Sloanea granulosa Ducke, Bol. Tech. Inst. Agron. do Norte
19: 13. 1950; 45.
Flora of Peru 439
Related to and resembling S. rufa but glabrate or shortly pu-
bescent; stipules deltoid-lanceolate, to 2 mm. long; petioles 8-15
mm. long; leaves subacute to rounded at base, obtuse to emarginate
at apex, 9-14 cm. long, 4.3-9 cm. wide, sparsely short-pubescent
beneath and with scattered long trichomes along the mid-rib; in-
florescences 1.5-2 cm. long, 3-7-flowered, the peduncles 1-4 mm.
long, i)edicels 4-12 mm. long, bracts 1-2 mm. long, rarely few-
dentate; flowers 4-9 mm. long, sepals 1.5-2 mm. long, glabrous
within; stamens 1.5-2.5 mm. long, the broad anthers with a short
stout glabrous awn; style deeply 4-parted; capsules about 3.5 cm.
long, ovoid, the 4 valves 4-5 mm. thick, smooth and red inside,
granulose and densely fine-pubescent without; seed 17 mm. long,
covered to the funiculus with a fimbricate 5-lobed red aril. — The
only locality in Peru (if that of Pearce) is on the Rio Maran6n.
Becomes a very tall tree with large radiating buttresses (Ducke).
Peru(?): Monterico (Pearce). Amazonian Brazil.
Sloanea guianensis (Aublet) Benth. Joum. Linn. Soc. 5: suppl.
69. 1861; 33. Ablania guianensis Aublet, PI. Guian. 585. pi. 23U.
1775.
Stipules 0.5-1 mm. long, lanceolate, obtuse; petioles 1-3 mm.
long, opposite or subopposite, not clustered at twig-tip; leaves
obovate to elliptic-lanceolate, obtuse to cuneately acute at base,
obtuse to acuminately obtuse at apex, 5-22 cm. long, 2.5-9.5 cm.
wide, glabrous to minutely pubescent above, sometimes sparsely
hirsute and barbate in the prominent nerve axils beneath, entire to
obtusely dentate; inflorescences 1-9-flowered, 1.5-3 cm. long,
sparsely to densely fuscous pubescent, the peduncles to 1.5 cm. long,
pedicels 3-8 mm. long, bracts 5-15 mm. long, these deltoid to
obtusely lanceolate; flowers white to yellow, 3-8 mm. across, 5-8
mm. long; sepals 4-9, variable in size, shape and pubescence, some-
times glabrous within; stamens 2-4 mm. long, the. awn a fourth to
a third as long as the sublanceolate anthers; style 4-parted; capsule
9-12 mm. long, 6-9 mm. in diameter, the usually 4 valves velvety
and rather densely spiny antrorsely pubescent spines 5-9 nmi. long;
seed to 1 cm. long, almost enclosed by a 5-lobed aril. — Said to attain
25 meters and to develop triangular buttresses; valued for firewood
and paddles (Mexia). S. trichosHcha Williams & Sandw,, Flora
Trinidad & Tobago 1: 110. 1929, has smaller flowers with exserted
awnless stamens, hirtellous capsule spines and has been found in
adjacent Brazil (Rio Acre).
440 Field Museum of Natural History — Botany, Vol. XIII
Loreto: Pongo de Manseriche, Rio Maranon, Mexia 6137.
Along the Ucayali, Tessmann 3213. Brazil to Trinidad and Venezu-
ela. "Cutana-cuspi" (Mexia). '
Sloanea laxiflora Spruce ex Benth. Journ. Linn. Soc. 5: suppl.
65. 1861; 82. S. acutiflora Uitt. Recueil Trav. Bot. Ne^rl. 22: 357.
1925, fide Smith. S. polyantha Ducke, Archiv. Inst. Biol. Veg. Rio
Jan. 2: 162. 1935, fide Smith.
Glabrous except for the finely pubescent branchlets and often
terminal inflorescences, these 5-11 cm. long; petioles alternate, to
1.5 cm. long; leaves ovate to elliptic, rounded at base, acuminate-
acute to obtuse, 4.5-15 cm. long, 2.5-8 cm. wide, subcoriaceous,
veins prominent beneath, margin more or less entire; peduncle
3-5.5 cm. long, pedicels 5-10 mm. long; flowers white or yellowish,
6-10 mm. long, sepals apparently soon falling, puberulent both
sides; stamens 3-7 mm. long, anthers dehiscing apically, with a short
glabrous awn; style entire; capsules 2.5-3 cm. long, subglobose, the
4 valves 2-2.5 mm. thick, densely spinose; spines subrigid, 8-10
mm. long, puberulent; seed white, to 2.3 cm. long, with a 3 lobed
pale yellow aril (fruit character after Ducke). — A buttressed tree
observed as high as 15 meters.
Loreto: Iquitos (Ducke 1833). To the Guianas.
Sloanea rufa Planchon ex Benth. Journ. Linn. Soc. 5: suppl.
68. 1861; 43. S. longicaudata Ducke, Bol. Tech. Inst. Agron. do
Norte 19: 14. 1950, fide Smith.
Twigs stout, cinnamon pubescent; stipules subpersistent, ovate,
laciniate, 4-12 mm. long, densely pubescent; petioles to 7.5 cm. long,
rufous-tomentose; leaves elliptic to obovate, rounded to cordate
at base, acutely or obtusely short-acuminate, 11-32.5 cm. long,
5.5-27 cm. wide, coriaceous, densely pubescent medially above,
rufous-tomentose. beneath, the veins prominent, obscurely but
sometimes spinulosely repand-dentate; inflorescence 5-9-flowered,
1.5-3 cm. long, densely pubescent, the peduncles 1-2 cm. long,
pedicels 3-10 mm. long, bracts 1-3 mm. long, laciniate; flowers
5-7 mm. long, pale green or yellowish; sepals 2-3 mm. long, within
pubescent only toward margin; stamens 2-4 mm. long, the glabrous
filiform awn to 1.5 mm. long; style 4-parted; capsules probably to
2.5 cm. long, valves 4(?), to 1.5 mm. thick, with many semirigid
sparsely hispid clavate spines about 4 mm. long. — Small tree to
7 meters tall (Klug) resembling S. spathulata C. E. Smith and
Flora of Peru 441
S. granulosa Ducke in the apically crowded leaves but distinctive
in pubescence and stipules. S. Duckii C. E. Smith, I.e. 44, is dis-
tinguished by glabrous foliage and much smaller capsules (author) ;
found as near Peru as S5o Paulo de Olivenca, it probably extends
westward.
Loreto: Mishuyacu near Iquitos, Klug 5S. To French Guiana.
Sloanea spathulata C. E. Smith, Contr. Gray Herb. 175: 43.
1954.
Twigs sparsely pilose or glabrate, the leaves crowded toward the
tips; stipules lanceolate, to 5 mm. long, pilose; petioles stout, 3-9
mm. long, canaliculate above; leaves obovate, abruptly obtuse at
base, mucronulately subacute, 12-33.5 cm. long, 4.5-nearly 14 cm.
wide, chartaceous, sparsely pilose beneath, the secondary nerves
very prominent, undulate or repand-dentate toward apex; in-
florescences immature, to 1 cm. long, pubescent, the peduncles
4 mm. long, pedicels 3-4 mm. long, bracts 0.5-1 mm. long, entire,
obtuse; flowers 3-4 mm. long; sepals mostly 5; anthers longitudinally
dehiscent, shortly apiculate; style 4-parted apically; capsules 2-2.5
cm. long, the usually 4 valves 2.5-3.5 mm. thick, the many sparsely
tomentose spines 6-8 mm. long, slightly curved. — Differs from
S. rufa Planchon in the narrow leaves with very short petioles and
in the short anther awn (author) ; probably occurs in adjacent Peru.
Type was a tree 18 meters high.
Rio Acre: near mouth of Rio Macauhan, Krukoff 53S7, type;
also 5S9S.
Sloanea terniflora (Moc. & Sess^) Standi. Trop. Woods 79: 10.
1944; 94. Lecostemon terniflorum Moc. & Sess^ ex DC. Prodr. 2:
639. 1825. S. (ruadrivalvis Seem. Bot. Voy. Herald 85. pi. 15. 1853.
Unique among Peruvian species in the umbellately 1-3-flowered
inflorescences, these 3.5-5 cm. long; indument on all parts a puberu-
lence or lacking; petioles alternate to opposite, 3-20 mm. long;
leaves subovate, elliptic or obovate, subcordate, rounded or cune-
ately obtuse at base, rounded to obtuse at apex, 6.5-15 cm. long,
3-7.5 cm. wide, chartaceous to coriaceous, entire to unevenly
repand-dentate; peduncles 1.5-3 cm. long, pedicels 8-13 mm. long;
flowers maroon, 5-7 mm. long; sepals ovate, puberulent on both
sides; stamens 3.5-4 mm. long, anthers apically dehiscent, the
puberulent awn 0.5-1 mm. long; style entire; capsule ellipsoid,
1.5-2.5 cm. long, 1-2-seeded, the 4 valves 1-3 mm. thick, densely
442 Field Museum of Natural History— Botany, Vol. XIII
covered with easily detached spines, these about 2 mm. long; seed
1-1.2 cm. long, nearly covered with irregularly margined aril. —
Buttressed tree (known to 30 meters high).
Loreto: above Iquitos, Mexia 6510. To Mexico and Brazil.
"Anallocaspi" (Mexia).
MALVACEAE. Mallow Family
Herbs or shrubs, rarely arborescent, the indument usually
stellate or lepidote, the stems often fibrous. Leaves alternate, entire
or variously lobed, the stipules free, except Nototriche. Flowers
hermaphrodite, rarely dioeceous or polygamous, often bracteolately
involucrate. Sepals (4) 5, more or less connate, valvate or the calyx
rarely truncate. Petals 5, distinct but often adnate at base to the
staminal column, contorted or imbricate. Stamens many (or 5 or
10), hypogynous, monadelphous, the anthers 1-celled. Ovary free,
2- or more-celled, the fruit usually consisting of 4-many carpels
around a central axis or rarely of 1 carpel, or the carpels seriate
vertically; style sometimes completely connate, as many or twice
as many as the carpels; ovules 1 or more, from the inner angle of
each cell. — The fruit is usually dry, separating into cocci, these
indehiscent, or bivalved, or the fruit rarely capsular.
Here belongs of course cotton, at least one species native to Peru.
And the family is scarcely less known for two beautiful plants widely
grown for ornament, hibiscus (H. rosa-sinensis L.) in the tropics
and subtropics, and mallows or hollyhocks (Althaea rosea Cav.) in
temperate regions. The alpine mallows of Peru, the Nototriches,
are among the most beautiful of high Andean flowers; the plants
often form little gray-green cushions in loose soils and rocks, usually
below melting snow, and in flower many varieties are colorful with
their small mallow-type blossoms in delicate or bright blues and
violets, scarlet or crimson, sometimes white.
The wording of this compilation of the Peruvian mallows is mine
since the well-known student of the group, B.P.G. Hochreutiner,
was unable to contribute it as long planned. However, he has been
kind enough to read it for errors and omissions, reminiscent of his
co-operation and encouragement during my Geneva sojourn. Most
of his many useful suggestions have been noted in the text by his
name or his initials. The technical key is naturally in part tra-
ditional but also in part after Thomas H. Kearney, Am. Midi. Nat.
46: 98-105. 1951, and I acknowledge my indebtedness and ap-
Flora of Peru 448
preciation also for his useful "The American Genera of Malvaceae,"
I.e., which contains references and critical notes detailing the basic
characteristics of most American genera, especially those whose
relationships are still incompletely understood; it has been my
endeavor to pass on accurately Kearney's observations and those
drawn by him from other students, notably from Hochreutiner's
great contributions to an understanding of the fruiting characters
of the family in Ann. Cons. Jard. Bot. Geneve 21: 347-387. 1920,
which is summarized lucidly; cf. Hochreutiner, Flora Madagascar.
Besides the technical key, one devised largely on characters other
than those of fruits for the determination of perhaps most at least of
the Peruvian species has been appended. The present taxonomy
of the group does not permit this to be entirely definitive; probably
someone will simplify this in due time, letting sectional names serve
for present generic ones maintained on relatively developed char-
acters of carpels and carpel-dehiscence. Of course, as in better
organized families, concomitance of characters should be stressed,
not differences largely, but similarities.
Omitted from the key is Lavatera assurgentiflora Kellogg, of the
Santa Barbara Islands; it was found cultivated at Huancayo by
Luis A. Chavez; in Puno by J. Soukup; it resembles Hibiscus, but
has a broadly 2-3-lobed involucel. L. arhorea L., with villous
inflorescences, was purchased in the markets of Lima (Cook &
Gilbert 2076), as "Malva real or comun" and, while similar to the
former, has deltoid-ovate bractlets longer than calyx lobes, erect
instead of recurved-assurgent pedicels, and violet instead of roseate
flowers.
No single "Flora" has contributed more to the preparation of
this one than the scholarly example. Flora of Jamaica, by Fawcett
and Rendle and it is fitting for me to give it credit here particularly
since so many Malvaceae occurring in Peru are also found in Ja-
maica.
Key (Peruvian Species)
Carpels in 2 or more superposed verticels (unless one species), at
maturity completely separate, indehiscent; herbs with solitary
flowers, no involucel 1. Palaua.
Carpels commonly as many or half as many as the style branches,
uniseriate; genera never completely as above.
Style branches normally 10, twice as many as the indehiscent
(or tardily) carpels; stigmas capitate or discoid.
444 Field Museum of Natural History — Botany, Vol. XIII
Petals auriculate on one side below; fruit, until dry, berry-like
with fleshy envelope 2. Malvaviscus.
Petals unappendaged; fruit not berry-like, even early, dry.
Inflorescence foliaceous-bracted 3. Malachra.
Inflorescence not foliaceous-involucrate.
Leaf midnerve beneath with gland near base; carpels
glochidiate 4. Urena.
Leaf nerves glandless; carpels sometimes aristate or
muricate 5. Pavonia.
Style branches and carpels same number or style unbranched.
Carpels never free, the fruit capsular; stamen tube rarely
filamentous at apex.
Style branches somewhat divergent, stigmas more or less
capitate or discoid; seeds usually reniform.
Calyx regularly 5-dentate or -lobed, persisting.
6. Hibiscus.
Cal3rx spathaceous, unevenly 2-3-lobed, deciduous.
7. Abelmoschus.
Style branches short, erect or style unbranched, clavate;
seeds usually angulate or obovoid; calyx with black
glands.
Bractlets small, more than 3, or wanting; calyx 5-cleft.
8. Cienfuegosia.
Bractlets 3, foliaceous or caducous; calyx dentate or
subentire 9. Gossypium.
Carpels finally free, the fruit a schizocarp unless Abutilon;
stamen tube filamentous at and often below apex.
Stigmas decurrent on the slender pointed style branches.
10. Malva.
Stigmas apical or nearly, distinctly larger than styles (except
a few Chilean species of Abutilon).
Ovules normally 2 or more in each carpel.
Involucel wanting.
Carpels more or less completely divided (by a fold of
the lateral walls) into 2 superposed cavities.
11. Wissadula.
Carpels 2-celled by a horizontal projection (endo-
glossum) of the dorsal wall .... 12. Pseudabutilon.
Flora of Peru 445
Carpels 1-celled, the cavity undivided ... 13. Abuiilon.
Involucel present; herbs 14. Modiola.
Ovule normally solitary (except Sphaeralcea, and solitary
in the Peruvian species).
Involucel present (sometimes obsolete or caducous in
Malvastrum, Sphaeralcea) ; ovule erect or ascending.
Carpel cavity more or less divided by transverse
septum, a horizontal projection of the dorsal
wall 14. Modiola.
Carpel cavity not divided or only slightly (one genus,
sens. lat.).
Carpels clearly diverse apically and basally, reticu-
late, lower part indehiscent. ... 15. Sphaeralcea.
Carpels not clearly diversely segmented, usually
not reticulate 16. Malvastrum.
Involucel none; (rarely present in Nototriche annuals)
also see Malvastrum, Sphaeralcea.
Carpels much inflated.
Carpel cavity entirely open 13. Abuiilon.
Carpel cavity with an organ maintaining the seed
inside the dehiscent carpel 18. Gaya.
Carpels little or not at all inflated.
Carpel bases separated from carpel body, uniting
into a cup, apices often alate 19. Cristaria.
Carpel bases not cupulate nor apices alate, unless
in Sida (rarely).
Carpels typically with diverse indehiscent basal
section 15. Sphaeralcea.
Carpel cavity partly to completely divided (by
fold of lateral walls) but dehiscence equal.
11. Wissadula.
Carpel cavity entirely open.
Stipules (rarely none) and petioles united;
ovule erect-ascending; stems, except few
annuals, polsterform 17. Nototriche.
Stipules free; ovule pendulous or horizontal.
Carpels loculicidal nearly to base; calyx
lobes 4 23. Tetrasida.
446 Field Museum of Natural History — Botany, Vol. XIII
Carpels septicidal, often septicidal and locu-
licidal, separating; calyx lobes 5.
Lateral carpel walls firm 20, Sida.
Lateral carpel walls fragile 21. Anoda.
Carpels loculicidal, persisting; calyx lobes 5.
22. Bastardia.
Vegetative Key (only for Peruvian Species)
Flowers subtended by large foliose bracts.
Bracts laciniate, exceeded by the showy flowers. . . .9. Gossypium.
Bracts entire, conspicuous, concealing the small flowers.
3. Malachra.
Flowers often with an involucel of more or less developed bractlets
but not notably foliose-bracted.
Involucel of 3-many distinct or connate bractlets present at calyx
base (sometimes deciduous, rarely caducous as in Malvastrum;
see also annual Nototriche).
Flowers showy, several cm. long.
Corolla funnelform, petals with 1 auricle on claw; fruit
berry-like 2. Malvaviscus.
Oorolla more spreading (unless rarely in Hibiscus); petals
not auricled nor fruit berry-like.
Calyx deeply 5-lobed, gland-dotted 8. Cienfuegosia.
Calyx shortly 5-dentate or -lobed or 2-3-lobed.
Calyx persisting, the teeth or lobes regular ... 6. Hibiscus.
Calyx deciduous, 2-3-lobed, spathe-like. .7. Abelmoschus.
Flowers small, at most a cm. or two long, rarely somewhat
longer (see one or two species of Hibiscus.)
Leaves with 1 or more glands on nerves beneath; bractlets
connate 4. Urena.
Leaves without nerve-glands; bractlets distinct.
Herbaceous annuals or biennials.
Style branches slender, introrsely stigmatic; plants often
low, spreading 10. Malva.
Style branches with apical or subapical stigmas.
Flowers solitary; carpels transversely subseptate.
14. Modiola.
Flora of Peru 447
Flowers not solitary, often racemose; carpels with
2 sections or slightly if at all septate.
Carpels (typically) with 2 diverse parts.
15. Sphaeralcea.
Carpels not clearly diversified 16. Malvastrum.
Suffrutescent or shrubby, sometimes acaulescent.
Style branches twice as many as carpels; leaves green or
greenish, longer than wide or flowers few, pedicellate;
inflorescences short, few-flowered or paniculate.
5. Pavonia.
Style branches and carpels same number; flowers race-
mose or somewhat congested; leaves about as wide
as long (HibisciLS hrasiliensis might be sought here).
16. Malvastrum.
Involucel none (specimens with only mature calyces of group
with involucel only into anthesis [as Malvastrum sp.] might be
sought here).
Flowers solitary, adnate as stipules to petioles; depressed alpine
perennials, rarely annuals 17. Nototriche.
Flowers, stipules and petioles not jointly adnate.
Styles as many as carpels, the latter in 2 or more superposed
verticels except one species; flowers axillary, solitary.
1. Palavxi.
Styles or branches as many as the uniseriate carpels.
Leaves entire or essentially, except Anoda, sometimes
obscurely or minutely crenulate, rarely serrate and
hastately lobed, never linear.
Flowers solitary on axillary peduncles; leaves often
hastately lobed 21. Anoda.
Flowers not solitary nor leaves lobed.
Calyx 4-lobed; carpels basally loculicidal; leaves soon
glabrous above 23. Tetrasida.
Calyx 5-lobed; leaves glabra te or pubescent on both
sides.
Petals white, medially adnate to stamen tube;
leaves green; tree 20. Sida.
Petals usually yellow, only basally adnate; leaves
canescent beneath; herbs or shrubs.
11. Wissadvia. 13. Abutilon.
448 Field Museum of Natural History— Botany, Vol. XIII
Leaves more or less distinctly serrate, rarely entire and
linear, sometimes palmately lobed or parted, not
hastately leaved annuals.
Leaves more or less palmately lobed or parted.
Carpel bases dilated, forming a cup; flowers long-
pedicellate 19. Cristaria.
Carpel bases cupulate, not dilated; flowers sessile or
pedicellate 20. Sida.
Leaves serrate or rarely entire, then linear.
Carpels much inflated; leaves usually canescent at
least beneath.
Carpels 1-celled 13. Abutilon.
Carpels partly divided 18. Gaya.
Carpels not or little inflated; leaves often green or fruit
capsular (Bastardia).
Carpels normally 1-seeded.
Indument glandular; carpels persisting, locu-
licidally dehiscent 22. Bastardia.
Indument eglandular; carpels separating, sep-
ticidally or septicidally and loculicidally
dehiscent 20. Sida.
Carpels normally 2-several-seeded.
Flowers rarely 1 cm. long; seeds 2 or 3 in capsule
divided by a fold or a dorsal projection or
a lateral fold.
12. Pseudabutilon. 11. Wissadula.
Flowers rarely as short as 1 cm.; seeds usually
several, in a single-celled capsule.
13. Abutilon.
1. PALAUA Cav.
Reference: Hochreutiner, Ann. Cons. Jard. Bot. Geneve 5:
170-173, 1901.
Tomentose or glabrate herbs, rarely suffrutescent, with axillary
peduncled solitary ebracteolate flowers and usually sinuately lobulate
or dissected leaves. Anthers clustered at apex of filament tube.
Ovary cells many, 1-ovuled, the filiform style branches same number,
enlarged apically and vertically stigmatose. Carpels at maturity
inordinately capitate-congested in 2 or more superposed verticals
Flora of Peru 449
(except in P. Guentheri), parting from the receptacle, indehiscent,
the seed ascending. — Genus name changed to Palava and Palavia
by later authors but honors Antonio Palau y Verdera, early botanist
of Madrid. See also Ulbrich, Bot. Jahrb. 42: 104-113. 1908 for
a detailed revision.
Perennials, suflfrutescent; leaves densely tomentose, obscurely or
not lobulate.
Leaves crowded, ovate-cordate, nerves impressed above, rugose
beneath P. moschata.
Leaves soon openly borne, ovate or elliptic, plane .... P. vdutina.
Annuals; leaves, at least cauline, rarely undivided, then lightly
pubescent.
Leaves all undivided or the upper more or less obscurely lobulate.
Petals little if at all exceeding the calyx.
Leaves green, the indument sparse P. malvifolia.
Leaves densely stellulate, especially beneath . . P. inconspicua.
Petals much longer than the calyx.
Leaves green, lightly stellulate only beneath in age.
P. rhomhifolia.
Leaves cinereous, greenish-gray even above only in age.
P. tomentosa.
Leaves more or less divided or subbasally trifid and lobulate un-
less the early basal.
Calyx lobes ovate; leaves dissected or trifid, at least upper.
Upper leaves trifid, the lobes lobulate P. Weberbaiieri.
Upper as lower cauline leaves more or less dissected.
P. dissecta.
Calyx lobes linear; leaves unequally pinnate or subentire.
P. Gueniheri.
Palaua dissecta Benth. Joum. Linn. Soc. Bot. 6: 101. 1862;
173. P. fiexuosa Mast. Card. Chron. 435. 1866. P. mollendoensis
Ulbr., P. geranioides Ulbr., P. pusilla Ulbr., Bot. Jahrb. 42: 108,
111, 112. 1908.
Decumbent ascending or early subacaulescent and suberect,
lightly (unless calyces) stellulate-hispidulous or subtomentose, the
cauline leaves palmately dissected, the lobes themselves more or
less and unevenly pinnate, 3-5-lobed or rarely subentire, the crenate
basal oblong-ovate, the dissected upper more deltoid-ovate, variable
450 Field Museum op Natural History— Botany, Vol. XIII
also in size, ordinarily 2.5-5 cm. long and somewhat narrower;
stipules subulate to 7 mm. long; peduncles slightly or early much
exceeding the leaves; calyx 7 mm. long, hispid-stellate, the broadly
ovate lobes acute to acuminate, 4 mm. long, not glandular-dotted;
petals (1) 2 cm. long or longer, mauve to magenta or white below;
anthers bright red; carpels obovoid, rugose reticulate. — Partly after
Hooker and Masters, the former under Bot. Mag. pi. 5768. The
type with much-dissected leaves was mixed with P. rhombifolia
Grab, with entire leaves; the purplish petals only 1 cm. long, in
Chile (Johnston). Flowers deep magenta, the centers white (Worth
& Morrison). Pennell 13329, referred by Ulbrich to his P. pusilla,
as "t3rpical but only 3 cm. high, unbranched, with few leaves."
P. mollendoensis is apparently merely a robust (certainly annual)
specimen, larger in all parts; Guenther & Buchtien specimen consists
of young plants with subentire leaves on the lower parts of the stem
(Ulbrich). P. geranioides apparently has slightly larger stipules
but nothing else to distinguish it. The carpels have been described
by Bruns on the basis of Guenther & Buchtien 229 as nearly 30,
brownish, tetragonous, laterally applanate, scarcely convex, gibbous-
verrucose, the elliptic seed with some long trichomes in the region
of the hilum. F.M. Negs. 9255 (P. geranioides); 9256 (P. mollen-
doensis); 9257 (P. pusilla).
Lima: San Lorenzo, Maclean, type. Without locality, Cuming.
— Arequipa: Posco, Vargas 2015; Cook & Gilbert 53 (det. Ulbrich);
{Guenther & Buchtien 212, det. Ulbrich). Lomas near Mollendo,
600 meters, Weberhauer 158Jt. (type, P. mollendoensis); Weherhauer
1^67 (tjrpe, P. geranioides); Weherhauer 1573 (det. Ulbrich); Mexia
05165; Worth & Morrison 15755; West 8199 (all det. Johnston).
East of Islay (Worth & Morrison 15711, det. Johnston). Mejia,
40 meters (Guenther & Buchtien 229, det. Ulbrich, P. geranioides).
Mejia to Chalascapa, (Guenther & Buchtien 219; 220, det. Ulbrich,
P. mollendoensis). Open gravelly places, Pampa de Arrieros, 3,750
meters, Pennell 13329. Chile. "Corilla" (Mexia).
Palaua Guentheri Bruns, Mitt. Inst. Allgem. Bot. Hamb. 8:
56, pi. 8. 1929. Malvastrum mollendoense Ulbr. Bot. Jahrb. 42: 120.
1908. P. mollendoensis (Ulbr.) Johnst. Journ. Arnold. Arb. 19: 260.
1938, not Ulbrich, 1908.
More or less ashy stellate-tomentose annual with usually several
spreading slender stems, the unequally pinnate leaves and flowers
crowded at the nodes and tips; leaves with 1-2 (type) or 3-5 (7)
Flora of Peru 451
narrowly linear-lanceolate segments 1-1.5 mm. wide; peduncles
1-3 cm. long; calyx 8 (-12) mm. long, parted nearly to the base,
tomentose especially below, the linear lobes strongly 1-nerved;
petals blue or pale rose-color, little longer than calyx, obtuse, ciliate
toward base; stamen tube 5-7 mm. long; styles 3-6 mm. long,
9-12-lobed, filaments 2-2.5 mm. long; carpels 9-12, convex, sub-
globose, glabrous. — Undoubtedly a species of Palaua (Johnston);
not a good Palaua, the carpels being uniseriate although somewhat
unequal in length (young) ; its most striking feature is the high and
narrow deeply lobed somewhat urceolate calyx, the slender lobes
marked by a dark central stripe (Kearney). It is a heresy to have
a plant with uniseriate carpels in the genus Palaua (B.P.G.H.);
and, finally, Krapovickas, Bol. Soc. Argent. Bot. 4: 187. 1952, has
studied material that confirms the original diagnosis and the ob-
servation of Johnston that the carpels actually are in two incom-
pletely superimposed series (Leafl. West. Bot. 6: 168. 1952). F.M.
Negs. 20924; 9319 (M. mollendoense) .
Arequipa: Prov. Islay, sand dunes, Mexia 7777 (det. Johnston);
Vargas 8ItS2. Mejia, lomas, Guenther & Buchtien 191, type. Near
Mollendo, Weberhauer 151^ (type, M. mollendoense)', Mexia JtlSO,
Palaua inconspicua Johnst. Contr. Gray Herb. 85: 151. 1929.
Herbaceous annual; stems erect or more or less decumbent,
solitary or many, 1-2 dm. long, slender, simple, rarely shortly few
branched, with minute stellate pubescence; leaves scarcely rosulate,
upper reduced, all orbicular-reniform-cordate, obtuse, 1-3.5 cm.
long and wide, more or less obscurely 3-5-lobate, minutely stellulate
above, paler and densely stellate beneath, the 5 palmate nerves
prominent; petioles slender, stellate; stipules subulate, persisting,
ciliate, 2-5 mm. long; peduncles slender, 1-3 cm. long, 1-5-flowered,
the articulate part 1-4 mm. long; calyx campanulate, 2-3 mm. long,
stellate, the lobes ovate; petals white, 2-3 mm. long, scarcely longer
than the calyx which conceals the depressed globose fruit, this 3-4
mm. broad; carpels 20-25, 1-seeded, 0.9 mm. long, compressed
ovoid, glabrous, rugose, the seeds 0.7 mm. long. — Differs from
P. modesta (Phil.) Reiche in distinctly annual, shorter, less con-
spicuously spreading trichomes, smaller corollas (Johnston). Not
seen (as indicated of course by the parenthesis of the collector's
name) but the description of fruit as "depressed" has caused Hoch-
reutiner to query: Is it sure that this is pluriverticillate with super-
posed verticils?
452 Field Museum of Natural History— Botany, Vol. XIII
Arequipa: Lower edge of fertile belt, Mollendo, (Johnston 3565,
type). Chile.
Palaua malvifolia Cav. Diss. 1: 40. pi 11. 1785; 172. P.
parviflora (L'H^r.) Hochr. Ann. Cons. Jard. Bot. Geneve 5: 173. 1901.
Malope parviflora L'H^r. Stirp. Nov. 1: 105, pi 50. 1789. P.
declinata Moench. Meth. 609. 1794. P. micrantha Ulbr. Bot. Jahrb.
54: Beibl. 117: 48. 1916, fide Johnston.
Prostrate annual, simple or well-developed plants with many-
stems, these rather remotely branched, the indument sparse or
nearly lacking, minutely stellulate except on the 1.5-2 mm. long
narrow stipules; leaves broadly ovate to subrotund, sinuate to
obscurely lobulate, variable in size, often 2-3 cm. long; peduncles
to 2.5 cm. long, usually shorter; flowers red; calyx 2-3 mm. long, the
lobes broadly ovate; petals suborbicular, 3-4 mm. long, the stamen
tube about half as long; carpels many, glabrous, rugulose, scarcely
1.5 mm. long, the seeds glabrous. — P. micrantha var. hirsuta Ulbr.
Notizbl. Bot. Gart. Berlin 11: 517. 1932 has white flowers and
larger leaves, these sparsely hirsute both sides, the petioles and
branches more densely. Ulbrich confused this with P. rhomhifolia
Graham, in spite of the revision of Hochr. I.e. 170-173. As pointed
out by Otto Kuntze, Rev. Gen. 3: 157. 1898, Fries and others
(cf. Sida palmata) the correct date of L'H^ritier's work (above page
and plate) is 1789. Dombey from the Lima sands is the type for
both names. My specimens were from the dryer seaward slopes
of sandy lomas. Hochreutiner restricted the name of L'H^ritier
to the type, the leaves subcuneate at base, but has indicated to me
he now feels both names apply to the same species. F.M. Negs.
9495; 23760.
Lima: Lomas de Chancay, Raimondi (det. Ulbrich). Barranca
near Lima, Weherhauer 1600; 1606; 5692 (type, P. micrantha).
Ruiz & Pavdn; Dombey. Lurin, 5939 (det. Johnston). — Arequipa:
Atiquipa, Raimondi (det. Ulbrich). Near Mejia, Loma formation,
200 meters (Guenther & Buchtien 198, type, var.). Chile.
Palaua moschata Cav. Diss. 1: 41, pi 11. 1785. P. lomageiton
Ulbr. Notizbl. Bot. Gart. Berlin 11: 516. 1932.
Sprawling, often forming patches, the defoliate older lignescent
stems and branches densely clothed with stipular and petiolar
remains; stipules lanceolate; petioles 1-8 cm. long; leaves many,
crowded at the branchlet tips, cordate-ovate, obtuse or rounded at
Flora of Peru 453
apex, crenate or crenate-serrate, sometimes obscurely lobulate,
2-5.5 (8) cm. long, nearly or quite as wide, densely tomentose on
both sides, more or less bullate above by the impressed veins,
rugulose beneath by the prominent nerves and veins; calyx tomen-
tose, about medially lobed, angled, 1-1.5 cm. long, the lobes acute;
p)etals reddish-lilac, obovate, obtuse or truncate, 1.5-2 (2.5) cm.
long; stamen tube 1 cm. long, little exceeded by the styles; fruit
enclosed in calyx, the carpels reniform. — Var. macrantha R.E. Fr.
Sv. Vet. Akad. Handl. ser. 3. 24, no. 2: 3. 1947 has larger leaves,
strongly cordate sepals 13-16 mm. long, 7-9 mm. wide, corolla to
2.5 cm. long, scarcely a genetic variant. P. lomageiton, unknown to
the lomas, with much larger leaves and flowers is allied to P. veliUina
Hill & Ulbr. of the lomas with different leaves, larger flowers (Ul-
brich). Entire plant is said to be strongly musk-scented. My
specimens were from loose stony upper slopes of seaside hills,
Murphy's from sheltered hollows among summit rocks. P.M.
Neg. 29772.
Lima: Huara, Ruiz & Pavdn; sands, Dombey, type. San Lorenzo,
Gaudichaud; Mathews 1010 (type, var. macrantha). Chorillos, 5872;
Weberbauer IW, 146. Callao, Andersson; Wilkes' Exped. San Gallen,
Murphy 3j^76 (det. Johnston). — lea: Lomas, 500 meters, Bahia de
la Independencia, Weberbauer 7961, (type, P. lomageiton). Chile.
Palaua rhombifolia Graham, Edinb. New Phil. Joum. 369.
1830.
Sprawling-ascending annual, the weak stems, slender petioles,
these 1-3 cm. long, and leaves beneath lightly hispid-stellate, the
latter as the carpels without soon glabrous or nearly so; leaves
alike but reduced above, ovate-rotund or -elliptic, often about 2 cm.
wide and long, sometimes three times as large, somewhat irregularly
crenate-lobulate; peduncles 3-4 cm. long, densely hirsute; calyx
about 8 mm. long, the broadly ovate lobes acute, canescent puberu-
lent within; petals about 12-18 mm. long, or longer, bright rose-red,
the narrow claws densely ciliate; fruits about 7 mm. across. —
Illustrated, Bot. Reg. pi. 1375; also Bot. Mag. pi. SIOO. F.M. Negs.
23761; 23762.
Ancash: Lomas de Mong6n, Goodspeed Exped. 9175 (det. John-
ston). — Lima: Near Lima, Cruckshanks, type. Sandy lomas along
the sea, Lurfn, 5921 (det. Johnston). Anc6n, Grant 7^69. Loma
Pasomayo, Stork & Vargas 9355 (det. Johnston).
454 Field Museum of Natural History— Botany, Vol. XIII
Palaua tomentosa Hochr. Ann. Cons. Jard. Bot. Geneve 5:
171. 1901.
Spreading-ascending more or less stellate-tomentose annual, the
broadly ovate or subrotund leaves less densely so above; stipules
subulate, slightly ciliate; petioles (1) 5-9 cm. long, the longer basal
especially sparsely long-pilose; leaves cordulate or subtruncate at
base, rather obscurely and unevenly 3-5-lobulate and coarsely
dentate the larger teeth mucronulate, the younger tomentose both
sides, the older greenish above, all 7-9-nerved, about 5 cm. long and
wide or the basal somewhat larger, progressively smaller toward the
tops of the elongating stems; i)eduncles 3.5-5 cm. long, articulate
above the middle; buds subglobose; calyx 8-9 mm. long, reddish
tomentose below, the subcordate ovate acute lobes 5-6 mm. long,
3-4 mm. wide near the base, sparsely setose within; petals obovate,
typically 1.5-2 cm. long, lilac or pink, paler below; stamen tube as
styles 8 mm. long; carpels many. — Stamens deep red, petals pink
to lavender, white in center (Worth & Morrison). This plant could
not be Mexican, as the type label indicates. F.M. Neg. 23763.
Arequipa: Loma near Atiquipa, Worth & Morrison 15672.
Sandy quebradas, Atiquipa, Worth & Morrison 15629. — Moquehua:
Torata, Weberbauer 7U20? (flowers smaller). Without locality,
Pavdn, (type. Herb. Geneve).
Palaua velutina Ulbr. & Hill, Bot. Jahrb. 42: 108. 1908.
Habit of the generally similar P. moschata but as to type much
more open in growth, the slenderer stems and branches with fewer
irregularly crenate and lobulate ovate-elliptic leaves 2-4.5 cm. long,
1-2.5 mm. wide, the close indument velutinous; flowers purplish,
2.5-3 cm. in diameter; calyx cupulate, 12 mm. long, the lobes ovate-
deltoid, acute, 6-7 mm. long; petals obovate, claw ciliate, 1.5-2
cm. long, 8-10 mm. wide; stamen tube 6-7 mm. long; carpels many,
2 mm. long, rugulose only dorsally. — F.M. Neg. 9258.
Arequipa: Loma sands (Guenther & Buchtien 192; and between
rocks, 192a, both det. Ulbrich). Lomas near Mollendo, Weberbauer
H93. — Tacna: Near Lacumba, 1,500 meters, Woitschach, type.
Palaua Weberbauer! Ulbr. Bot. Jahrb. 42: 110. 1908.
Annual, simple or laxly branched, stellulate pubescent including
the calyces, only the subulate-lanceolate persisting stipules (4-5
mm. long) pilose with simple trichomes; petioles 2-3 (5) cm. long;
basal leaves rotund obovate or suborbicular, irregularly coarsely
crenate or sublobulate-crenate, the upper leaves rotund-ovate.
Flora of Peru 455
3-5-lobed the larger terminal lobe obovate, the lateral obliquely
ovate, all coarsely and unevenly crenate-lobulate, all forms variable
in size, 2-4.5 cm. long, 1.5-4.5 cm. wide, cordate to subtruncate
at base, rather densely stellulate, the nerves prominent beneath;
flowers subumbellate in the upper axils on pedicels 2-6 cm. long;
calyx cupulate, 5-6 mm. long, the lobes obovate-acuminate, 4 mm.
long and wide above the base; petals about 1 cm. long, nearly as
wide, red or purple, ciliate and inconspicuously glandular at base;
stamen tube 6-7 mm. long, glabrous, as the styles and ovary. —
P. rhombifolia var. acaulis Hochr. seems to be a young plant. This
probably should be included in P. tomentosa. F.M. Neg. 9259.
Arequipa: In Lomas near Mollendo, Weberbauer H70, tjrpe;
Hitchcock 22^20; Mexia 0U168 (det. Johnston, P. dissecta); (Giinther
& Buchtien 227, det. Ulbrich). Mejia Giinther & Buchtien 228.
Pasco, 575 meters. Cook & Gilbert 50 (det. Ulbrich); {Giinther &
Buchtien 226, det. Ulbrich). — Moquehua: Torata, Weberbauer 7Jt20;
7It20A? Without locality, Mathews 912. "Gorilla" (Mexia).
2. MALVAVISCUS [Dill.] Adans.
Reference: Schery, Ann. Mo. Bot. Gard. 29: 183-244. 1942.
Shrubs, sometimes arborescent, the solitary axillary or terminal
or subterminal flowers usually bright red, the corolla funnelform,
the petals connivent (or spreading only apically) and auriculate on
one side of the claw, the stamen column usually long-exserted.
Involucel present. Stigmas apical. Ovules solitary, ascending.
Fruit baccate, the 5 uniseriate indehiscent carpels enclosed in a fleshy
envelope until full maturity but this character is not apparent in
old fruits (Schery). — Schery considers the Peruvian shrub as con-
stituting one species as indicated below in the sjaionjnny; this may
be the correct interpretation but types (unfortunately) have not
been designated nor typical forms described even for the earliest
name. The Peruvian shrub is at least expediently divisible into two
variants or species and since there is disagreement as to their
disposition a key to them is given after the description of the original
species without an implied opinion by me. See Hibiscus spiralis
for M. Poeppigii (Spreng.) G. Don, possibly M. aboreus Cav.
according to Schery.
Malvaviscus arboreus Cav. Diss. 3: 131, pi U8. 1787; 209.
Af. BaLbisii DC. Prodr. 1: 445. 1824; also M. mollis DC. I.e., fide
Schery.
456 Field Museum of Natural History — Botany, Vol. XIII
Shrub or straggling in other vegetation, rarely several meters
tall, the indument tomentose to scabrous-stellate sometimes nearly
lacking especially on the leaves above, these more or less lobed to
entire; flowers about 3-5.5 cm. long; involucel usually densely
pubescent the linear-lanceolate to spatulate segments subequaling
or exceeding the variously pubescent calyx, its lobes commonly
subdeltoid; petals often deeply retuse; stamen column finally ex-
serted ordinarily for one-fourth to one-third its length (after Schery).
The type from Mexico with the following characters: Tree with
cordate crenate subtrilobed acuminate tomentose leaves, the middle
lobe more produced; stipules setaceous, small, marcescent; peduncles
villous, axillary, solitary, shorter than petioles; bractlets 8, linear;
calyx tubular, lO-striate; corolla suffussed-red (Cavanilles). —
Illustrated, Schery, I.e. text figures and pis. 14^-17.
Peru (see below). Central America; Mexico; West Indies. —
Doubtfully in Peru at least typically but varietally, subspecifically
or perhaps specifically divisible as follows:
Calyx subturbinate or short-cylindric, at anthesis scarcely twice
longer than wide or at maturity scarcely enclosing the fruit.
M. arboreus, tjT)ical.
Calyx at least twice as long as broad at anthesis, at maturity ob-
viously enclosing the fruit.
Bractlets distinct in anthesis, rarely 3 mm. wide.
var. longifolius or M. longifolius.
Bractlets subimbricate even in anthesis; petal auricle 1 cm. long.
var. Williamsii or M. Williamsii.
Malvaviscus longifolius Garcke in Otto & Dietr. Allg. Gartenz.
22: 321. 1854. M. arboreus Cav. var. longifolius (Garcke) Schery,
Ann. Mo. Bot. Card. 29: 218. 1942. M. cuspidatus Turcz. Bull. Soc.
Nat. Mosc. 31, pt. 1: 190. 1858. M. maynensis Huber, Bol. Mus.
Goeldi 4: 583. 1906. M. Ulei Ulbr. Notizbl. Bot. Gart. BerHn 6:
328. 1915.
Like M. arboreus highly variable in leaf-lobation and pubescence
but the calyx long-cylindric, contracted above to enclose the fruit
within the tube and perhaps not stabilized even as regards its
character, as suggested by Schery but in Peru apparently distinct.
— Synonymy after Schery; more or less villous-stellate specimens
det. Schery as M. aboreus; M. maynensis as to type has subglabrous
leaves truncate or obscurely cordate at base, the branches glabrous
Flora of Peru 457
or puberulent, M. Ulex subentire openly cordate or scarcely cordate
leaves, indument a short puberulence. F.M. Neg. 9430 {M. Ulei),
San Martin : Juanjuf, KIilq 4S82; S919. Boquer6n, AUard 2207 Jt.
— Hu^nuco: Divisoria, Woytkowski Sj^59. — Junln: Juaja (Univ. of
Lima IS). Chanchamayo Valley, Schunke A116; 21; H92. San
Ram6n, Killip & Smith 2j^7H. Puerto Bermudez, KiUip & Smith
266^8.— Ay acucho: Estrella, Killip & Smith 2S065. —Loreto: Rio
Ucayali, Huber 1S83 (type, M. maynensis). Balsapuerto, Klug
SOI 5. Iquitos, WiUiams 8068 (det. Ulbrich, M. maynensis). Rio
Nanay, Williams 508. La Victoria, WiUiams 2675 (det. Ulbrich,
M. cuspicUUus) . To Colombia.
Malvaviscus Williamsii Ulbr. Notizbl. Bot. Gart. Beriin 11:
545. 1932. M. arboreus Cav. var. Williamsii (Ulbr.) Schery, Ann.
Mo. Bot. Gard. 29: 227. 1942.
Terete younger branches scabrous- tomentulose; stipules tri-
angular-lanceolate, 4 mm. long, half as wide; petioles tomentose,
5-8 cm, long; leaves cordate-orbicular, 8-11 cm. long, 9-12 cm. wide,
coarsely but rather regularly crenate, membranous, minutely and
sparsely stellulate on both sides, a little tomentose on the nerves;
peduncles 1.5-3 cm. long; involucel bractlets 9-11, ovate, acutish,
membranous, obscurely stellulate, 1 cm. wide, 1.5 cm. long, nearly
as long as the campanulate scabrous tomentose calyic, this softly
tomentose within the lobes, 4 mm. long, 5 mm. wide at base; corolla
roseate, 4-4.5 cm. long, 6-7 cm. across, the oblong petals 2-2.5 cm.
wide with a lateral triangular appendage nearly 1 cm. long, fimbriate,
also slightly stellulate and simply pilose; ovary subglabrous; stamen
tube 6 cm. long, glandular styles pilose; fruiting involucre and
calyx subindurate, carpels 8 mm. high. — Nearest M. Balbisii DC.
with smaller involucre and leaves (Ulbrich). Probably only a var.
of M. longijolius or a subvar. if that is treated as a variant of M.
arboreus. Curiously, type apparently not seen by Schery, who
wrote, "Only variety with large broad involucral lobes."
Loreto: Forest, Rio Nanay to Rio Napo, Williams 706, type.
Florida, Klug 2077. Colombia.
3. MALACHRA L.
Reference: Hochreutiner, Ann. Cons. Jard. Bot. Gendve 20:
144-149. 1917.
Perennial herbs or more or less suffrutescent, marked by the
clustered flowers often attached to conspicuously venose foliaceous
458 Field Museum of Natural History— Botany, Vol. XIII
bracts, rarely with involucre, each flower provided with subulate-
filiform bractlets, or sometimes the inner flowers subtended by
narrow stipules. Anthers subsessile, the tube truncate or 5-denticu-
late. Ovary cells 5, the solitary ovule ascending; style branches 10,
apically capitate-stigmatose. Carpels uniseriate, indehiscent or
opening vertically toward base. — Probably not more than nine
species, some extremely variable and difficult to define (Kearney);
see also Gurke, Bot. Jahrb. 16: 345. 1893. Hochreutiner suggested.
I.e. 144-145, that varieties could take care of the more stabilized
characters of the two or three established entities.
Indument canescent-tomentose or appressed on stems; inflorescences
more or less peduncled M. capitata.
Indument spreading, hispid; inflorescences sessile or subsessile.
M. alceifolia.
Malachra alceifolia Jacq. Coll. Bot. 2: 350. 1788. M. fasciata
Jacq., I.e. 352.
Abundantly hispid or rarely sparsely with spreading yellowish
simple and stellate trichomes, erect, sometimes a meter or two tall;
lower leaves cordate, 3-5-lobed or angled, the upper rounded at
base and more or less serrate or lobate, all subrotund, glabrous or
asperous or sparsely hispidulous, usually at least about a dm. long
and wide; stipules 1-2 (3) cm. long; capitate inflorescences sessile or
more or less pedunculate; bracts broadly ovate, deeply to slightly
cordate, acute to acuminate, sessile to petiolate, often pellucid-
membranous especially below in age, dentate or entire, the outer to
2.5 cm. long; calyx membranous, the lobes aristate, 5-8 mm. long
in flower, longer in fruit; petals yellowish, white or pink-tinted,
1-1.5 cm. long; carpels 3-3.5 mm. long, usually persistent. — Giirke,
I.e., retained M. fasciata and described several forms but the leaves
and bracts are variously subentire to lobed, obtuse to acute, the
carpels pilose to reticulate and glabrous (M. fasciata); varieties
could be designated, as M. fasciata, the bracts rounded to sub-
cordate, the leaf -lobes acute. F.M. Neg. 32649 (M. fasciata).
Ancash: Santa, Stork & Horton 91 5 If (det. Johnston). — Lima:
To Ancon, Mexia 8103 (det. Killip). — Loreto: Pucallpa, Soukup
3076 (det. Rudd). Yurimaguas, Williams U0Jf7. Rio Nanay,
Williams Uk7. Rio Itaya, Williams 212. To the West Indies.
"Malva."
Flora of Peru 459
Malachra capita ta L. Syst. Nat. ed. 12: 458. 1767. M. ruder-
cUis Giirke in Mart. Fl. Bras. 12, pt. 3: 460. 1892. M. Poeppigii
Gurke, Bot. Jahrb. 16: 347. 1893.
Entire plant canescent with a fine indument mostly of stellulate
and simple trichomes intermixed or the latter lacking; lower leaves
subrotund, usually 5-lobed, the upper 3-lobed or undivided, all
rounded or obtuse at base, unequally serrate, the larger leaves 6-10
cm. long, 4-7 cm. wide; heads solitary, or 2-3 in the upper axils
or terminal; bracts plane or conduplicate, the sometimes revolute
margins entire or with 1-2 teeth, sessile or subsessile, cordate, to 2
cm. long; cal)rx 6-8 mm. long, the ovate lobes obtuse; petals 1 cm.
long; carpels 3 mm. long. — M. ruderalis is the form or variety with
medially conduplicate revolute marginal bracts which appears to be
a condition scarcely of genetic origin. A photograph of the type
shows clearly, I think, that the species of Linnaeus is the basic entity
here; the two species of Giirke are the same as pointed out by Hoch-
reutiner, and may constitute at least a variety. A common weed.
Illustrated, Cav. Diss. 1 : pi. 33, fig. 1 . F.M. Neg. 9420 {M. ruderalis) .
San Martin: San Roque, WiUiams 7775. Juanjul, AUard 22512
(det, Lyman Smith). Tarapoto, WiUiaTus 5656. — Loreto: Yuri-
maguas, Poeppig 2232 (type, M. ruderalis, M. Poeppigii) ; Williams
1^368. Rancho Indiana, Mexia 6^1 9a. Near Iquitos, Williams
1311; lUU; Klug m; 78^. Rio Nanay, Williams 779; 1252. Rio
Itaya, Williams 323 J^. Pebas, Williams 1586. La Victoria, Wil-
liams 3029. Bolivia to Colombia. "Malva."
4. URENA L.
Reference: Hochreutiner, Ann. Cons. Jard. Bot. Geneve 5:
131-146. 1901.
Herbs or shrubby, the leaves usually angled or lobulate, the
small flowers commonly glomerate in the axils and involucrate by
the 5 connate bractlets. Stamen tube truncate or 5-denticulate,
the anthers subsessile. Carpels 5, usually glochidiate. — Otherwise
like Malachra, the styles also 10. The leaves commonly have 1-3
narrow thick-margined glands at the base of the medial nerves
beneath. See also Giirke, Bot. Jahrb. 16: 330-385. 1893, for the
original revision.
Urena lobata L. Sp. PI. 692. 1753; 136.
Highly variable vegetatively; leaves typically subrotund, angu-
late, usually cordate at base, the obscure lobes acute or palmately
460 Field Museum of Natural History— Botany, Vol. XIII
parted or serrate or deeply sinuate, glabrous to tomentose; calyx
5-7 mm. long, usually slightly shorter than the involucel; petals
1-2 cm. long; carpels about 6 mm. long. — U. sinuata L., if found,
may be recognized by its leaves lobed medially or more deeply.
Hochreutiner recognized and defined (modifying Giirke's treatment)
a dozen variants; a meter- tall shrub with bright lilac flowers, in
clearing (Klug). Illustrated, Bot. Mag. pi. SOJtS; Fawcett & Rendle,
Fl. Jam. 5: 127.
Loreto: Mishuyacu, Klug 9Jt^. (det. Ulbrich). Nearly cosmo-
politan in warm regions.
5. PAVONIA Cav.
More or less ligneous, or rarely completely herbaceous with ser-
rate, entire or angulate-lobed often pellucid-punctate leaves and
variously colored flowers solitary and peduncled in the axils or
clustered at the ends of the branches. Bractlets 5-many, distinct
or more or less connate basally and with the calyx. Carpels obo-
void-trigonous, never glochidiate but sometimes echinulate and
often 1-3-aristate or -rostrate, often separating septicidally. Ac-
cording to Hochreutiner carpels early or tardily dehiscent, also
loculicidally, except in species with obsolete dorsal nerve, while
those of P. spinifex (et cetera with thick-walled carpels) split open
only upon germination. Style branches and stigmas always 10 but
ovary with 5 monospermed loculi (B.P.G.H.).
Two keys are given, the one based on fruiting characters kindly
contributed by Dr. Hochreutiner; however, the vegetative key,
while not accounting for leaf-variations, will lead to most flowering
specimens, otherwise scarcely determinable.
Carpels with long hamate-pilose awns; leaves usually more than
twice as long as wide; flowers solitary in the axils or grouped
at end of stem or branches.
Awns narrow, nearly filiform; carpel body transversely rugose.
Leaves cordate or subcordate; bractlets 10-16 mm. long.
P. spinifex.
Leaves not cordate; bractlets 4-8 mm. long P. sepium.
Awns unequal or shorter than carpels, these not distinctly trans-
versely rugose.
Bractlets 5-6, ovate; inflorescence densely umbellate.
P. fruticosa.
Flora of Peru 461
Bractlets 8-10, linear; inflorescence laxly paniculate.
Bractlets much longer than caljrx.
P. oxyphyUaria, P. peruviana.
Caljrx much longer than bractlets P. leucantha.
Carpels awnless, sometimes only mucronate, smooth; inflorescence
capituliform P. Riedelii.
Carpels awnless, smooth or rugose, muticous or gibbous.
Stipules lanceolate; petals yellow P. spicata, P. paniculata.
Stipules filiform; petals yellow P. sidaefolia.
Petals red P. mollis.
Vegetative Key
Flowers solitary in the axils (rarely lateral and 2), at least 1.5 cm.
long (unless rarely in P. septum).
Leaves ovate to oblong-elliptic, at most 2-3 times longer than
wide.
Indument soft to touch.
Bractlets linear; petals cuspidate, red P. mollis.
Bractlets lanceolate; petals yellow P. sidaefolia.
Indument of leaves scabrous, at least upper surface (Peru).
Petals 2.5 cm. long or longer; leaves often somewhat cordate.
^ 'P. spinifex.
Petals less than 2 cm. long; leaves usually narrowed to base.
P. sepium.
Leaves lanceolate to oblong-lanceolate, several times longer than
L: wide.
m Flowers subsessile; bractlets expanded apically. . . .P. Riedelii.
^' Flowers long-peduncled; bractlets linear, hispid.
P. oxyphyUaria.
Flowers all or mostly several together and always shorter than 1.5
cm.
Leaves ovate or rotund to about twice as long as wide; carpels
weakly armed if at all.
Bractlets hispid, exceeding calyx P. panicuUda.
Bractlets short-stellate, shorter than caXyx. P. spicata.
Leaves usually much more than twice as long as wide; carpels
spinose at tip.
462 Field Museum of Natural History— Botany, Vol. XIII
Tall, erect, with peduncled or capitate inflorescences; calyx
and involucel subequal.
Stipules conspicuous; flowers capitate; carpel awns unequal.
P. fruticosa.
Stipules minute; flowers laxly corymbose; carpel awns sub-
equal P. leucantha.
Low, ascending; stipules conspicuous; csdyx and bractlets very
unequal P. peruviana.
Pavonia fruticosa (Mill.) Fawcett & Rendle, Fl. Jam. 5: 130.
1926. Sida fruticosa Mill. Gard. Diet. ed. 8. 1768. P. typhalaea
(L.) Cav. Diss. 3: 134. 1787. Urena typhalaea L. Mant. 2: 258. 1771.
More or less ligneous with few if any branches, sparsely and
minutely stellate pubescent, at least the tips and the leaves both
sides, these oblong- or lanceolate-elliptic, sometimes rather obovate,
more or less acuminate, cuneate to rounded at base, coarsely and
irregularly serrate; flowers usually 5-15 or more, capitate at tips of
stems and branchlets; bractlets 5-8, ovate-lanceolate, united about
one-fourth, (6) 8-10 mm. long; calyx 5-7 mm. long; petals white,
10-13 mm. long, longer than stamen tube; carpels trigonous, 5-6
mm. long, dehiscing from base, smooth dorsally but with 3 apical
retrorsely setose spines, the longer medial 5-7 mm. long. — Often
a meter or two high. Determinations by Standley except as noted.
Uittien in Pulle, Fl. Surinam 3: 434. 1941, who overlooked. I.e. 14,
the Miller name, suggests that it can be discarded by calling the
latter's work an "opus negligendum," Fawcett & Rendle identified
the Miller plant; the comer stone of plant naming rests on the law
of priority; if Uittien's idea is followed his own may well be discarded
some day by new students; once disrespect for established basic
nomenclatorial law occurs, it becomes a modus operandi; compare
Journ. Wash. Acad. Sci. 19: 247-252. 1929. P. rosea Schlecht.,
perhaps occurring, is similar but the often roseate flowers are
corymbose or loosely capitate, the bractlets linear-lanceolate.
Illustrated, Mart. Fl. Bras. 12, pt. 3: pi. 87.
San Martin: Cainarachi, Klug 2732. Tarapoto, Williams 5852
(det. Ulbrich). San Roque, Williams 7080. — Junin: La Merced,
5366 (det. Blake, P. rosea). Puerto Yessup, Killip & Smith 26266
(det. Killip).— Hudnuco: Tingo Maria, Allard 21163; 21904- (det.
Lyman Smith). Pampayaca, 5128 (det. Blake, P. rosea). — Loreto:
Rio Paranapura, Klug 3953. Iquitos, Williams 13^2 (det. Ulbrich);
8103. Rio Nanay, Williams U29; J^U8 (det. Ulbrich). Caballo-
Flora of Peru 463
Cocha, Williams 21H (det. Ulbrich). Mishuyacu, Klug 1US9.
Iquitos, KiUip & Smith 27259 (det. Killip). To Central America
and the West Indies.
Pavonia leucantha Garcke, Jahrb. Bot. Gart. Berlin 1: 211.
1881.
Stem more or less appressed hirsute-stellate especially toward
the apex, slender, irregularly branched above, a meter or so high;
stipules 2-3 mm. long, deciduous; petioles about 1 cm. long; leaves
oblong-elliptic or lanceolate, often 3-4 times longer than wide, the
lower to about 1.5 dm. long, 3-6 cm. wide, rounded to somewhat
acute at base, acute or acuminate, serrate, minutely appressed
stellate both sides and beneath, hirtellous; corymbs 3-6-flowered
the lower p)eduncles 5-6 mm. long, the basal bracts 5-parted, in-
volucral bractlets 8-10, connate about medially, lanceolate, 3-
nerved, 4-5 mm. long, the campanulate stellate calyx 5-6 mm. long,
its deltoid acute lobes ciliate; petals 10-12 mm. long, exceeding the
glabrous stamen tube; carpels glabrous, dorsally 1-nerved, 7 mm.
long, the retrorsely pubescent awn 3 mm. long. — Determinations by
Standley. Illustrated, Mart. Fl. Bras. 12, pt. 3: pi. 89. F.M
Neg. 9452.
San Martin: San Roque, Williams 697U; 7158; 762S; 7683; 7722.
Lamas, Williams 6437. Vitoc, Ruiz. Zepelacio, Klitg 35ItO. Tara-
poto, Williams 6713; Woytkowski 35069 (det. Cuatrecasas). — Loreto:
Rio Itaya, WiUiams 3U; 2^1 (det. Ulbrich). Yurimaguas, Poeppig
2302, type; Williams 5239; 5305; 5092. Near Iquitos, Klug 1032;
WiUiams 8101; 8206. Santa Maria, AUard 22^63 (det. Lyman
Smith). Ecuador; Bolivia.
Pavonia mollis HBK. Nov. Gen. & Sp. 5: 282. 1822. Astero-
chlaena cv^pidata Garcke, Bot. Zeit. 8: 668. 1850. P. Kunthii
GUrke var. mollis (HBK.) GUrke in Mart. Fl. Bras. 12, pt. 3: 511.
1892. Hibiscus cordifolius L.f. Suppl. 309. 1781, fide Glirke, not
P. cordifolia Wawra.
Shrub, a meter or so high, the slender flowering stems stellate
puberulent and more or less hispid with somewhat glandular spread-
ing trichomes, the subrotund-ovate cordate-based acuminate leaves
softly stellate tomentose both sides or rarely sparsely so, the flowers
all axillary on slender peduncles as long or longer than the leaves;
stipules 3-4 mm. long; petioles to 10 cm. long; leaves to a dm. long,
and nearly as wide or the upper considerably smaller, crenate-serrate;
bractlets 7-9, linear, ciliate and puberulent, 8-16 mm. long, about 1
464 Field Museum of Natural History— Botany, Vol. XIII
mm. wide; calyx 4-5 mm. long, stellate and somewhat hirsute; petals
1.5-2 cm. long, rose-colored, glabrous or nearly without; stamen
tube glabrous, 1-1.5 cm. long; carpels muticous, reticulate-rugose,
pubescent, coriaceous, about 5 mm. long, the sparsely puberulent
seeds about 3-3.5 mm. long. — Fruit tardily dehiscent, loculicidally
and finally septicidally (Hochreutiner) . The type of A. cuspidata
was from Guayaquil. F.M. Negs. 9438 (A. cuspidata); 9450.
San Martin: Tarapoto, Williams 5792 (det. Ulbrich, A. cus-pi-
data). Juanjui, Klug ^221 (det. Killip). Zepelacio, Klug 3538.
Ecuador; Colombia.
Pavonia oxyphyllaria Donn. Sm. Bot. Gaz. 23: 237. 1897.
P. costaricensis Hochr. Ann. Cons. Jard. Bot. Geneve 10: 18. 1906,
fide Standley.
Ligneous cylindrical stems more or less pubescent, with long
reddish trichomes; stipules subulate, filiform, rufous villous as the
petioles (2-5 mm. long) and peduncles, these about 1.5 cm. long, to
5 cm. long in fruit; leaves lanceolate-oblong or oblong-elliptic, acute
or acuminate, serrate; peduncles axillary and congested at apex of
stems, articulate in upper part; involucral bracts about 13, linear-
filiform, conspicuously yellowish-red villous and setose, about 1.5
cm. long, scarcely 1 mm. wide at base; sepals about 4 mm. long,
tomentose; petals obliquely obovate, retuse, pubescent both sides;
fruit depressed, the 5 carpels dorsally nervose, the 3 spines about
1 cm. long with reflexed setae; seeds 5 mm. long. — Related to
P. Pseudo-Typhalaea with larger fruits and differing from P. Warm-
ingiana by the form of the fruit (Hochreutiner; description of petals
after Smith). The second record for Peru, according to Ulbrich.
Loreto: Pebas, Williams 1H7 (det. Ulbrich). Costa Rica;
Panama.
Pavonia paniculata Cav. Diss. 3: 135, pi. U6, fig. 2. 1787.
Half-shrub sometimes a meter or two high, more or less glandular
and stellate pubescent, often setose with some longer simple spread-
ing trichomes; stipules lanceolate, to 1 cm. long; petioles several to
many cm. long, about equaling the ovate, angled or somewhat
lobate leaves these glabrescent above or the nerves and veins
stellate puberulent, cordate at base, acute or acuminate at apex
(as lobes); flowers usually in a compound corjnnb or panicle, or
solitary in the upper leaf axils; bractlets 7-12, distinct, linear,
hispid, much exceeding calyx, this 5-8 mm. long, the yellow petals
Flora of Peru 465
sometimes twice as long; stamen tube antheriferous from base, 5-6
mm. long; carpels trigonous, 3-4 mm. long, dorsally a little rugulose,
edges scabrous, apex rarely with a weak short spine. — Determina-
tions by Standley except as noted. Illustrated, Mart. Fl. Bras. 12,
pt. 3: pi. 96.
Lima: Rio Rimac, Safford. — San Martin: Chuzuta, Klug U008.
Zepelacio, Klug SSS3. Juanjul, Klug It221. Shores of Lake Rilami-
Cocha, Woytkowski 35126 (det. Cuatrecasas). — Hudnuco: Chinchao,
Mexia 0^151. Rio Maran6n, Domhey, type. Cuchero, Poeppig.
Near Hudnuco, in thickets and along fences, 20^7; 3538 (det.
Hochreutiner). — Junin: Colonia Peren6, KiUip & Smith 2U992 (det.
Killip). Oxapampa, Soukup 2J!t29 (det. Rudd). Satipo, Soukup
2850. La Merced, Killip & Smith 231^72; 2j^051. Puerto Bermudez,
Killip & Smith 26622.— Ay acucho: AIna, Killip & Smith 23106;
22554 (det. Killip). — Loreto: Common field weed, Iquitos, Williams
7933; 8209. Florida, Klug 20U- Leticia, Williams 303U (det.
Ulbrich, var. corymbosa Gurke). To the West Indies. "Malva-
malva."
Pavonia peruviana Giirke in Mart. Fl. Bras. 12, pt. 3: 487.
1892. P. parva Ulbr. Notizbl. Bot. Gart. Berlin 11: 543. 1932.
P. nana Ulbr. Verb. Bot. Ver. Brandenb. 50: 85. 1908, not R.E.
Fries, Bull. Herb. Boiss. s^r. 2. 7: 999. 1907.
Ascending-erect, the usually arcuate simple stem 1-few dm. tall,
more or less simply hirsute especially above and on the leaves both
sides (typically) or glabrate in age; lower petioles 1-2 cm. long;
stipules linear-lanceolate, rigid, deciduous, about 1 cm. long; leaves
oblong-elliptic, rounded to cuneate at base, acuminate, the larger
intermediate ones to about a dm. long, a third as wide, rarely some-
what stellate pubescent, coarsely and irregularly serrate; flowers
corymbosely congested, in age more or less laxly few- to several-
flowered, the peduncles 5-15 (20) mm. long with a triparted basal
bract; involucel turbinate, the 9-10 linear acuminate 3-nerved
segments 8-11 mm. long, 1 mm. wide, sparsely long-hirsute without
and ciliate; calyx campanulate, 4-5 mm. long, stellate-pubescent
without, the broadly deltoid lobes ciliate; petals obovate, very
obtuse, 8-10 mm. long, puberulent; stamen tube 5-6 mm. long,
glabrous; carpels 5, trigonous, 1-nerved, glabrous, 9-10 mm. long,
the awns about (2.5) 4-5 mm. long, erect, retrorsely setose; seeds
glabrous. — P. Warmingiana Giirke, I.e. 488, of Brazil, to which some
Peruvian specimens have been referred, was distinguished by more
466 Field Museum of Natural History— Botany, Vol. XIII
dense inflorescence, less connate involucel bractlets, pilose calyx
lobes, and P. parva from it by rose-colored exserted flowers, sub-
equal awns; none of these characters seem to be constant or con-
comitant, but should be studied when more collections are available.
Compare also P. rosea Schlecht. under P. fruticosa, with entire
bracts, the lateral carpel awns much shorter than the medial. Illus-
trated, Giirke, I.e., pi 88. F.M. Negs. 9458 (P. parva); 23705.
San Martin: San Roque, Williams 7735 (det. Ulbrich, P. parva)',
7637 (distr. as P. Warmingiana) ; 6988 (det. Ulbrich, P. Warmingi-
ana, with query). — Hudnuco: Pampayacu, Poeppig 158^, type.
Without locality (Mathews 6J^Jf). — Loreto: Cerro near St. Antonio
de Cumbaso, Ule 6861 (tjrpe, P. parva). Brazil; Colombia.
Pavonia Riedelii Giirke in Mart. Fl. Bras. 12, pt. 3: 493. 1892.
Stems slender, more or less lineately stellate-puberulent, gla-
brescent below, this indument also on the petioles (4-5 mm. long),
peduncles (1-4 mm. long) and oblong-lanceolate leaves, especially
above; stipules subulate-filiform, 3-4 mm. long; leaves subrotund
or slightly cordate at base, acuminate, mostly 4-6 cm. long, 1-1.5
(2) cm. wide, irregularly serrate-crenate, gradually smaller above,
becoming bractiform; inflorescences capituliform, 3-5-flowered, in
the upper axils or at the tips of accessory branches; involucel bract-
lets 9-11, linear, hispid with simple trichomes, 5-7 mm. long, usually
with a broader reflexed apical appendage; calyx cupulate, hirsute,
7-9 mm. long, the acuminate lobes 3-nerved; petals yellowish, to
about 2.5 cm. long; carpels trigonous, dorsally convex, membranous,
hirtellous, obtuse but shortly mucronulate, 3-4 mm. long. — F.M.
Neg. 9465. Illustrated, Giirke, I.e., pi. 91, fig. 2 (analysis).
San Martin: San Roque, Williams 7120; 7358 (det. Ulbrich).
Zepelacio, Klug 3581 (det. Standley). Brazil.
Pavonia sepium St. Hil. Fl. Bras. Merid. 1: 225. 1827.
Much like the related P. spinifex and in Peru apparently merging
but in general smaller in all parts; leaves usually oblong-ovate,
narrowed or obtuse at base, rarely rounded or lightly cordate, often
7-10 cm. long, 3-4 cm. wide; petals 12-15 mm. long; stamen tube
8-10 mm. long; carpels 4 (-6) mm. long with spines 5-7 mm. long.
— The subsp. macrocarpa R.E. Fr. Sv. Vet. Akad. Handl. 24, no. 2:
24. 1947 has ovate leaves long-narrowed to tip, shortly acute, rounded
or rarely lightly cordate at base, early densely stellate beneath, the
carpels 11-12 mm. including the 5-6 mm. long spines, a variant
Flora of Peru 4ffl
apparently better treated as merely a variety. Illustrated, Mart.
Fl. Bras. 12, pt. 3: pi. 86, fig. 1.
Cajamarca: Huancabamba, Sandeman 4^22. — Hudnuco: Near
Hudnuco (Asplund 1SU6S, det. Fries, subsp. macrocarpa). — Junin:
Pichis Trail, Killip & Smith 25880 (det. Killip).— Cuzco: Machu-
picchu, Herrera 3219 (distr. as P. paniculata). Southern South
America to Colombia.
Pavonia sidaefolia HBK. Nov. Gen. & Sp. 5: 283. 1822.
Velvety tomentose even to the involucels and calyces and
typically also more or less spreading hirsute with long simple tri-
chomes; stipules 3^ mm. long; petioles 1-4 cm. long; leaves cordate-
ovate, acute, crenate, usually 4-7 cm. long, 3-4 cm. wide; peduncles
axillary, 2-6 cm. long; bractlets of the involucel 6-9, lanceolate,
1-2 cm. long, 2.5-5 mm. wide; calyx 5-10 mm. long, the lobes
deltoid, 5-nerved; petals yellow, red at pubescent base, 2-2.5 cm.
long; stamen tube glabrous, 6-7 mm. long; carpels coriaceous,
obsoletely reticulate-rugulose, 3^ mm. long, gibbously obtuse;
seeds striate, 2.5-3 mm. long. — The Peruvian plant is the var.
diuretica (St. Hil.) Giirke without elongate trichomes. Illustrated,
St. Hilaire, PI. Usu. pi. 58. F.M. Neg. 9796.
The diphthong in the species name may be written "t" in accord
with recommendation 44 of the International Rules of Botanical
Nomenclature; however, when this work started it was the intent
at least to follow the spelling of the author and indeed this has been
found to be the sensible solution also as regards place names, native
names and even the name of the author himself, since an arbitrary
attempt at uniformity may be more trouble than it is worth and may
even lead to greater confusion.
San Martin: Tarapoto, Woytkowski 850^3 (det. Cuatrecasas) ;
Williams 5539 (det. Ulbrich); also 5406; 5515. To Venezuela,
Paraguay.
Pavonia spicata Cav. Diss. 3: 136. 1787. Malache acabra Vogel
in Trew, PI. Sel. 50, pi. 90. 1772, not P. acabra Presl.
Shrub, the younger parts even to the calyces finely stellate-
tomentose or the ovate leaves glabrate both sides except toward the
cordate base; glandulosity lacking; stipules linear-lanceolate, often
a cm. long; petioles about a third as long as the leaves, these nar-
rowly acuminate, entire or minutely serrate-dentate, often about
a dm, long or longer; flowers in terminal sometimes more or less
468 Field Museum of Natural History— Botany, Vol. XIII
corymbose racemes; bractlets 8-10, oblong-lanceolate, shorter than
calyx, this 12-13 mm. long; the greenish-yellow petals 1.5-2 cm.
long; stamen tube 1.5 cm. long; carpels acutely trigonous, 9-11 mm.
long, dehiscent within medially, sometimes crested dorsally, bluntly
3-cusped apically.
Peru (probably). Tropical South America to Florida.
Pavonia spinifex (L.) Cav. Diss. 3: 133, pi. It5, jig. 2. 1787.
Hibiscus spinifex L. Syst. Nat. ed. 10. 1149. 1759.
Glabrescent or lightly stellate-puberulent shrub or half-shrub
with rather large mostly solitary axillary yellow flowers and irregu-
larly serrate ovate leaves; stipules to about 1 cm. long; petioles 1-3
cm. long or the lower twice as long; leaves usually somewhat cordate,
the lowest to 12 cm. long, 8 cm. wide; peduncles usually 1-4 cm.
long; bractlets nearly distinct; calyx deeply parted, 9-12 mm. long;
petals yellow, 2-3 cm. long, sometimes larger, usually exceeded by
the stamen tube; carpels 4-6 mm. long, indehiscent, transversely
rugulose dorsally and with 3 ribs terminating at apex in retrorsely
setose spines 5-10 mm. long. — Illustrated, Fawcett and Rendle,
Fl. Jam. 5: 129; Mart. Fl. Bras. 12, pt. 3: pi. 85.
Cajamarca: Prov. Hualgayoc, Soukup 3852. — Hudnuco: Pam-
payacu, Sawada 65. Hudnuco, 20^8. Mito, 1570. Huacho, Stork
& Horton 9^11 (det. Standley). — Cuzco: Abandoned field, Echarate,
Goodspeed Ezped. 10U52 (det. Standley). San Miguel, Cook &
Gilbert 1037; 1071. Machupicchu, yargras ^OSi . Warm and tropical
America. "Anguia," "taroca-asta," "cuemo de venado" (Cook &
Gilbert).
6. HIBISCUS L.
Reference: Hochreutiner, Ann. Cons. Jard. Bot. Geneve 4:
23-191. 1900.
Shrubs or vigorous herbs, rarely arborescent, with mostly showy
flowers involucrate by several to many usually narrow, rarely
reduced free or somewhat connate bractlets. Leaves various.
Stamens disposed evenly or unevenly below the truncate or dentate
column apex. Ovary cells 5, each (2) 3-many-ovuled. Style
branches 5, the stigmatic part globose or spatulate-dilated. Capsules
loculicidal, rarely ligneous, sometimes with false partitions, the
seeds usually reniform, glabrous or pubescent. Bractlets rarely
bifurcate at tip, sometimes simple in part in the same involucel.
Flora of Peru 469
The detailed monograph of Hochreutiner is without descriptions
but replete with sjmonymy and keys, with many critical notes and
observations. Since the Peruvian species are mostly relics from culti-
vation and popular ornamentals in many plazas, a purely artificial
key is appended to facilitate their identification.
Peduncles shorter than flowers.
Leaves rotund-reniform, entire, green above, canescent beneath.
H. tiliacexis.
Leaves serrate to lobed, glabrescent or pubescent on both sides.
Flowers about 4 cm. long; plants glabrescent. . .H. cannabinus.
Flowers larger; plants more or less pubescent.
Bractlets typically forked apically; leaves often rather cor-
date-ovate.
Bractlets longer than calyx, stellate-hirsute.
Stems aculeate- verruculose H. bifurccUus.
Stems not aculeate H. furcellatus.
Bractlets shorter than calyx, puberulent H. peruvianus.
Bractlets simple; leaves all oblong-lanceolate, often tomen-
tose H. Lamhertianus.
Peduncles mostly (or all) at least soon longer than flowers.
Petioles, peduncles, often leaves beneath densely pubescent.
Leaf lobes acute; calyx soon inflated H. miUabilis.
Leaf lobes rounded; calyx little accrescent H. HUchcockii.
Petioles, peduncles, leaves glabrous or only early pubescent.
Flowers about 2 cm. long; bractlets often exceeding calyx.
Sepals more than half connate H. spiralis.
Sepals free or less than half connate.
H, Cavanillesianiis, H. brasiliensis.
Flowers much larger; bractlets at most equaling calyx.
Petals entire; bractlets and calyic subequal.
Bracts as calyces glabrous or trichomes fine; leaves often
lobed, minutely serrate H. Sabdariffa.
Bracts as calyces setose; leaves coarsely serrate.
H. rosa-sinensis.
Petals dissected; bractlets minute H. schizopetalus.
Hibiscus bifurcatus Cav. Diss. 3: 146, pi. 51. 1787; 108.
Glabrescent or setose, but stems, i)etioles and leaf midnerve
more or less prickly, the lower leaves typically deeply and acutely
470 Field Museum of Natural History— Botany, Vol. XIII
lobed, sometimes sub-hastate, often nearly glabrous, cordate, the
upper leaves ovate-lanceolate, acuminate, all acutely serrate and
with a gland on midnerve; peduncles shorter than calyces these
becoming coriaceous, nervose with a medial nerve for each lobe and
sinus, the latter nerve parted at sinus to extend marginally on each
lobe anastomosing at apex with the medial nerve; flowers axillary
and apically spicate, pink or violet; bractlets linear-filiform, con-
spicuously forked at tip, free but joined at base to calyx; capsules
barely included, ovoid, acute, flavescent-pubescent; seeds 3 mm.
long, glabrous.
Peru (probably) . Tropical and subtropical America.
Hibiscus brasiliensis L. Sp. PI. ed. 2. 977. 1763; 87. H.
phoeniceus Jacq. Hort. Vindob. 3: 11, pi. 1^. 1776, fide Hochreutiner.
Slender-stemmed soon glabrescent shrubs sometimes several
meters high; petioles about 1.5 cm. long; leaves ovate, truncate or
subcordate at base, acuminate, sometimes the lower deeply 3-lobed,
those of the flowering branches often about 6 cm. long, 4 cm. wide;
peduncles typically elongate, articulate about medially; bracts
narrowly linear, glabrous or nearly, usually 2 cm. long, equaling or
much exceeding the subglabrous calyx, nearly equally wide from
base to apex; calyx deeply lobed, 10-12 mm. long, the ovate lance-
olate lobes acuminate; flowers rose-colored or rarely white, 1.5-2
cm. long. — The var. sylvaticus (Benth.) Hochr. has the larger lower
leaves deeply 3-lobed, 8-9 cm. long, 12-16 cm. wide; var. luteus
[Pavon] Hochr. has yellowish rigid stellate trichomes especially on
the stems and leaves beneath, peduncles articulate in the upper
one-fifth and bractlets densely ciliate. This could well be H.
Cavanillesianus HBK. but compare note under that species.
Tumbez: Rainy-green formation, southeast of Hacienda La
Choza, Weberhauer 7717 (det. Ulbrich, H. phoeniceus). — Loreto:
In garden, Caballo-Cocha, Williams 2373 (det. Standley). Northern
South America and the West Indies.
Hibiscus cannabinus L. Syst. Nat. ed. 10. 1149. 1759; 114.
H. unidens Lindl. Bot. Reg. 10: pi. 878. 1825.
Highly variable in size and contour of leaves, indument and
presence of glands on caly^, the bracts simple or more or less bi-
furcate; petioles and leaves subequal, the latter usually glabrescent,
more or less deeply 3-lobed or -parted, the ovate or oblong-lanceolate
lobes acute or acuminate, serrate; peduncles short or flowers even
Flora of Peru 471
subsessile, these often about 3 cm. long; bracts connate at base,
linear, commonly simple, rarely one or more somewhat 2-forked
at tip, subequaling the calyx, its acuminate lobes as the bracts with
a few rigid hyaline trichomes, these also present on the shorter
ligneous pointed capsules. — The var. unidens (Lindl.) Hochr, I.e.
115, has the involucel more or less forked.
This has been cultivated as a source of fiber.
Loreto: In garden, Fortaleza, Williams U508; also at Nanay,
but apparently wild, Jt22 (both det. Standley). Old and New World
tropics.
Hibiscus Cavanillesianus HBK. Nov. Gen. & Sp. 5: 290.
1822; 124.
Shrubby, a meter high or higher, with smooth glabrous stems;
branchlets hispid-stellate; stipules persisting, 6-8 mm. long; petioles
8-10 mm. long; leaves subrhombic to ovate-oblong, acute at base,
narrowly acuminate, 3.5 cm. long or longer, about 2.5 cm. wide,
lightly pubescent on both sides or especially beneath, the indument
appressed with rigid scattered stellate trichomes, coarsely and
irregularly serrate; peduncles at apex of axillary branchlets about
8 mm. long, shortly hispid as the 10-11 linear subspreading bractlets;
calyx lobes ovate, acute, 3-nerved; petals violet, suborbicular,
stellate-pubescent without, 14-16 mm. long; styles 5; ovules 6;
carpels depressed, ovate, hirsute; seeds 4. — Hochreutiner remarks:
very doubtful species, placed in Furcaria group with simple bractlets
from the description of the calyx, but type not at Paris. However,
compare H. brasiliensis, var. liUeiis with pubescent bractlets.
Hochreutiner suggests to me that species should be suppressed as
diagnosis omits seeds and permits only h5rpothesis; however, it will
probably be identified by recollection at type locality.
Cajamarca: Near Tomependa, Bracamores de Ja^n (Bonpland,
type).
Hibiscus furcellatus Desr. in Lam. Encycl. 3: 358. 1789; 107.
Similar to H. hijurcaius but bractlets often less or not at all
forked and all younger parts more or less tomentose-scabrous with
brownish-gray stellate trichomes; leaves usually entire or angulate,
more or less obscurely dentate; calyx shorter than involucel the
acute lobes glanduliferous; petals 6-8 cm. long, red. — The var.
Diodon (DC.) Uitt., Pulle, Fl. Surinam 3: 21. 1932, has 3-5-lobed
leaves, bractlets more or less bifurcate, var. afurca Uitt., the bract-
I
472 Field Museum of Natural History — Botany, Vol. XIII
lets entire; the species therefore seems to merge with typical H.
bifurcatus and probably represents a phase of it, the characters
apparently variable, not consistently concomitant, even intangible.
F.M. Negs. 7988; 23725; 23726 (type and vars.). Illustrated,
Rodrigo, Rev. Mus. de la Plata n. ser. 7, Bot.: 119.
Hudnuco: Divisoria, Woytkowski 3j^55? Tropical and subtropical
America.
Hibiscus Hitchcockii Ulbr. ex Kearney, Leafl. West. Bot. 7:
271. 1955.
Shrub, about 2 meters tall, the younger parts minutely stellate-
tomentose, the leaves becoming glabrescent above; pyetioles about
a third as long as the suborbicular blades, these at least 11 cm. long,
with or without a shallow open sinus, shortly 3-5-lobed, the rounded
lobes sparsely crenate; peduncles often longer than 9 cm., solitary
in the upper axils or subcorjrmbosely clustered at the apex of the
stem and the few branches; involucel of 8-9 bractlets 12-18 mm.
long, the conduplicate ovate blade — this nearly 1 cm. wide — much
longer than the subcylindric lower portion; calyx at anthesis 19-22
mm. long, cleft nearly to base, the lanceolate acuminate 5-nerved
lobes eglandular; corolla funnelform-campanulate, the roseate
petals scurfy puberulent without, 6-7 cm. long; stamen tube much
shorter than style, this included, the stout branches clavate; capsule
ovoid, 2-2.5 cm. long, scurfy puberulent, equaling or subequaling
calyx, the rigid valve-cusps 4 mm. long, very sharp; seeds densely
lanate. — Characters of involucel suggest relationship to H. sororins
L. f., which was placed by Hochreutiner in his revision. I.e. 166,
167, in his section Spatula, but he characterized this section as
having glabrous seeds (Kearney) ; in Peru H. sororius L. f . resembles
most H. furcellatus Desr. but the similarly dilated bractlets are not
forked, the leaves not lobed; as Kearney remarks, Ulbrich's species
appears to be very distinct; type from near Guayaquil.
Piura: between Canchaque and Serrdn, Prov. Huancabamba,
Stork llJf21 (det. Johnston). Ecuador.
Hibiscus Lambertianus HBK. Nov. Gen. & Sp. 5: 291, pi.
U78. 1821; 142. H. salviaefolius St. Hil. Fl. Bras. Merid. 1: 249.
1827, fide Hochreutiner.
Slender, typically canescent-tomentose, velvety in Peru, or in
type leaves more hirsute above (in a variety glabrous) these oblong-
or ovate-lanceolate, serrulate, truncate at base, acute to acuminate.
I
Flora of Peru 473
often about 1 dm. long, 3-3.5 cm. wide; peduncles short; bractlets
linear, free, simple, shorter than the soon inflated shortly lobed
cal3rx, the broad lobes without marginal nerves; petals about 1 dm.
long; capsules included, short-setulose. — The flowers of the Peruvian
specimen noted by collector as pale purple, darker at base, column
purple, stigma white; the leaves are about equally velvety-tomen-
tulose both sides and acute rather than acuminate, in this resembling
the St. Hilaire type to which it was referred. F.M. Negs. 35500;
23729 (last two, vars.); 35499 (H. salviaefolius) . Illustrated, Rev.
Mus. de la Plata n. ser. 7, Bot.: 133.
San Martin: Shores of Lake Rikuri-Cocha, Tarapoto, Woytkow-
ski S5124. Colombia to Brazil and Paraguay.
Hibiscus mutabilis L. Sp. PI. 694. 1753; 147.
Shrubs or small trees with ample cordate-rotund less than
medially lobed leaves, the lobes acute, and large flowers that charac-
teristically change color, commonly opening white, becoming
roseate and finally yellowish; indument close, scurfy-stellate,
especially dense on the upper stems, bracts and calyces, the second
free, narrowly lance-linear, soon shorter than the quickly enlarged
or in fruit much inflated cal}^, its broadly ovate acuminate lobes
nervose but not marginally; capsules included, the seeds lanuginose.
— Probably originally from China or Japan, but long established in
South America and tropical America. Determinations by Standley.
Loreto: La Victoria, Williams 2760. Lower Rfo Huallaga,
WiUiams j^55. Rio Maraiion Valley, Dennis 29167 (det. Killip).
Rio Itaya, WiUiams 215. Tropical regions. "Flor variable."
Hibiscus peruvianus R. E. Fr. in Sv. Vet. Akad. Handl. ser.
3. 24, no. 2: 31. 1947.
Branchlets, petioles and leaves beneath canescently stellate-
puberulent, glabrescent in age, and more or less minutely aculeate
with recurved yellowish prickles; petioles to 7 cm. long; leaves
ovate-triangular, basally truncate or slightly and openly cordate,
to one-third sinuately 3-5-lobate, the divergent deltoid lobes acute
or the uppermost leaves little if at all lobed, all irregularly crenate,
the largest 6-10 cm. long and broad, membranous, green, glabrous
above except slightly hirsute on the principal nerves, early minutely
stellate-pubescent beneath; flowers solitary in the axils; pedicels 2.5-3
cm. long, to 6 cm. long in fruit, shortly and densely cinereous-tomen-
tulose, sometimes abundantly aculeate; involucral bractlets 9-10,
474 Field Museum of Natural History— Botany, Vol. XIII
densely puberulent with rigid stellulate trichomes, linear, about 12
mm. long and 1 mm. broad, bifurcate apically; calyx cupulate, canes-
cent-tomentulose, about 1.5 cm. long, the lanceolate-deltoid acute 3-
nerved lobes with a small round gland on the medial nerve; an-
droecium about 3.5 cm. long; ovary narrowly ovoid, acute, densely
appressed yellowish-setose. — Similar to H. bifurcatus Cav. (and H.
jurcellatiLS Desr.), both of which have hirsute or hispid-stellate
involucels and calyces. Illustrated, Fries, I.e., pi. 2, figs. 11-13.
Loreto: Florida, Ucayali (Tessmann 8072, type). Florida, Klug
2182. Santa Rosa, Killip & Smith 28956. Near mouth Rio Mara-
fion, Dennis 29237. Mishuyacu, Klug 1J^73 . Caballo-Cocha, Wi7^taws
2304' (det. Ulbrich, H. furcellatus) . Pebas, Williams 1862. Naza-
rete, Osgood 23; 2Jt.. — Huancayo: Near Huanta, Dennis 29239.
"Binaqui-ey" (Huitoto, Klug).
Hibiscus rosa-sinensis L. Sp. PI. 694. 1753; 133.
Glabrous or essentially glabrous shrub with ovate or ovate-
elliptic acute or shortly acuminate crenate serrate leaves, these
mostly rounded at the base, about 8 cm. long, 4-5 cm. wide; pe-
duncles articulate above the middle; bractlets 5-8, sublinear,
shorter than the rather tubular clayx, this about 2.5 cm. long,
deeply dentate; flowers 7-10 cm. long, the petals entire, the stamen
column exserted, the anthers evenly disposed; capsules obovoid,
seeds not lanate. — Conforming with present custom the specific
name is hyphenated; commonly grown for its beauty it not in-
frequently is collected as an escape, only a few of many collections
cited. H. syriacus L., totally different but conveniently noticed
here as perhaps also cultivated and escaping, has basally cuneate
leaves, somewhat 3-lobate, the bractlets at least equaling the calyx,
the stamen column not exserted.
San Martin: Tarapoto, Williams 59 U5. — Hudnuco: In the plaza,
Hudnuco, Woytkowski. — Loreto: Clearing, Mishuyacu, near Iquitos,
Klug 678. In field near Iquitos, Williams 82^5. Yurimaguas,
Williams U08U. Rio Itaya, Williams 219. Rio Nanay, Williams
J^33. Caballo-Cocha, Williams 2375; 2376. Leticia, forest edge,
Williams 3057. La Victoria, Williams 2519. Generally in the
tropics. "Cucarda," "flor Betun" (Williams).
Hibiscus Sabdariffa L. Sp. PI. 695. 1753; 116.
Glabrous or essentially except for the ciliate or setose bracts,
calyces and ovoid acute capsules; leaves oblong-lanceolate, some of
Flora of Peru 476
them more or less hastately lobed, many merely minutely serrulate;
peduncles shorter than calyces, these rigid-fleshy as the connate
simple sublinear bracts, both about equal and enclosing the fruits;
flowers pink or red, about 3 cm. long; capsules 2 cm. long; seeds
reniform, laciniate-lepidote.
The early fleshy calyces and bracts, often reddish, are rather
acid and not infrequently serve to flavor pleasantly beverages —
called "Karkardi" in the Old World (B.P.G.H.)— or for confitures,
and, according to Killip & Smith, the fruit is used for vinegar.
A common name in many parts of the world is "Roselle" or "Ro-
zelle," and the leaves according to the monographer are cooked
and eaten as sorrel.
Loreto: Mishuyacu, near Iquitos, Killip & Smith 29979. Gen-
erally in warm and tropical regions.
Hibiscus schizopetalus (Mast.) Hook. f. Bot. Mag. pi. 6524-
1880; 131. H. rosa-sinensis L. var. schizopetalus Mast. Gard. Chron.
282. 1879.
Similar to H. rosa-sinensis but in flower at least strikingly
distinct by the extremely long peduncles and the dissected petals,
many-lobed with small spatulate lobules, and the almost minute
bractlets. — It is very popular as a cultivated shrub in the tropics.
Huancayo: San Lorenzo, Rio Huanta to Rio Pastaza, Dennis
29218 (det. Killip).— Loreto: Iquitos, WiUiams 35^. In forest,
Caballo-Cocha, Williams 2^71. Edge of forest. La Victoria, Wil-
liams 2750. Clearing, Leticia, Williams 3061. East tropical Africa.
Hibiscus spiralis Cav. Icon. 2: 47, pi. 162. 1793; 90. Mal-
vaviscus Poeppigii (Spreng.) G. Don, Gen. Syst. 1: 475. 1831.
Achania Poeppigii Spreng. Syst. 3: 100. 1826. H. Poeppigii
(Spreng.) Giircke, Jahresb. Nat. Ver. Halle, 133. 1850. H. tubi-
florus DC. Prodr. 1: 447. 1824.
In general like H. brasiliensis; indument more or less developed,
stellate-hirsute; leaves often angulately 3-lobed, 1-4 cm. long,
obtuse or acutish, truncate or cordate at base, serrate; bractlets to
nearly 10 mm. long, little shorter than calyx, this with acuminate
lobes; corolla crimson, 2-2.5 cm. long, cylindric or the petals spread-
ing only at tip; capsules more than 1 cm. long; seeds lanate. —
Sjrnonymy after Hochreutiner, but Schery, Ann. Mo. Bot. Gard. 29:
231. 1942, suggested that H. Poeppigii may be MaXvaviscus arhoreus
L. var. m£xicanus Schlecht.
Peru(?): {Pavdn, fide Hochreutiner). Warm regions.
476 Field Museum of Natural History— Botany, Vol. XIII
Hibiscus tiliaceus L. Sp. PI. 694. 1753; 62. H. ahutiloides
Willd. Enum. Hort. Berol. 736. 1809, fide Kearney.
Tree, the branches conspicuously recurved by the bases of the
large stipules; leaves rotund-reniform, deeply cordate, subentire,
green and soon glabrous above, canescent beneath with a close
stellulate indument; bracts foliaceous, connate, more or less distant
from the calyx, this multidentate; petals nervose, broadly elliptic-
rotund, 4-5 cm. long, nearly as wide, yellowish to pinkish, harshly
puberulent; fruit ellipsoid, pseudo-partitioned, the many biseriate
seeds reniform. — A large shrub or small tree with ligneous capsules
and carpels incompletely divided longitudinally by a septum that
tends to split into two membranes; sometimes regarded as generi-
cally distinct, Pariti Adans. or Paritium St. Hil. Killip and Smith
(2981), as Ruiz and Pavon, found the species cultivated, the former
at Iquitos.
Tumbez: Inner edge of Mangrove, Condesa Island, Rio Tumbez,
Raimondi (det. Ulbrich). All tropical regions.
7. ABELMOSGHUS Medic.
Like Hibiscus but at most subligneous and with an irregularly
2-3-lobed spathaceous calyx that is deciduous before the fruit
matures. — The calyx actually is adnate at base to corolla so that it
falls with the latter and stamens (Hochreutiner, Candollea 2: 83-85.
1924).
Okra or gumbo, A. esculentus (L.) Moench. and A. moschatus
Medic, of the Old World, are popular in the New, particularly in
warmer regions, and sometimes are found persisting after cultiva-
tion has been abandoned.
Abelmoschus moschatus Medic, Malv. 46. 1787. Hibiscus
Abelmoschus L. Sp. PI. 696. 1753.
Green but rather conspicuously spreading-hirsute, often a meter
or so high; leaves variable, more or less palmately lobed, caudate,
hastate, often strikingly when the lobes are narrow, the basal widely
spreading, coarsely crenate-serrate, sometimes merely angulately-
lobed on the same branch; flowers often sulphur-yellow, 5 cm. or
more long; bracts linear, 6 or more, much shorter than the lance-
ovoid long-pedunculate capsules. — A. esculentus (L.) Moench. differs
particularly in having ovoid long-attenuate capsules, truncate at
base, shortly peduncled. It probably also occurs as an escape from
y
Flora op Peru 477
gardens. Determinations by Standley, except as noted. Illustrated,
Pflanzenfam. 3, Abt. 6: 49.
Loreto: Clearing, Peila Blanca, Rio Itaya, Killip & Smith 29669.
Florida, King 20U1. Rio Nanay, WiUiams S52 (det. Ulbrich).
Puerto Arturo, edge of forest, Williams 50^5. Pasture weed. La
Victoria, Williams 26U5; 2735; 2761. Old Worid tropics. "Aya
murillu" (Williams).
8. CIENFUEGOSIA Cav.
Fugosia Juss. Gen. PI. 274. 1789.
Reference: Hutchinson, New Phytol. 46: 125-131. 1947.
Similar to the related Hibiscus but the 3-5 bractlets narrow and
minute, often deciduous, rarely wanting, the calyx deeply 5-cleft, the
3-4-celled ovary with 3-many ovules in each cell and the apically
clavate style 3-4 sulcate or with 3-4 clavate short stigmatose
branches. Oil glands seriate. Fruit a loculicidal capsule, the walls
becoming dry, brittle. Seeds obovoid-globose, often pubescent.
— Oil glands, according to Hochreutiner, are always present.
Cienfuegosia heterophylla (Vent.) Garcke, Bonplandia 8: 148.
1860. Redoutea heterophylla Vent. Descr. PI. Gels. 11, pi 11. 1800.
Redoutea tripartita HBK. Nov. Gen. & Sp. 5: 293. 1822. Fugosia
tripartita (HBK.) Steud. Nom. end. ed. 2, 1: 649. 1840.
Flowering as an annual but often more enduring, the slender
acutely angled stems sometimes several dm. tall, nearly glabrous,
the minute stellate trichomes much dispersed; leaves variable, ovate
to lanceolate, shallowly to deeply lobed or tripartite, often 3-5 cm.
long, 0.5-3 cm. wide; pedicels in the upper axils, about 4 cm. long
or longer, enlarged beneath the gland-dotted filiform-caudate lobed
calyx; flowers yellow, reddish-brown at base, 2-4 cm. long; seeds
white or fulvous tomentose, the cottony indument to about 10 mm.
long. — According to Svenson the Peruvian plant differs from the
description of Ventenat in its pubescent style, broader calyx lobes;
he suggests that FtLgosia cuneata Benth., Bot. Voy. Sulphur 68.
1844, from Guayaquil, described as procumbent, may be a weak
state. Hutchinson however separates it as an herb with entire or
shallowly divided leaves, characters not apparently significant but
he also found the calyx more deeply divided; the plant of southern
Ecuador and Peru and probably northwards seems at most to be
a variant, for convenience var. cuneata (Benth.) Macbr., comb.
478 Field Museum of Natural History— Botany, Vol. XIII
nov. Fugosia cuneata Benth. I.e. Incidentally the species name of
Bentham has not been transferred to Cienfuegosia as Hutchinson
failed to cite its publication. The species becomes shrubby at base.
F.M. Neg. 9790.
Piura: River gravels, La Brea (Haught & Svenson 1158U). Tu-
lara, Haught W- Cerro Prieto, Haught 191. Amotape Hills, Haught
37. — Amazonas(?): Maranon Valley, Weberhauer 621Jf; 155. —
Cajamarca: Ja^n de Bracamoras, Bonpland (type, R. tripartita). \
To Brazil and Florida.
9. GOSSYPIUM L.
Reference: Hutchinson, Silow & Stephens, Evolution of Gossyp-
ium, i-xi, 1-160. 1947.
Tall herbs or shrubby, sometimes arborescent, more or less
dotted with black oil glands, the leaves usually palmately 3-9-lobed
to entire, the large flowers involucrate by 3 usually ample rarely
minute or caducous entire to incised bracts. Calyx cupulate,
truncate or shortly 5-dentate. Stamens many, united below. Ovary
cells 3-5, many-ovuled; style apically clavate, 5-sulcate and 5-
stigmatose. Capsules loculicidal, the seeds sub-globose or angled,
almost glabrous to (ordinarily) more or less densely lanate. — Besides
the above well-presented and informative book entitled "The
Evolution of Gossypium and the Differentation of the Cultivated
Cottons," see Guy Roberty's conscientious work, Candollea 7:
297-360. 1938; also 10: 345-398. 1946.
Cotton in Peru is of such interest that the reference work cited
(Oxford University Press) may be recommended here especially to
those concerned with cotton as a crop, for the book contains an
extensive bibliography and basic information of agricultural as well
as botanical significance.
The domestication of cotton and its association with man's
development is also presented, graphically, under the general con-
sideration of the evolution of the various species, and interesting
accounts of the cottons of both the New and the Old World,
Peruvian publications of the Est. Expl. Agric. de La Molina
(Peru) concerning the cultivation of cotton include: Insects of cotton,
J. F. Wille & 0. Beingolea, Informe 88. 1954; Fungus of cotton,
"Damping Off," by Jos^ M. Lamas & Consuelo Bazdn de Segura,
89; Control of cotton insects by use of maiz between rows and
insecticides, by Juan E. Simon F., 90; Insectos e insectidas en la ;
Flora of Peru 479
compaiia algodonera, by Juan E. Wille, Juan E. Simon, Juan E.
Gonsalez, 97. 1955; also, 25th Memorial Anual (1952) of cultivated
cotton within Peru.
Nine (or ten) of the fifteen known native species, all without
true lint hairs, are endemic to the western side of the Americas
and adjacent islands; five cultivated species are recognized, all with
lint hairs, but many forms of these have been described as wild or
semi-wild, probably however persisting from abandoned planta-
tions.
Leaves deeply lobed ; bractlets coarsely laciniate G. barbadense.
Leaves entire; bractlets finely laciniate G. Raimondii.
Gossypium barbadense L. Sp. PI. 693. 1753. G. peruvianum
Cav. Diss. 6: 313. pi. 168. 1788. G. vitifolium Lam. Encycl. 2: 135.
1786. G. barbadense L., subsp. vitifolium (Lam.) Roberty, Can-
dollea 10: 386. 1946.
Shrub or annual, glabrous or pubescent with long trichomes;
fruiting branches many-jointed; leaf-lobes 3-5, somewhat con-
stricted at base, rather long-acuminate, often plicate at the sinuses;
bractlets about as wide as long, with usually 10-15 narrowly lanceo-
late-acuminate teeth commonly exceeded by the subtubular corolla,
not much expanding; stamen tube long, the anthers evenly and
closely disposed; stigmas never spreading, often connate; carpels
ordinarily 3.5-6 cm. long, 3-(4)-celled, ovoid, acute, glandular
punctate, the sutures glabrous; seeds usually 5-8 per cell, with
(in cultivation) abundant even lint, sometimes also more or less
tomentulose. — A wild type with sparse lint was found and recorded
by Boza.
Inca cultivations of this species gave place to those of Indian
tribes and from these a commercial crop has been re-established in
the coastal valleys of Peru, the dominant variety, Tanguis, being
perennial, wilt-resistant; but the modem trend is in favor of annual
cottons (after Boza, as given by Hutchinson, et al., I.e. 102). Egyp-
tian cotton of commerce may be annual varieties descended from
hybrids of Sea Island with G. barbadense. Roberty in his revision,
CandoUea, I.e. considers G. peruvianum as a species which may be
useful in some types of investigations.
Tumbez: (Boza). — Piura: (fide Boza & Madoo). — Cajamarca:
Pacasmayo, Rose 18516 (det. Cook). — San Martin: Tarapoto,
480 Field Museum of Natural History — Botany, Vol. XIII
Williams 576^. — ^Ayacucho: Thickets, Aina, Killip & Smith 228It3.
— Junin: Thickets, La Merced, Killip & Smith 23562. San Ram6n,
Killip & Smith 2JI^88Jlt. Rio Pinedo, Killip & Smith 236J^3.—Cuzco:
Valle de Lares, Hacienda Pabellon, Herrera 789. Valle de Santa
Ana, Cook & Gilbert 1^99. San Miguel, Cook & Gilbert 1023.—
Loreto: Clearing, Puerto Arturo, Killip & Smith 2786U. Cliff
edge. La Victoria, Williams 3103. Lower Rio Nanay, Williams
351; J^j^. Yurimaguas, Killip & Smith 280^^. Northwestern
Argentina and northwards in tropical America. "Algodon," "utju,"
"ampi" (Campa), "uchto" (Cook and Gilbert).
Gossypium Raimondii Ulbr. Notizbl. Bot. Gart. Berlin 11: 548.
1932. G. Klotzschianum Anderss. subsp. Raimondii (Ulbr.) Roberty,
Candollea 13: 29. 1950.
Branches terete or subangled, the younger as the promptly
caducous subulate stipules, these 6 mm. long, grayish-tomentose;
petioles erect, 3-5 cm. long, densely and softly tomentose as the
truncate or subcordate based acute or subacuminate entire leaves
beneath, these glabrescent in age above except the 5-7 prominent
nerves, 8-14 cm. long, 9-12 cm. wide; involucral bracts semi-
orbicular, nearly 3 cm. long, 1.5-2 cm. wide with subulate-linear
lacinulae to 2 cm. long, tomentose as the campanulate calyx; corolla
15-16 mm. across, the obcordate-cuneate petals especially toward
the base sparsely tomentose, glabrous within with more punctiform
glands, dark purple toward the base as the conical stamen tube, this
1 cm. long, filaments long, the upper longer than those below;
capsules acuminate, the sutures with a line of trichomes between the
2 rows of seeds, these 4-8 per cell, densely pubescent with greenish
trichomes. — Apparently related to G. Klotzschianum Anderss. of the
Galapagos with glabrous branches and less tomentose more or less
3-lobed leaves, larger involucres, smaller flowers; and the glands
present on calyces and leaves of G. peruvianum Cav. are here present
only on the petals (Ulbrich). See Boza & Madoo, Est. Exp. Agric.
Molina, Peru, Bol. 22. 1941, for a study of this species and excellent
illustrations. Roberty's disposition may be correct or logical but
for this work it is useful to consider the Peruvian plant as a distinct
species.
Cajamarca: Ascope to Cascas, Raimondi 326, type. Playa del
Rio de Santa Ana o Jaguey, Raimondi. Near Cascas, Raimondi. —
Libertad: Near dry stream bed among Prosopis trees. Hacienda
Chiclin, West 8081 (det. Kearney).
Flora of Peru 481
10. MALVA Linn.
Annual or biennial, usually somewhat hirsute, often procumbent
or erect-ascending herbs, the leaves frequently angulately lobed or
dissected, the flowers solitary or fasciculate in the axils, sessile or
peduncled, rarely borne in terminal racemes. Bractlets 3, distinct.
Calyx 5-parted, at least medially. Petals emarginate, rarely den-
ticulate. Anthers clustered at the top of the filament tube. Ovary
cells many, 1-ovuled, the ovule erect or ascending. Style branches
7 or more, stigmatic longitudinally within. Fruit circular, more or
less flattened, the many erostrate indehiscent uniseriate carpels
parting at maturity, the seed ascending.
Flowers pedicelled M. parviflora.
Flowers sessile or subsessile M. verticiUata.
Malva parviflora L. Amoen. Acad. 3: 416. 1756.
Erect divaricate-branched annual or biennial, a dm. to some-
times 2 meters high, glabrous or sparsely stellate pubescent; leaves
cordate-suborbicular, shallowly 5-7-lobed, dentate-crenate; pedicels
2-10 mm. long, usually clustered in the axils, slender; bractlets
linear-lanceolate, 3-5 mm. long, the pubescent calyx 4-6 mm. long
at anthesis but spreading in fruit to form a rotate-scarious disk
12-16 mm. wide, the lobes deltoid-ovate; petals obovate, emarginate,
4-6 mm. long, glabrous, white except for purplish tips and veins;
carpels 8-12, dorsally reticulate, dentate in angles, puberulent as the
seed. — The "Cheeses" or "Cheese-weed" of English-speaking
children. The similar M. rotundifolia L. is a much branched pro-
cumbent weed with mostly solitary flowers, villous petal-claws, re-
ticulate acutely margined carpels, while the simulating M. neglecta
Wallr. has smooth rounded carpels. Another weed-like species to be
expected is M. nicaeensis All. marked by ovate bractlets.
Cuzco : Hacienda Macju, Pampa de Anta {Herrera 652). "Malva
sylvestre." Almost cosmopolitan.
Malva verticiUata L. Sp. PI. 689. 1753.
Erect, branching, glabrous or pubescent, petioles elongate, about
as long as the leaves, these cordate-suborbicular, 5-6-lobate, 2.5-7
(12) cm. long; flowers subsessile, densely verticillate; bractlets
linear-lanceolate, 6 mm. long; calyx lobes ovate, acute, about 8 mm.
long but accrescent; petals 6-12 mm. long; carpels 10-12, trans-
482 Field Museum of Natural History— Botany, Vol. XIII
versely rugose dorsally, reticulate laterally. — Apparently like M.
parviflora except for the densely clustered flowers.
Leaves used as a poultice (Mexia) .
Hudnuco: Chinchao, 2,200 meters, Mexia 0^^150 (det. Johnston).
"Malva crespa" (Mexia). Widely distributed.
11. WISSADULA Medic.
Reference: R. E. Fries, Sv. Vet. Akad. Handl. ser. 2, 43, no. 4:
1-114. 1908.
Like Ahutilon but carpels more or less completely divided by
horizontal or oblique constriction of lateral walls; flowers, at least
in Peru, about half a cm. long except W. disperma (1 cm.) and W.
stellata (1-2 cm.).
Leaves obviously and unevenly dentate or repand-dentate; petals,
about 8 mm. long or longer unless W. Pavonii.
Carpels muticous; pedicels short W. Pavonii. ;
Carpels aristate; pedicels 2 (-4 in fruit) cm. long W. disperma.
Leaves entire or finely and evenly, often obscurely crenulate-ser-i
rulate or -repand; petals shorter than 8 mm. except W. stellata.
Corolla 8-10 mm. long or longer, yellow; leaves velvety both sides.
W. stellata. .
Corolla 3-5 (7) mm. long; leaves various.
Flowers yellow or brownish-red, sometimes light yellow.
Leaves velvety both sides unless rarely in age, cordate.
Leaves unless uppermost at least obscurely or remotely
crenate-serrulate W. fuscorosea. ,
Leaves all entire.
Carpels soon longer than calyx.
Peduncles and calyces setose-stellate and tomentose.
W. suhpeltata\
Peduncles and calyces puberulent . . .W. hernandioides^
Carpels not or barely exceeding calyx W. microcarpa\
Leaves except nerves soon glabrous or subglabrous above,
rounded or little cordate at base W. excelsior 1
Flowers white or purplish W. zeylanica]
Wissadula disperma Hochr. Ann. Cons. Jard. Bot. Gten^ve 20:
114. 1917. Pseudabutilon Hitchcockii Ulbr. Notizbl. Bot. Gartj
Berlin 11: 522. 1932?
Flora of Peru 488
Glandular-villous, especially the cylindrical somewhat zigzag
stems, petioles, these 2-6 cm. long or longer, peduncles (to 2 cm. or
4 in fruit, medially articulate), calyces without, and obconic fruits;
stipules subulate, to 6 mm. long; leaves broadly ovate, deeply cor-
date, abruptly and acutely acuminate (acumen to 2 cm. long),
unevenly and coarsely dentate or sinuate-dentate, often 5-8 cm.
long, 1.5-3 cm. wide, velvety both sides, densely villous- tomentose
beneath, less so and slightly glandular above, palmately 9-nerved
at base; leaves reduced above; calyx cupulate, glabrous toward base
within the nectary conspicuous, the lobes elongate, 8 mm. long,
fruiting cal)rx nearly 1.5 cm. long; petals about 1 cm. long, basally
pilose on margins; stamen column 2.5 mm. long, glabrous basally,
densely pilose above; carpels 5, bicomiculate, 8-9 mm. long without
beaks these about 3 nmi. long, the transverse fold little developed,
with 2 superposed lenticular pilose seeds, the upper erect, the lower
pendent. — Very remarkable in that it constitutes an obvious con-
nection between Wissadula and Abutilon (author). Here would be
sought the apparently similar Pseudabutilon Hitchcockii Ult^r.
Notizbl. Bot. Gart. Berlin 11: 522. 1932 of Guayaquil "which seems
to be very well attributed by you to W. disperma but I have not
seen the plant of Hitchcock" (B.P.G.H.). F.M. Neg. 23755.
Peru(?): without data, Pavdn, type.
Wissadula excelsior (Cav.) Presl, Rel. Haenk. 2: 118. 1835; 44.
Sida excelsior Cav. Diss. 1 : 27, pi. 5. 1785. Abutilon ferrugineum
HBK. Nov. Gen. & Sp. 5: 271. 1822?
Virgate, little branched, sometimes a meter tall, the terete upper
stems, petioles, leaf nerves beneath and younger panicles notably
ferrugineous with a partly stipitate-stellate indument; leaves entire,
ovate, rounded or lightly cordate at base, acuminate, soon green
and glabrescent or glabrous above, canescent between the nerves
and reticulate veins beneath, often 6-12 cm. long, about half as
wide; panicles rather oblong-ovoid, the peduncles to 1 cm. long or
much shorter; calyx ovate at base, puberulent and rusty hirsute,
2.5-3.5 mm. long; petals yellowish, 3.5-4 mm. long; carpels 5,
maturing dark, membranous, fragile, puberulent, about 8 mm. long,
the beak 0.5-1 mm. long; seeds pulverulent, hilum sparsely pilose,
2 mm. long. — Fries has discussed the identity of A. ferrugineum,
I.e. 92-93; however, it is from Yoja, Ecuador, and a later name.
The native name "Palo de Balsas" noted by Cavanilles was probably
a mistake. Determinations by Standley. F.M. Negs. 29799
(Jussieu); 32628 (Poeppig).
484 Field Museum of Natural History— Botany, Vol. XIII
San Martin: Juanjul, Klug Jt387. — Junin: Near Peren^ Bridge,
Killip & Smith 253 J^3. Rio Maraiion Valley, Dennis 29127 (det.
Killip).— Hudnuco: Tingo Maria (Asplund 12067; 12311, det.
Fries). Zepelacio, Klug 36^7. Yurimaguas, Poeppig 2W; Williams
39H. Mouth of the Ucayali, Tessmann 3090. Without locality,^
Jos. de Jussieu, type. Ecuador; Brazil.
Wissadula fuscorosea Ulbr. Bot. Jahrb. 54: Beibl. 117: 59. 1916.
Tomentose or more or less lanate-tomentose including the ample
paniculate inflorescence of brownish-red flowers; stipules lanceolate,
7 mm. long; petioles subequaling the leaf-blades; lower leaves about
orbicular, deeply cordate, more or less acuminate, to 10 cm. long,
6-7 cm. wide, the upper ovate to lanceolate, obscurely or distinctly
serrate, the 7-9 palmate nerves and reticulate veins prominent
beneath; pedicels 1-2 mm. long, to 5 mm. after anthesis; calyx
glabrous within, 4.5 mm. long, the ovate acuminate lobes 2 mm.
long; corolla spreading, 6 mm. long, connate 1 mm. with stamen
tube, this subconoid, glabrous; styles 3-4 mm. long, with a few
scattered trichomes, stigmas globose, glabrous; fruits obconoid-
globose, the calyx broken to base, to 6 mm. high, the 4 tomentulose
apiculate carpels 2-3-seeded, the seeds in the upper cell mostly
collaterally binate, one in the lower cavity, densely lanate near the
hilum. — To 2 meters high. Resembles W. contracta (Link) R. E.
Fries and W. densiflora R. E. Fries, both with shorter pedicelled
yellow flowers. F.M. Neg. 9299.
Huancavelica: Grasslands, Prov. Tayacaja, left of Rio San
Bernardo, Weberbauer 6556, type.
Wissadula hernandioides (L'H^r.) Garcke, Zeitschr. Naturw.
Halle 63: 122. 1890; 48. W. amplissima R. E. Fr., Sv. Vet. Akad.
ser. 2. 43, no. 4: 48. 1908, excl. syn., fide Fawcett & Rendle, Fl.
Jam. 5: 95. 1926. Sida hernandioides L'H^r. Stirp. 2: 121.
Branchlets, petioles and peduncles minutely stellate-tomentose
or finally glabrate as the leaves above or these more or less densely
and permanently canescent tomentose at least beneath, typically
deeply and narrowly cordate at base or apparently sometimes
openly cordate, entire or essentially, rotund-ovate or very broadly
ovate, sometimes rather abruptly but in general gradually acuminate,
variable as related species in size and length of petioles; flowers
axillary or mostly in diffuse terminal panicles, the peduncles slender,
soon 2-3 cm. long or longer, merely puberulent as the calyx, this
Flora op Peru 485
3-4 mm. long; corolla yellow, 4-6 mm. long, the spathulate petals
ciliolate at base; fruits 8-10 mm. in diameter, at maturity only
sparsely puberulent, the 4-5 carpels 7-8 mm. long, acuminate or
beaked, the beak 0.5-1.5 mm. long; seeds 3, subsimilar or diverse,
the upper globose-cordiform, punctate, minutely puberulent, the
lower subovoid, pilose especially at the hilum. — Probably in Peru
and quite possibly the specimens referred to the similar W. sub"
peltata should rather be included here. Illustrated, Fries, I.e. pi. Ut
jig. 1 (plant); pi. 6, figs. 12-1 U (fruits).
Peru (probably). Tropical America; Africa.
Wissadula microcarpa R. E. Fr., Sv. Vet. Akad. Handl. ser.
2. 43, no. 4: 55. 1908.
Younger parts densely canescent with stellate-tomentose in-
dument, this shorter on the leaves, these greenish above in age,
paler beneath, rotund-ovate, entire, deeply cordate, acuminate, to
7.5 cm. long, about 6 cm. wide or the lower probably larger, the
nerves and reticulation prominent beneath; petioles (except those
of the upper reduced leaves) 3.5-4 cm. long; inflorescence terminal,
many-flowered, ample, the branches spreading, the puberulent
peduncles 5-12 mm. long; calyx tomentulose and with some larger
yellowish stellate trichomes, 3.5-4 mm. long, the ovate-triangular
acute lobes 2-2.5 mm. wide; corolla lemon yellow, about 6 mm. long,
the petals pilose at base as the very short stamen tube; fruit sub-
globose, the 5 puberulent carpels included in the calyx, acute or
apiculate but not rostrate; seeds 3, similar, black, subreniform,
sparsely hirsute, to 2 nmi. long. — Illustrated, Fries, I.e. pi. 6, figs. 15,
16 (fruit). F.M. Neg. 9302.
Loreto: Salinas de Tilluana on the Huallaga, Ule 6710, type. —
Hudnuco: Chulque, 1,700 meters, Mexia 0^107 (det. Johnston).
"Utquicha" (Mexia).
Wissadula Pavonii Hochr. Ann. Cons. Jard. Bot. Gendve 20:
113. 1917.
Younger stems irregularly angled, densely tomentose-hirsute as
the petioles, these 1.5-4 cm. long, peduncles (1-2 cm.) and calyces
without; leaves broadly ovate, deeply cordate at base, acute or
subacuminate, about 5 cm. long, 3 cm. wide, stellate-pilose but green
above, velvety ashy-tomentose below and unevenly and rather
coarsely dentate, the 7-9 palmate nerves prominent beneath;
486 Field Museum of Natural History— Botany, Vol. XIII
flowers solitary, axillary and crowded at ends of secondary branch-
lets; calyx 4, in fruit 5 mm. long, the ovate lobes glabrous within
except tips; petals high-connate with short pilose stamen column,
the free part about 6 mm. long, the shorter stamens sparsely pilose;
fruit subglobose, about 8 mm. in diameter, the 5 subreniform
muticous carpels tomentulose, about 5 mm. high, 3 mm. broad, with
2 upper collateral, 1 lower seed, all brown-black, softly pilose. —
Allied to W. decora Sp. Moore and W. sordida Hochr. but distinguish-
able from both by the indument and by the tendency of some leaves
to be trilobate by the presence of 2 slightly larger teeth (author).
Here might be sought W. andina Britton, Bull. Torrey Club 16:
153. 1889; 76, Bolivian, with corolla 10-12 mm. long and only 3 or 4
carpels. According to Fries, Mathews 50U from Cuesta de Pur-
rochuco does not belong to W. andina as indicated by Baker; the
collection has not been seen. F.M. Neg. 23758.
Lima: Chancay, (Ruiz &) Pav6n, type.
Wissadula Stella ta (Cav.) Schum. in Mart. Fl. Bras. 12, pt. 3:
445. 1891. W. nudiflora (L'H^r.) Benth. Bot. Voy. Sulphur 69. 1844;
65. Sida nudiflora L'H^r. Stirp. Nov. 123, pis. 59, 59h. 1789.
Abutilon nudiflorum (L'H^r.) Sweet, Hort. Brit. ed. 1. 53. 1827; ed.
2. 64. 1830. Sida stellata Cav. Diss. 1: 27. 1785.
Velvety yellowish stellate-tomentose to the pedicels, these as
calyces rusty stellulate-puberulent, the former 5-13 mm. long, the
latter in terminal lax subsimple or sparsely branched leafless panicles,
4-5 mm. long the broadly ovate acute lobes half as long; stipules
linear-filiform; petioles 2-7 cm. long; leaves broadly ovate or the
upper almost suborbicular but gradually acuminate, openly but not
widely cordate at base, entire or obscurely crenulate, prominently
nerved and somewhat reticulate beneath, greenish and impressed
reticulate above in age, commonly 5-10 cm. long, 3.5-6 cm. wide or
larger; corolla 1-2 cm. long, the spathulate-orbicular petals more
or less stellate-hirsute basally, as often the 1 mm. long stamen tube;
carpels 5, acute, puberulent, 5 mm. long, the upper seed slightly,
lower densely, especially hilum, stellulate. — A 1-2 meter shrub.
F.M. Negs. 9303; 7987 (as Sida periplocifolia, var., ined.).
San Martin: Juanjui, Klug ^S30 (det. Standley). — Hudnuco:
Common near Hudnuco, 20^; Sawada P6U (det. Ulbrich, W.
microcarpa) ; Ruiz & Pavdn; Domhey, type; Stork & Horton 9^06
(det. Standley, W. periplocifolia).
Flora of Peru 487
Wissadula subpeltata (Ktze.) R. E. Fr. Sv. Vet. Akad. Handl.
ser. 2, 43, no. 4: 56. 1908. AbiUilon amplissimum (L.) Ktze. var.
subpeltata Ktze. Rev. Gen. 3, pt. 2: 17. 1898.
Canescent puberulent-tomentose especially the leaves beneath,
their upper surfaces greenish-sericeous; stipules linear-lanceolate;
petioles to 1.5 cm. long; leaves broadly ovate the uppermost almost
suborbicular but gradually acuminate, deeply and narrowly cordate
the basal lobes often overlapping, prominently nervose and reticu-
late beneath, entire, various as all species in size, those of the
flowering branches often only a few cm. wide, the lower to 1.5 dm.
long and wide; panicles ample with slender spreading branches;
peduncles puberulent and as calyces (3.5 mm. long) early at least
with a few simple and stellate rigid trichomes, 2-4 cm. long in fruit;
petals cuneate, pilose below as stamen tube, about 5 mm. long;
fruit subglobose, dark brown or blackish, about 8 mm. wide, the
5 carpels glabrous obovoid, 7 mm. long,'beak 0.5 mm. long; seeds 3,
globose-reniform, the upper strongly rugose, glabrous, the lower
hirsute especially at the hilimi. — Characters of pubescence and seeds
not developed as indicated but apparently determinations correct.
However, the older specimens of Cook and Gilbert with coarser
trichomes nearly lacking seem very much like W. hernandioides,
which compare, and perhaps would better be placed there or in
W. boliviana R. E. Fr., 40, with openly cordate leaves, 4 carpels
(always only 4?); these forms seem to crowd each other and their
stabihty may be open to question. Illustrated, Fries, I.e. pis. 5
(plant); 6, fig. 27 (fruit); 7, fig. 15 (androecium).
San Martin: Tarapoto, Williams 5806 (det. Ulbrich).— Cuzco:
Santa Ana, Cook & Gilbert 1509? To eastern Brazil and Argentina.
Wissadula zeylanica Medic. Malv. 25. 1787; 32. Sida peri-
plocifolia Presl, Rel. Haenk. 2: 117. 1835, not L. as to herb.
Younger parts more or less brownish tomentulose and stellate
pubescent, the leaves especially so beneath, sometimes glabrescent
above, membranous or firmer, ovate- or lanceolate-triangular,
gradually acuminate, truncate or openly cordate, entire, the largest
lower to a dm. long or longer, nearly half as wide, reduced upwards,
the petioles too becoming much shorter, even the lowest at most
3 cm. long; inflorescence (Peru) terminal, lax, more or less ample,
the slender pedicels in fruit to 5 cm. long, often a few flowers solitary
in the upper axils; caljrx 2-3 mm. long, pulverulent or glabrous, the
ovate acute lobes half as long; petals about 5 mm. long, white or
violet-tinted; fruit 8-10, the pulverulent carpels (5) 7-8 mm. long.
488 Field Museum op Natural History— Botany, Vol. XIII
with beak 0.5-1 mm. long; upper 2 seeds subglabrous the lower one
densely hirsute especially the hilum. — The South American form,
designated W. periplocifolia Presl var. gracillima R. E. Fr., I.e., 34,
has a tendency to more cordate lower leaves, glabrate above, more
ample panicles and carpel beaks to 1 mm. long. Still more distinct
is W. diffusa R. E. Fr., I.e., 37, from near Guayaquil, the carpel
beaks 2-3 mm. long, but scarcely, as the author suggests, more
than a variety. Distinct but similar and occurring within Peru
is W. hernandioides (L'H^r.) Garcke (W. amplissima R. E. Fries,
I.e., 48, fide Fawcett & Rendle) ; the leaves are narrowly and deeply
cordate at base, the smaller petals yellow.
Junin: Cabello, river canyon, 1830 (det. Hochreutiner) . Puerto
Yessup, stream bed, Killip & Smith 26316 (det. Standley). Ceylon;
Africa; tropical America.
12. PSEUDABUTILON R. E. Fries
Reference: Fries, Sv. Vet. Akad. Handl. ser. 2. 43, no. 4: 96-108.
1908.
Like Wissadula but the carpels more or less divided by the
horizontal projection of the dorsal wall, that is, the two cavities are
separated by an endoglossum, a membranous tongue-like organ,
instead of by a horizontal fold formed by constriction of the lateral
walls. — The carpels are 5-11 and the flowers at least in Peru to
about 8 mm. long. In Wissadula the endoglossum is developed in
varying degrees, an impression gained particularly from Kearney's
interpretation, Amer. Midi. Nat. 46: 115. 1951, but Hochreutiner
has called to my attention his belief that the character is important;
however, it is characteristic also for Modiola, belonging to a different
generic alliance, and therefore may not be a significant indicator of
generic relationship.
Leaves ovate, long-acuminate or acute.
Petals about 8 mm. long; leaves as calyx black punctate.
P. nigripunctulatum.
Petals about 5 mm. long; leaves not black punctate.
P. Weberhaueri.
Leaves suborbicular, abruptly acuminate P. spicatum.
Pseudabutilon nigripunctulatum (Ulbr.) R. E. Fries, Sv.
Vet. Akad. Handl. ser. 3. 24, no. 2: 11. 1947. Abutilon nigripunctu-
latum Ulbr. Bot. Jahrb. 54, Beibl. 117: 57. 1916.
Flora of Peru 489
Branchlet tips and cordate-ovate leaves especially beneath
black puncticulate and also more or less scabrous with minute
stellate trichomes, the filiform stipules (3-4 mm. long), petioles
(5-20 mm. long) and calyces without somewhat tomentose; leaves
rather long-acuminate, 3-5 cm. long, 1.5-3 cm. wide, crenate-
serrate, yellowish-green and black punctulate beneath, the nerves
prominent; flowers white, axillary or in the axils of a highly varied
inflorescence, the slender peduncles 2-3 cm. long, articulate about
3 mm. below the calyx, this cupulate, 5 mm. long, the tomentose
lobes 2 mm. long, black punctulate at base, glabrous within; petals
suborbicular, glabrous, 8 mm. long; stamen tube 4-5 mm. long,
conoid at base; styles 3 mm. long, connate 1 mm., the capituliform
stigmas globose; carpels 6, grayish subtomentose, shortly aristate,
not splitting apart, the 2-3 seeds tomentulose near the hilum.
— FVies, I.e., noted that the species has the endoglossum of Pseuda-
btUUon. Shrub about a meter high with an aromatic fragrance.
F.M. Neg. 9281.
Lima: San Bartolom^, 1,500 meters, Weberhauer 5301, type.
— Apurfmac: Rio Pachachuca, Goodspeed Ezped. 1052U (det.
Standley).
Pseudabutilon spicatum (HBK.) R. E. Fries, Sv. Vet. Akad.
Handl. ser. 2. 43, no. 4: 98. 1908. Wissadula spicata (HBK.) Presl,
Rel. Haenk. 2: 117. 1835. Abutilon spicatum HBK. Nov. Gen. &
Sp. 5: 271. 1822.
More or less suffrutescent, often a meter or so high, canescent
tomentose but the leaves green above at maturity, the older stems
glabrescent; stipules tardily deciduous, linear-subulate, to 1 cm.
long; petioles 5-15 cm. long, but upper leaves often subsessile;
leaves suborbicular, deeply cordate, abruptly acuminate, 5-15 cm.
long, about as wide, or larger, dentate, the 7-9 primary nerves most
marked beneath; panicles to 4 dm. long, leafless, with short appressed
subspicate branchlets, the fruiting peduncles about 3 mm. long;
calyx 3-4 mm. long, lobes acute; petals retuse, glabrous except basal
margins, 6-7 mm. long; fruit 5-merous, turbinate, 7-8 mm. in
diameter, the carpels stellate-pilose, acute, 4-5 mm. long; seeds
globose-cordiform, puberulent except glabrous hilum, scarcely 2
mm. long and broad, 2 collateral in upper cavity, 1 in lower. —
Illustrated, Mart. Fl. Bras. 12, pt. 3: pi. 78; Fries, I.e., pi. 7 (carpel).
San Martin: Juanjul, Klug Jt2Jt5 (det. Standley). Tarapoto,
WiUxams 6097 (det. Ulbrich).— Junin: Colonia Peren^, KiUip &
490 Field Museum of Natural History— Botany, Vol. XIII
Smith 25006 (det. Ulbrich). Bolivia to Mexico and the West
Indies.
Pseudabutiion Weberbaueri Ulbr. Bot. Jahrb. 54: Beibl. 117:
60. 1916.
Yellowish tomentose, even the axillary subpaniculate inflo-
rescences; stipules linear-lanceolate, 2 mm. long; petioles angulate,
1.5-3 cm. long; leaves ovate, deeply cordate, acute, 4-8 cm. long,
3.5-5 cm. wide, crenulate-serrate, the 7-8 nerves and reticulate
veins prominent beneath; pedicels 3-8 mm. long, articulate 2-3 mm.
below the calyx, this cupulate, 3 mm. long, the broadly ovate
acuminate lobes 2 mm. long; corolla yellow, spreading, the 10-nerved
oval obtuse petals 4.5-5 mm. long, nearly 1 mm. connate with the
stamen tube, this stellate above; styles with capitate stigmas gla-
brous, free; fruits cylindric-globose, calyx more or less reflexed,
5 mm. high, with usually 8 compressed ovoid carpels, nearly divided
dorsally below the middle, each cavity with 1 cordiform seed, its
sparse indument simple, and stellulate trichomes. — Tjrpe 2 meters
high among shrubs and small trees. Belongs in the neighborhood
of W. paniculata Rose with 6-11 carpels.
Cajamarca: Shumba Valley, 700 meters, Weberbauer 6169, type.
13. ABUTILON Adans.
Gayoides (Gray) Small, Fl. S. E. U. S. 764, 1335. 1903. Bogen-
hardia Rohb. Repert. Gen. PI. 1: 200. 1841.
Reference: Schumann, Mart. Fl. Bras. 12, pt. 3: 253-456. 1891;
and Pflanzenfam., Nachtrag 1: 235-239. 1897.
Herbs or shrubs generally with somewhat cordate angled or
lobed leaves and mostly axillary often showy flowers. Involucel
wanting. Ovary cells 5-many, normally 2-9-0 vuled (cf. A. pulveru-
lentum, A. Weberbaueri); ovules erect-ascending except 1-ovuled
species. Styles filiform or clavate, stigmatose apically. Carpels
slightly to much inflated if at all usually acute and mucronate to
aristate, the walls thin, firm-membranous or coriaceous, various in
dehiscence but always primarily loculicidal sometimes early sep-
ticidal; in most species the carpel halves fall as units or when they
remain attached it is the halves of adjacent carpels that form the
pair (Hochreutiner, Ann. Cons. Jard. Bot. Geneve 21: 364-365.
1920). — The 1-ovuled species as noted may be referable to Sida or
Bastardia (Kearney, in herb.). However the same variation has
Flora op Peru 491
been accepted for WissddiUa. Maybe these questionable species
are connecting entities in this remarkably close-knit family and are
most conveniently placed in the genus in which their general facies
seems least out of place. A. dianthum Presl, accredited in litt. to
Peru, came from Ecuador. Svenson, Amer. Joum. Bot. 33: 463.
1946, following Fawcett & Rendle retained the single species of
Gayoides in AbtUilon a procedure that has been emphatically en-
dorsed by Hochreutiner who has observed to me: it is distinguish-
able only by the consistency of the carpel wall which is very variable
in the genus.
Flowers large, normally (full-grown) 3-4 cm. long.
Petals soon spreading, finally reflexing, narrow.
Leaves softly tomentose on both sides; stems short-stellate.
A. reflexum.
Leaves soon scabrous-stellate or glabrous above; stems pilose
or hirsute.
Leaves coarsely crenate; peduncles to 7 cm. long.
A. lateritium.
Leaves subentire; peduncles to 2 dm. long. . .A. pedunculare.
Petals erect or tardily spreading above, broad.
Leaves distinctly lobed A. striatum.
Leaves not lobed.
Indument of broadly ovate or subrotund leaves velvety on
both sides.
Peduncles longer than subtending leaves A. Umgipes.
Peduncles shorter than subtending leaves, . . .A. arboreum.
Indument of triangular-ovate leaves scabrous above, soon
sparse or lacking A. sylvaticum.
Flowers rarely little longer than 2 cm., usually shorter.
Indument of branchlets in part spreading, pilose-hirsute; flowers
mostly or all solitary.
Flowers yellow; carpels 1-3-ovulate or -seeded.
Petals erect.
Indument in part viscid-glandular.
Carpels muticous or nearly, 3-seeded; leaves acute or
obtuse A. hirtum.
Carpels rostrate, 1-2-seeded; leaves shortly acuminate.
A. WeberbaiLeri, A. cor datum.
492 Field Museum of Natural History— Botany, Vol. XIII
Indument neither viscid nor glandular.
Carpel walls firm; petals 1.5 cm. long or longer.
Calyx-lobes acute; carpels muticous or nearly, ovules
as seeds 3 A. indicum.
Calyx-lobes acuminate; carpels cusped, ovules as
seeds about 4 A. mollissimum.
Carpel walls thin, finally much inflated; petals to 12
mm. long A. crispum.
Petals soon reflexed A. giganteum.
Flowers not yellow; carpels 6-8-0 vuled A. pauciflorum.
Indument of branchlets stellate-puberulent to lanate, often minute
or lacking, or flowers mostly not solitary.
Flowers not yellow, solitary except A. pulverulentum.
Petals promptly reflexed; indument soon scurfy-stellate.
A. arequipense.
Petals tardily if at all reflexed.
Flowers solitary on long slender pedicels A. piurense.
Flowers glomerate in efoliate cymes.
A. pulverulentum, A. cymosum.
Flowers yellow, in several-flowered inflorescences unless the
lowest, paniculate or in axils.
Branchlets lanate or velvety-tomentose.
Flowers paniculate A. ramiflorum.
Flowers in crowded cymes A. cymosum.
Branchlets scabrous stellate or glabrate, or also somewhat
villous.
Petals not reflexing.
Flowers mostly in several-flowered inflorescences; car-
pels 5-7 (11) A. umbellatum.
Flowers mostly axillary-clustered; carpels 8-9.
A. virgatum.
Petals reflexing after anthesis; uppermost flowering branch-
lets somewhat panicled A. giganteum.
Abutilon arboreum (L.) Sweet, Hort. Brit., ed. 1: 53. 1827.
Sida arhorea L.f. Suppl. 307. 1781. S. peruviana Juss. ex Cav. Diss.
1: 36. 1785. S. grandiflora Poir., Encycl. Suppl. 1: 31. 1810. S.
mollis Ortega, Decad. 5: 65. 1798, at least as to Peru.
Flora of Peru 493
Shrub or small tree, sometimes 6 meters high, the upper branches
and petioles sericeous-tomentose or -puberulent, sometimes also
pilose; leaves broadly ovate, deeply cordate, more or less acuminate,
softly tomentose both sides, usually about 1.5 dm. long and nearly
as wide, or the lower twice as large, the petioles about as long;
stipules subulate, acuminate, 1 cm. long or longer, tardily caducous;
peduncles commonly 1 dm. long or twice as long in fruit, sometimes
binate, always axillary, but frequently on an accessory branchlet;
cal)rx campanulate, canescent-sericeous, 1.5-2 cm. long, the oblong
triangular acute 3-nerved lobes tomentose within and without;
petals whitish, 3.5-4 cm. long, about 3 cm. wide at the retuse apex,
marginally pilose on the long claw; androecium 3 cm. long, the
stamens in 5 fascicles; ovary white- villous, the cells about 8-ovulate;
carpels chartaceous, ultimately 1.5 cm. long, 1 cm. broad, fer-
rugineous-tomentose dorsally; seeds 3 mm. in diameter, glabrous
except for the conspicuously villous hilum. — Synonjmiy after Schu-
mann; type of S. peruviana by Jos. de Jussieu without locality;
probably also S. arborea, the type locality not Africa as given by
Linnaeus f. (Schumann). Vargas noted the flowers as white. For
some reason most of the Peruvian collections have been referred to
A. molle (Orteg.) Sweet. Determinations by Kearney except as
noted. Illustrated, L'H^ritier, Stirp. Nov., pi. 63.
Junin: San Ram6n, Killip & Smith 2^802 (det. Ulbrich). Car-
papata, Killip & Smith 2^76 (det. Killip). — Hudnuco: Cuchero,
Dombey; Poeppig. — Cuzco: Ruins of Machupicchu, West 6U2J!t (det.
Johnston). — Apurlmac: Am pay, Vargas 788. — Cuzco: Valle de San
Miguel, Herrera 1985; 1986. Valle del Urubamba, Herrera 1570;
SSU7. San Miguel, Cook & Gilbert 920. Machupicchu, Vargas 783;
78U. Ollantay, Soukup 567. Near Cuzco, Soukup 138. "Rata-
rata" (Herrera).
Abutilon arequipense Ulbr. Bot. Jahrb. 54: Beibl. 117: 49.
1916.
Younger branches and leaves densely yellowish cinereous lanate-
tomentose, becoming scurfy-stellate or glabrescent, the gray bark
nodulose rugulose, the leaves cordate-oblong, obtuse or subacute,
2-3 cm. long, 13-18 mm. wide; stipules linear, 8 mm. long; petioles
5-10 mm. long, short-tomentose as peduncles, these 2.5-3 cm. long,
and calyces, these nearly 11 mm. long, the ovate cuspidate lobes
about 8 mm. long, 6 mm. wide at base; corolla violet, soon reflexed,
darker toward base within; petals suborbicular, to 13 mm. long,
494 Field Museum of Natural History— Botany, Vol. XIII
15 mm. wide, about 15-nerved; stamen tube lanate, 6 mm. long,
anthers globosely capitate; styles filiform, clavate, stigmas sub-
capitate; fruit subglobose, about 9 mm. across, carpels 9-10, oblong-
ovoid, 6 mm. high, 4 mm. broad, 2.5-3 mm. thick, tomentose except
laterally, the obliquely cordiform seeds fuscous-lanate. — Type
a 2-meter shrub. Related to A. cor datum with larger cordate leaves,
different pubescence on branches and petioles; styles, stigmas and
carpels suggest those of A. Seineri Ulbr. of Africa, and it appears
to me that there is a relationship; there are similar cases of alliance
between plants of South Africa and the Andes; cf. Engler, Sitzungsb.
Kgl. Preuss. Akad. Wiss. 20: 564. 1914 (Ulbrich).
Arequipa: Above Cotahuasi, 2,800 meters, Weberhauer 6863,
type.
Abutilon cordatum Garcke & Schum. Mart. Fl. Bras. 12, pt. 3:
369. 1891.
Slender terete flowering branches abundantly tomentose, with
stellate and simple glandular viscid trichomes above; stipules
linear-subulate, hirsute, promptly caducous; petioles 5-10 cm. long;
leaves cordate, rather shortly but acutely acuminate, repandly
dentate, 1-1.5 dm. long, about 1 dm. wide, softly tomentose on both
sides; flowers all axillary, solitary, on peduncles 2.5-3.5 cm. long,
articulate above the middle, sometimes with an accessory branchlet;
calyx campanulate, to 14 mm. long, the oblong triangular acuminate
lobes pubescent within and without with bulbous-based trichomes
and also stellate- tomentose; petals 13-15 mm. long, about 1 cm.
wide above, ciliolate-pilose toward the base, apparently yellow;
ovary globose villous, the cells biovulate. — Species distinct by the
two-ovulate cells and the stellate, simple and glandular indument,
intermixed (authors). It is really so; Baker f. made a mistake in
putting it into the uniovulate Abutilon (B.P.G.H.). — Type from
Guayaquil; in Asplund 7667 with more obtuse leaves, the calyx j
shorter than the corolla, the carpels are 9-10 mm. long, beaks 2.5-3'
mm. long, glabrous below, densely glandular-pubescent above, the
reniform sparsely pubescent seeds about 3 mm. in diameter (Fries).]
F.M. Neg. 9264.
Libertad: Prov. Patdz, Raimondi (det. Ulbrich). Ecuador.
Abutilon crispum (L.) Medic. Malv. 29. 1787. Gayoides cris-
pum (L.) Small, Fl. S.E. U.S. 764, 1335. 1903. Sida crispa L. Sp.
PI. 685. 1753. Bogenhardia crispa (L.) Kearney, Leafl. West. Bot.
7: 120. 1954.
Flora of Peru 495
Softly tomentose herb or half-shrub, the prostrate or sprawling
to suberect stems somewhat hirsute-pilose; stipules subulate, 5-7
mm. long, more or less persisting; leaves cordate, the upper sub-
sessile the lower long-petioled, all acute or shortly acuminate,
coarsely serrate; peduncles solitary or binate in the axils, articulate
above the middle; caljrx about 6 mm. long, the ovate lobes acuminate,
the rounded yellowish petals slightly to twice as long; fruit yellowish,
the inflated carpels about 12, usually 3-ovuled, or 1-ovuled (Svenson),
at maturity minutely pilose and setose, 10-15 mm. long. — Svenson,
besides noting the single seeded carpels of his collections, recorded
the flowers as minute, orange; however variable the species, this
local variant seems worthy of recognition to call attention to it if for
no other reason and may be recorded as var. Svensonii Macbr.,
var. nov., ovulis solitariis; floribus minutis intense flavibus. Illus-
trated, Fawcett & Rendle, Fl. Jam. 5: 99.
Piura: In shrubs along creek north of Talara, Horton 11596.
Amotape Hills, trailing {HaughX & Svenson IISJ^S, type, var.). —
Hudnuco: Trailing, stony hill, 3165. — Junin: Sandy valley floor,
La Merced, 5It5S. — Ayacucho: Aina, Killip & Smith 228 j^. —
Apurimac: Rio Pachachuca, prostrate-radiating in gravel. Good-
speed Ezped. 10517 (det. Standley). Abancay, Vargas 466 (det.
Standley). American and Old World tropics.
Abutilon cymosum Tr. & PI., Ann. Sci. Nat. s^r. 4. 17: 185.
1862. A. rufinerve Seem., Bot. Voy. Herald 83. 1853, not St. Hil.
A shrub about 2 meters high, the branches velvety with a reddish-
yellow tomentum; stipules linear, erect; petioles long; leaves cordate,
acutely acuminate, unequally serrate, thick, 5-nerved; cymes axil-
lary, often geminate, the rather long erect peduncles with 1-3 small
leaves at apex; flowers 5-15 or more in crowded cymes, the pedicels
sometimes much longer than the calyces, these cuspidate-lobed and
shorter than the erect yellow petals; carpels about 8, very acutely
subulate-rostrate, finally biparted, the 3 glabrous seeds sparsely
muricate-papillose. — Imperfectly known but according to Baker
occurring in Bolivia and if so, no doubt in Peru.
Peru (cf. note above). To Panama.
Abutilon giganteum (Jacq.) Sweet, Hort. Brit. ed. 1: 53. 1826.
Sida gigantea Jacq. Hort. Schoenbr, 2: 8, pi. lU. 1797.
Herb or shrub, the young branches and elongate petioles with
or without spreading pilose indument, the large round-ovate leaves
496 Field Museum of Natural History — Botany, Vol. XIII
stellately and simply pubescent above, tomentose beneath, the
trichomes on the 9 prominent nerves simple; flowers solitary, axil-
lary, accompanied by a flowering branchlet forming spreading
foliose panicles; calyx about 1 cm. long, the lanceolate lobes 1-nerved
within, the orange (or lilac and yellow according to Klug) petals
finally reflexed, somewhat longer, villous at base about the stamen-
tube; carpels 8-14, tomentose, rostrate, the 3 seeds tuberculate-
pubescent. — According to R. E. Fries the species occurs in two
forms as to pubescence, one lacking the spreading pilose trichomes.
San Martin: Juanjui, Klug 3887; If370 (both det. Fries, det. n.
sp. in herb, by Standley). West Indies; Central America; Colombia.
Abutilon hirtum (Lam.) Sweet, Hort. Brit. 1: 53. 1826. Sida
hirta Lam. Encycl. 1: 7. 1783.
Somewhat viscid, flowering as an herb persisting as a shrub, the
indument short-stellate and simple trichomes intermixed; petioles
to a dm. long; leaves suborbicular, sometimes obscurely 3-lobed,
cordate, sometimes obtuse, often several cm. long and nearly as
wide; flowers solitary, axillary but more or less corymbose above,
tawny-yellow or orange with darker base, the oblique subretuse
petals 1.5-2 cm. long; calyx 13-16 mm. long, the lobes acute or
acuminate; stamen tube stellate; carpels about 20, 3-seeded, early
densely stellate, 10-12 mm. long, the dark brown seeds minutely
pitted and stellulate, 2.5-3 mm. thick. — Tomentum of branches
generally ferrugineous, the long trichomes simple and somewhat
glandular; accessory axillary branchlets develop rapidly by the
flowers (Hochreutiner). Treated as a variant of A. indicum by
Grisebach, and apparently with reason.
Lima: Botanical Garden, Killip & Smith 21521 (det. Killip).
Tropical Asia and Africa; West Indies; Florida.
Abutilon indicum (L.) Sweet, Hort. Brit. 1: 54. 1826. Sida
indica L. Cent. PI. 2: 26. 1756; Amoen. Acad. 4: 324. 1759.
Herb or becoming suffrutescent the younger parts canescent-
tomentose with usually some longer trichomes intermixed; petioles
short or elongate; stipules 3-5 mm. long; leaves round-ovate to
broadly ovate, cordate, acute, more or less clearly 3-lobed, irregu-
larly crenate or serrate, often about a dm. long, finally glabrescent;
peduncles solitary or often corymbose, articulate below the calyx
this 1 cm. long with ovate acute lobes; petals nearly 1.5 cm. long,
yellow, oblique, pubescent at base; stamen tube glabrous; carpels
Flora of Peru 497
12 mm. long, coarsely tomentose the 3 seeds glabrous except at
hilum, pitted. — Indument canescent, short; leaves ovate, more
canescent below than above; long solitary axillary peduncles usually
longer than leaves; caljrx ordinarily shorter than the black hirsute
carpels these 15 or 16 (Hochreutiner). F.M. Neg. 23776 (var.).
Cuzco: Lucumayo Valley, Cook & GiWert ISH (det. Ulbrich).
Tropical Regions.
Abutilon lateritium Ulbr. Bot. Jahrb. 54: Beibl. 117: 56. 1916.
Young branches scabrous and hirsute-pilose; stipules lanceolate,
6 mm. long, tomentose as the suborbicular leaves below; petioles
S-5 cm. long, hirsute; leaf -blades deeply cordate, acute or acuminate,
5-9 cm. long, 4-8 cm. wide, irregularly and coarsely crenate-serrate,
glabrescent or subscabrous above, the 7 nerves prominent; flowers
subnutant, the erect axillary peduncles 5-7 cm. long, articulate
5 mm. below the calyx, this 2.5 cm. long, tomentose within and
without, the lanceolate lobes to 2 cm. long; corolla red, campanulate,
the obtuse petals soon reflexing, glabrous except at base, obtuse,
3.5-4 cm. long, connate 7-8 mm. with stamen tube, this conoid, to
3 cm. long, stellate only within toward base; styles many, to 1 cm.
connate, glabrous as capitate stigmas; fruit subglobose, to 2.5 cm.
across, the many compressed carpels 10-11 mm. high with beaks
2-3 mm. long, laterally glabrous, not connate, the 2 or 3 seeds
vemiculose, minutely pubescent. — Type a meter high shrub in rocks
comparable to A. reflexum with subentire leaves, smaller flowers
and fruit and different pubescence (Ulbrich).
Lima: Near Chosica, 2,000 meters, Weberbaiier 58^9, type.
Abutilon longipes Ulbr. Bot. Jahrb. 54: Beibl. 117. 52. 1916.
White velvety tomentose except leaves above, even to the
calyces; petioles subequaling the broadly ovate or suborbicular
deeply cordate leaves, these to 12 or 13 cm. long, acuminate, mi-
nutely dentate, tomentulose but greenish above, prominently about
7-nerved beneath; peduncles solitary, axillary, much exceeding the
leaves, articulate 1-2 cm. below the campanulate calyces, these to
3 cm. long, the lobes tomentose on both sides, 3-nerved, acute;
corolla campanulate, subspreading, glabrous, 3.5 cm. long, petals
yellow, nearly 2 cm. wide, claw 7 mm. long; stamen tube glabrous,
cylindrical; styles many, connate only at base, glabrous, the stigmas
subglobose; fruit subglobose, to 3 cm. thick, the tomentose carpels
498 Field Museum of Natural History — Botany, Vol. XIII
obtuse, about 18 mm. high with 3-5 finely verruculose seeds, pilose
around hilum, the funicle indurate. — Related to A. globiflorum Don
with smooth leaves and smaller flowers (Ulbrich) but seems to be
very near A. arbor eum, for which Kearney has kindly supplied the
key-difference; however the character "breaks" in some material
as that of Cook & Gilbert. Type 2 meters high, among other shrubs.
Ayacucho: Tambo Osno, Huanta, 2,500 meters, Weberbauer
5599, type. Ccarrapa, Killip & Smith 22305 (det. Killip). — Cuzco:
Marcapata Valley, near Chilechile, 2,200 meters, Weberbauer 7875
(det. Ulbrich). Ollantay tambo. Cook & Gilbert 27 U; 812 (det.
Ulbrich). "Phancho," "jarul-jarul."
Abutilon moUissimum (Cav.) Sweet, Hort. Brit. 1: 53. 1826.
Sida mollissima Cav. Diss. 2: 49. pi. IJ4.. 1786. A. calycinum Presl,
Rel. Haenk. 2: 116. 1835. A. sordidum Schum. in Mart. Fl. Bras.
12, pt. 3: 406. 1891. S. cistiflora L'H^r. Stirp. Nov. 127, pi 61. 1789.
Erect shrubs, the flowering branchlets abundantly pilose with
long (to 4 mm.) trichomes and some minute stellate ones intermixed;
petioles 5-10 cm. long, the promptly caducous subulate stipules
shorter; leaves ovate, sometimes sublobate, cordate at base, acumi-
nate, more or less densely stellate-tomentose on both sides and with
scattered simple trichomes above, crenate, often 8-15 cm. long,
6-12 cm. wide; inflorescence axillary, few-flowered (mostly 3-
flowered), peduncles 4-5 cm. long; bractlets simulating stipules;
calyx campanulate, plicate-angulate, the acuminate lobes tomen-
tulose within, tomentulose and hirsute without, 10-14 mm. long;
petals 15-18 mm. long, sulphur-yellow, glabrous, except the margin-
ally ciliate base; ovary subcylindric, costately angled, sparsely
pilose, the cells 5-ovulate; carpels 14-17 mm. long, 7-8 mm. broad,
yellowish-villous, finally dehiscent to base; seeds ovoid, obscurely
papillose, 2-2.5 mm. long. — Woytkowski 35133, a bush to 3.5 meters
tall, flower pale orange-yellow. Calyx turbinate at base; synonymy
after Baker. Type of A. calycinum, indument in part hispid-
spreading, by Haenke from "mountain valleys of Peru." F.M.
Negs. 7989; 29784.
Cajamarca: Below Guerocotillo, 1,600 meters, Prov. Cutervo,
Weberbauer 7120 (det. Ulbrich).— San Martin: Tarapoto, Woyt-
kowski 35133 (det. Cuatrecasas) . — Hudnuco(?): On the Marafion,
Dombey, type (also of S. cistiflora). — Junin: La Merced, Killip &
Smith 2356 Jt (det. Killip). — Lima: Yanga, Dombey.
Flora of Peru 499
Abutilon pauciflorum St. Hil. Fl. Bras. Mend. 1: 206. 1827.
Growing parts pubescent with long spreading trichomes and
tomentose with short stellate ones, the soft leaves paler and es-
pecially tomentose beneath; stipules narrowly subulate or filiform
to 1 cm. long; petioles soon elongate or nearly equaling the cordate-
ovate, acuminate leaves, these crenate-serrate, 9-nerved, the larger
about 12 cm. long; flowers solitary, axillary, on long stout peduncles,
the roseate obovate emarginate petals exceeding the calyx, densely
pubescent marginally at base; calyx cupulate, 12-15 mm. long in
flower, the lobes acuminate; carpels 8-10 (12), villous, about 15
mm. long, shortly beaked; seeds puncticulate, tuberculate-hispidu-
lous. — Dr. Hochreutiner has written me that he believes Baker f.
was correct in identifying Grisebach's interpretation of A, pedun-
ciUare HBK. with this species and wonders if the type of HBK.
should not include it, another problem beyond the scope of this
work. However, Kearney, Leafl. West. Bot. 7: 252. 1955, thinks
that the plant of North America and the West Indies may be
distinct. F.M. Negs. 19682; 35458.
Peru (probably). West Indies and Mexico to Paraguay.
Abutilon pedunculare HBK. Nov. Gen. & Sp. 5: 273. 1822.
A. pionense Ulbr. Notizbl. Bot. Gart. Berlin 9: 53. 1924.
Branchlets herbaceous, slender, stellate and pilose the later
trichomes spreading; stipules linear-lanceolate, to 1 cm. long, ca-
ducous; petioles finally subequaling the lower leaves; these all
deeply and narrowly cordate, subrotund, rather abruptly long-
acuminate, subentire or repand-denticulate, to 16 cm. long, 13 cm.
wide, the upper much smaller, puberulent or nearly glabrous above,
softly canescent tomentulose beneath; peduncles a dm. or two long,
slender, axillary, solitary, the pendulous blood-red flowers 3.5-4
cm. long; calyx tomentose, about 3 cm. long, cleft to below the
middle with lanceolate acuminate lobes; petals oblong-lanceolate,
finally reflexing, irregularly denticulate at tip, hirsute marginally
toward base, 6-8 mm. wide; stamen tube glabrous; caipels (as
styles) 15- about 20, acute, coriaceous, chartaceous, the 3-^ seeds
glabrous (HBK.), the fruit 1.5 cm. high, 2 cm. across at base, not
enclosed in calyx (Ulbrich). — Related to A. reflezum, with leaves
tomentose both sides and with smaller flowers and fruits, the latter
enclosed in caljrx (Ulbrich); differs also in the pilose branches,
longer peduncles, form of leaves and number of styles (HBK.).
A. pubistamineum Ulbr, Repert. Sp. Nov. 13: 500. 1915 has yellow
500 Field Museum of Natural History— Botany, Vol. XIII
flowers, lanate stamen tube. See also A. pauciflorum; length of
peduncles is very variable (B.P.G.H.). F.M. Neg. 35459.
Cajamarca: Among shrubs, trees, Ron, Prov. Cutervo, 1,300
meters, Weberbauer 714^0 (type, A. pionense). Chamaya to Tome-
penda, Prov. Ja^n, Bonpland, type.
Abutilon piurense Ulbr. Bot. Jahrb. 54: Beibl. 117: 55. 1916.
Branches weak, slender, tomentulose; stipules lanceolate, 4 mm.
long, promptly caducous; petioles 1-3 mm. long; leaves deeply
cordate-amplexicaul, ovate, long-acuminate, 2-8 cm. long, 1-6 cm.
wide, glabrous above, ashy green and rather sparsely appressed
stellate beneath, entire or obscurely dentate, the 5-7 palmate nerves
prominent; peduncles 4-8 cm. long, solitary, axillary, the roseate
flowers nutant; calyx 17 mm. long, the lanceolate acute lobes 9 mm.
long, sparsely tomentose without, glabrous within; corolla early
campanulate, finally reflexed, 2.5 cm. long, connate nearly 7 mm.
with stamen tube, this conoid, glabrous without, stellate within,
2 cm. long; petals oblanceolate, obtuse, glabrous except for some
stellate trichomes near base; ovary pilose, multicarpellate; styles
1 cm. long, glabrous as the capitate stigmas. — Compared by author
with A. megapotamicum St. Hil. & Naud. with stouter erect branches
and upright petals. Among other evergreen shrubs. F.M. Neg.
9284.
Piura: Chauro to Hacienda San Antonio, 800 meters, Weber-
bauer 6010, type.
Abutilon pulverulentum Ulbr. Bot. Jahrb. 54: Beibl. 117: 51.
1916.
Younger branches, petioles, these to 6 cm. long, and peduncles,
these to 3.5 cm. long, lustrous yellowish puberulent-tomentulose;
leaves broadly ovate, angulate to sub-trilobed, obtuse or slightly
cordate at base, cuspidate, to 12 cm. long, 6-7 cm. wide, irregularly
denticulate, sparsely stellulate above or somewhat densely on the
nerves, softly tomentose beneath, the 6-7 palmate nerves and
pinnate-reticulate venation there prominent; pedicels in anthesis
very short; flowers glomerate on terminal peduncles, soon forming
a lax efoliate irregular cyme; calyx yellowish tomentose, 6-8 mm.
long, the lobes acuminate; corolla rotate, lilac- tinted, the broadly
obovate obtuse petals about 2 cm. long; stamen tube glabrous,
4 mm. long; styles nearly 3 mm. connate, stigmas large, capitate;
fruit sparsely pilose, 9 mm. thick, carpels about 15, suborbicular.
Flora of Peru 601
apically subangled, glabrous, compressed, the black pyriform solitary
seed with an indurate funicle. — Allied to A. umbellatum Sweet with
pedicellate flowers, beaked fruit, more sparsely pubescent leaves
(Ulbrich) ; but Kearney has pointed out to me that the sketch on the
type specimen shows a solitary pendulous ovule and he therefore
considers it some species of Sida. However, this character varies
greatly in the genus, and in the family, for that matter, as in some
other families and its facies is that of this genus. Type 3 meters,
in wet places. F.M. Neg. 9286.
Cajamarca: San Miguel, 2,600 meters, Weberbauer 390U, type.
Abutilon ramiflorum St. Hil. Fl. Bras. Merid. 1: 199. 1827.
Shrubby at least below, the virgate branches, petioles (to 13
cm. long) and leaves on both sides densely sometimes rather loosely
tomentose; stipules stellate-tomentose, subulate-acuminate, tardily
caducous; leaves broadly ovate, shortly and usually acutely or
mucronately acuminate, 8-18 cm. long, 6-16 cm. wide, those at base
of the terminal ample panicles much smaller; flowers subtended by
stipuliform bracts, the peduncles at anthesis about as long or 5-10
mm. long, to twice as long in fruit; calj^ 3-4 mm. long, yellowish-
ferrugineous, the yellow petals to 9 mm. long, pilose at base as
stamen tube; carpels 6 or 7, chartaceous, stellate-tomentose, dorsally
dehiscent to the middle, 7-8 mm. long, the 3 seeds minutely puberu-
lent. — Simulates species of Wissadula and ovules in same position
but carpels constructed as in Abutilon (Schumann), to which state-
ment Hochreutiner replies: there is not the slightest resemblance
with any species of Wissadula! The Peruvian material seen, as
determined in herbaria, is apparently too young to show that in fact
the Peruvian specimen may not rather be a Wissadula species, as
W. stellata. F.M. Neg. 35461.
Junin: La Merced, KiUip & Smith 2JtOJ^ (so det. in various
herbaria). To Paraguay and Brazil.
Abutilon reflexum (Juss.) Sweet, Hort. Brit. ed. 1: 53. 1826.
Sida refleza Juss. in Cav. Diss. 1 : 36, pi. 7, fig. 7. 1785. Sida retrorsa
L'H^r. Stirp. Nov. 133, pi. 6U. 1789.
Slender stems, petioles, these 1-4.5 cm. long, and peduncles,
5-12 cm. long, minutely stellate-tomentulose or puberulent; stipules
subulate, acuminate, tomentose, 7-8 mm. long, more or less cadu-
cous; leaves ovate, basally cordate, shortly and acutely acuminate,
minutely or subrepand serrate, mostly about 1 dm. long, half as
502 Field Museum of Natural History— Botany, Vol. XIII
wide or the lower considerably larger, conspicuously soft-tomentose
beneath, green above with some scattered minute stellate trichomes;
flowers axillary, solitary, erect or nodding; calyx about 1.5 cm. long,
campanulate, truncate at base, the oblong triangular lobes erect or
reflexing with the petals, puberulent within; petals purple, narrowly
spatulate, dentate at apex, glabrous except marginally stellate-
pilose toward base where adnate with stamen- tube for nearly 1 cm.;
androecium glabrous; ovary densely villous, cells 3-ovulate, carpels
12 or 14; fruit unknown. — Leaves of the Asplund specimen obtuse,
not "acutely acuminate," and more coarsely serrate, thus simulating
A. lateritium, but may be separated by flower color and pubescence
(Fries). No type given for A. reflexum. Raimondi determinations
by Ulbrich, Goodspeed by Johnston. Flower glowing red (Good-
speed).
Piura: Talara, Haught 71. Parinas Valley, Haught 67 (det.
Ulbrich). — Cajamarca: Prov. Ja^n, Raimondi. Trujillo to Caja-
marca, Raimondi. — Lima: San Bartolom^ {Asplund 10869, det.
Fries). Cerros de Matucana, Raimondi; Goodspeed 11318. Eulalia
Valley, Goodspeed & Stork 11500. Rio Rimac, Goodspeed 30211;
33110 (det. Leonard); Safford. Prov. Hoara, Domhey (type, S.
retrorsa). Ecuador.
Abutilon striatum Dicks, in Lindl. Bot. Reg. 25, Misc. Not.
39. 1839.
Distinguished in Peru by the glabrous or glabrescent mostly
deeply lobed leaves and the showy more or less nodding deep yellow
flowers, 1-3 in the upper axils of the slender branches; peduncles
elongate; stamen column usually conspicuously exserted; petals
2.5-3.5 cm. long; carpels about 11, 7-9-ovulate. — Probably only in
cultivation in Peru or possibly established as an escape. According
to Kearney, A. pictum (Gill.) Walp. of Argentina is the same.
Illustrated, Paxt. Mag. Bot. 7: 53.
Cuzco: Yucay, Soukup 563. Uruguay; Argentina?
Abutilon sylvaticum (Cav.) Schum. in Mart. Fl. Bras. 12,
pt. 3: 418. 1891. Sida sylvatica Cav. Diss. 2: 56. 1786.
Branchlets rather stout, soon glabrate, the tips as petioles, more
or less tomentulose; leaves ovate-oblong, attenuate-acuminate,
cordate, more or less densely tomentose on both sides, lower leaves
(typically) deeply and narrowly cordate, the upper narrowly
triangular-cordate; internodes of the flowering branchlets sparsely
Flora of Peru 503
and minutely stellulate, otherwise glabrous (as to tjrpe); calyx to
17 or 18 mm. long, densely yellowish stellate, the lobes tjrpically
acutely acuminate, 10-12 mm. long; petals yellow, to 3.5 cm. long,
22 mm. wide, glabrous except sparsely and shortly pubescent with-
out; stamen tube multistriate (type), little ampliated at base; styles
as carpels about 11, the latter inflated, muticous, 1.5 cm. long,
ferrugineous-tomentose; seeds 2.5 mm. long, canescent-pilose. — In
part after Fries (subsp. genuinum R. E. Fr. Sv. Vet. Akad. Handl.
ser. 3. 24, no. 4: 7. 1947), who distinguishes two variants, subsp.
Btichtienii R. E. Fr. and subsp. Klugii R. E. Fr., which probably
more accurately could be designated varieties; both have densely
pubescent flowering intemodes, the indument of the second fer-
rugineous, both more acuminate leaves, more openly cordate, the
latter even ovate and more or less sagittate, the former with caudate
acuminate calyx-lobes, these for the latter narrowly deltoid, acute
and more ferrugineous; further, in the variant Klugii the stamen
tube is 5-sulcate, the styles and carpels 16. My specimens from
stream banks, to 8 meters high, the long graceful branches with
short floriferous branchlets. F.M. Neg. 29760.
San Martfn: Juanjuf, Klug J^39 (subsp. Klugii). Zepelacio,
Klu^ 37^9 (type, subsp. Klugii). — Hudnuco: Rio Azul {Asplund
125It2, subsp. Bucldienii). On the Rio Maranon, Dombey, type.
Near Muna, j^158. Mito, 1510. Cuchero, Poeppig 1255. — Aya-
cucho: Ccarrapa, KiUip & Smith 22UU2 (det. Killip). — Cuzco: San
Miguel, Cook & Gilbert 1176.— Vwao: Oconeque, MetcalJ 30587?
Bolivia. "Papagaru."
Abutilon umbellatum (L). Sweet, Hort. Brit. ed. 1: 53. 1826.
Sida umbellata L. Syst. Nat. ed. 10. 1145. 1759. A. Anderssonianum
Garcke in Anderss., K. Sv. Freg. Eugenics Resa, Bot. 230. 1855;
98, pi. 15. 1861, fide Svenson.
Stellate-tomentose or glabrate to the upper corymbose or sub-
umbellate calyces, the indument of the round to ovate leaves minute,
denser beneath, of the stems and petioles (1-4 cm. long) glandular-
subvillous or sparse; stipules linear, acute, 5-10 mm. long; leaves
sometimes somewhat 3-lobed, crenate-serrate, cordate or subtrun-
cate, more or less abruptly acuminate, 3-6 cm. long or longer; bracts
2-3 mm. long; calyx 5-6 mm. long, villous-tomentose, the acute
lobes half as long, slightly longer in fruit; petals yellow, 8 mm. long,
pubescent at base; carpels 5-7 (11), hirsute-tomentose, the awns
about 2 mm. long, each carpel with 3 brown tessellate-tuberculate
504 Field Museum of Natural History— Botany, Vol. XIII
seeds. — Svenson, I.e., has discussed the variation of the species in the
Galapagos. My collection has 11 carpels, to 9 mm. long. Illustrated,
Cav. Diss. 1: pi. 6; Svenson, Amer. Joum. Bot. 33: pi. 15, figs. 1-3,
opposite p. 465.
Piura: Chulucanas to Morropon, Weherhauer 5967 (det. Ulbrich).
— Hudnuco: Near Hudnuco, 3493 (det. Kearney). To Mexico;
Venezuela; West Indies.
Abutilon virgatum (Cav.) Sweet, Hort. Brit. 1: 53. 1826. Sida
virgata Cav. Icon. 1: 53, pi. 73. 1791. A. mendocinum'PhW. Sert.
Mend. Alt. 6. 1870, fide Kearney.
Perennial herb, more or less lignescent at the base; the many
erect strict stems stellate-tomentulose above as the petioles, these
1-3 cm. long, and the leaves on both sides; stipules 8-10 mm. long,
scarcely 1 mm. wide; leaves ovate, angled or more or less 3-lobed,
the middle lobe the largest, cordate at base, serrate or crenate, often
3.5-5 cm. long and nearly as wide; peduncles spreading, 1- or 2-
flowered, more or less congested above; calyx about 1 cm. long,
campanulate, the lanceolate lobes long-acuminate, puberulent within,
tomentose without; petals 7 or 8 mm. long, half as broad near the
tips, glabrous even at the base; stamen-tube hispid-stellulate; ovary
tomentose, the cells 3-ovulate; carpels complanate-trigonous,
bicomiculate, 7 mm. long, 3.5 mm. broad above, dorsally tomentose,
finally bivalved even to the base; seeds trigonous, slightly stellulate.
— F.M. Negs. 8000; 32633 (A. mendocinum) ; 35549.
Cajamarca: Prov. Chota, Raimondi. Prov. Contumazd, Rai-
mondi (both det. Ulbrich). — Hudnuco: Near Hudnuco, Soukup 2225.
— ^Ayacucho: Huanta, Raimondi (det. Ulbrich). To Chile; Argen-
tina; Brazil.
Abutilon Weberbaueri Ulbr. Bot. Jahrb. 54: Beibl. 117: 53.
1916.
Young branches scabrous- tomentose and viscid-hirsute; stipules
subulate, hirsute, to 7 mm. long; petioles subequaling the orbicular
leaves, these cordate at base, acuminate, 2-4 cm. long, coarsely
serrate, the 7 nerves inconspicuous; flowers solitary in the upper
leaf-axils or at the axils of short branchlets, the peduncles mostly
about 5 mm. long, articulate 1-2 mm. below the calyx, 
