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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.
¥
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. — ■
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, this cupulate,
7 mm. long, tomentose, within black-dotted, glabrescent, the ovate
acute or acuminate lobes 5 mm. long; petals yellow, to 14 mm. long,
12-13 mm. wide, obtuse, glabrous; stamen tube conical, 8 mm. long.
Flora of Peru 505
sparsely pubescent with stellate and simple trichomes; styles about
6 mm. long, glabrous as capitate stigmas; fruit with 6 tomentose
aristate carpels 8 mm. high, the solitary seeds cordate-ovoid, mi-
nutely appressed pubescent. — Allied to A. virgatum, which is less
pubescent, with long-petioled ovate-lanceolate leaves, long-pedi-
celled flowers (Ulbrich). Kearney, in herb., allies this to Bastardia
bivalviSf as the carpels are 1-ovulate, separating from the axis.
A meter tall shrub among cacti and other xerophjrtes (Weberbauer).
Cajamarca: Montaiia de Nanch6, Raimondi (det. Ulbrich). —
Ayacucho: Below Coracora, 2,500 meters, Weberbauer 5815, type.
14. MODIOLA Moench
Modiolastrum Schum. in Mart. Fl. Bras. 12, pt. 3: 276. 1891.
A creeping herb with coarsely crenate to palmately parted leaves
and an involucel of (2) 3 bractlets, technically like Sphaeralcea but
ovary cells many (14-20), carpels biaristate, more or less trans-
versely septate between the seeds or seed (Modiolastrum), this
reduced by Hochreutiner, Ann. Cons. Jard. Bot. Geneve 20: 124.
1917, maintained by Krapovickas, who found the chromosome
count 5 instead of 9 for the type (M. caroliniana).
Modiola caroliniana (L.) G. Don, Gen. Syst. 1: 466. 1831.
Malva caroliniana L. Sp. PI. 688. 1753. M. malvifolia Griseb. Sjonb.
Argent. 45. 1879, fide Hochreutiner.
Prostrate stems rooting at the nodes, the stems and leaves (or
the latter glabrate) pubescent with simple and stellate trichomes;
stipules caducous; leaves broadly ovate, the narrow lobes sub-
pinnately lobulate; peduncles solitary or binate, 2-^ cm. long;
calyx 6-7 mm. long, the slightly^ shorter bractlets persisting; petals
red, 7-8 mm. long; carpels about 4 mm. long, hirsute dorsally
toward the 1 mm. long beak, glabrous and tuberculate-rugose below,
the seeds glabrous. — Also to be expected, perhaps, is the variant
M. geranioides (Hook.) Walp., the plant as the carpels somewhat
tuberculate. Hochreutiner has written: surely only one species;
all others are varieties or forms. Curious that there seems to be no
record of this herb from Peru, so widely distributed as a weed in
warmer South America. Illustrated, Mart. Fl. Bras. 12, pt. 3.
pi. 80 (M. lateritia). F.M. Neg. 9337 (M. malvifolia).
Peru (probably). Tropical and subtropical America; South
Africa.
506 Field Museum of Natural History — Botany, Vol. XIII
15. SPHAERALGEA St. Hil.
Urocarpidium Ulbrich, Bot. Jahrb. 54: Beibl. 117: 63. 1916?
Like Malvastrum but the carpels in two sections usually separated
by a deep notch, the apical dehiscent and smooth, the lower in-
dehiscent and utriculate; the stigmas are capitate, the bractlets
commonly deciduous soon after anthesis, the species all perennial
herbs, at most suffrutescent (Kearney) or annual as interpreted
here. — Emphasizing the carpel differentiation as basic, the number
of ovules as secondary (as in Modiola), the genus includes mostly
pluriovulate but also many uniovulate species. The chromosome
number is 5 or 10 according to Webber, Cytologia 7: 313-323. 1936,
as cited by Krapovickas, Lilloa 17: 179-221. 1949, who has found
no species in Peru unless M. Sandemanii Sandw. which (in a letter
to Kearney) he has decided is a very aberrant species of Sphaeralcea
for which he would propose subgeneric rank. Inasmuch as this (at
this writing — August, 1952) has not been done, it seems inconvenient
to retain Sandwith's imperfectly known species in Malvastrum.
Kearney has noted S. Endlichii Ulbr. of Mexico as having almost
completely dehiscent carpels, only faintly reticulate at base while
the variation of ovule number and carpel-structure within the
closely allied Wissadula (and Pseudabutilon) is suggestive that when
all the evidence is in, so to speak, the taxonomy of the tribe may
have to be reorganized, as suggested by Jepson, Svenson, Sandwith
and others; probably there are more "aberrant" species yet un-
discovered. My feeling that the species of Ulbrich belongs here
has been supported by Hochreutiner as follows: may be kept distinct
if a genus can be characterized by one single character, here the
caudate carpels, which is rather variable; if not, may be united with
Sphaeralcea as an exceptional species with awns.
Sphaeralcea arequipensis (Jofinst.) Krapov. Bol. Soc. Argent.
Bot. 3: 71. 1950. Urocarpidium albiflorum Ulbr. Bot. Jahrb. 54:
Beibl. 117: 64. 1916 — not S. albiflorum Rose, 1890. M. arequipense
Johnst. Contr. Gray Herb. 70: 74. 1924. S. Weberbaurii Krapov.,
I.e., fide Krapovickas.
Sparsely and minutely stellate pubescent but becoming glabrous,
often somewhat branched, 1-several dm. tall; stipules membranous,
persisting; petioles 1-1.5 cm. long; leaves ovate-rhombic, obtuse or
subtruncate at the broadly cuneate base, subacute, often indistinctly
3-5-lobed, to 3.5 cm. long, 2 cm. or so wide, irregularly serrate;
peduncles axillary, to 1 dm. long, the white subsessile flowers
I
Flora of Peru 507
secundly congested toward the tips; subulate bractlets 3-5, caducous,
(2) 3-4 mm. long; calyx 4 (-5) mm. long, medially ovate-lobed;
petals glabrous, obovate, 4 mm. long; stamen tube 3 mm. long,
strongly dilated at base, connate with corolla and falling with it;
styles many, free for only 1 mm.; ovary sparsely pilose; carpels
about 15, much compressed, 2.5 mm. high, lower part only dorsally
reticulate, the single apical spine ciliate, nearly 8 mm. long, the
glabrous seed borne in the non-dehiscing reticulate part of the basal
cavity. — Petals white (Ulbrich) or light wistaria- violet (Pennell).
Syonymy after Krapovickas in letter to Kearney dated June 22,
1952. Ulbrich suggested in herb, that the imperfectly known M.
congestiflorum Johnst., Contr. Gray Herb. 70: 74. 1924, may be
shown to belong here, but for convenience it is retained in Mal-
vastrum. Illustrated, Krapovickas, I.e. 73 {S. Weberbaurii).
Lima: Rocky outcrops, 1,500 meters, Weberbauer 5326, type.
San Bartolom^, Weberbauer 5285. Ambar, 2,010 meters. Stork 11^6
(det. Johnston). Open rocky slopes, Tingo, Pennell 131 UU- —
Arequipa: Near Arequipa, 2,500 meters, Pennell 13168 (det. Ul-
brich). — Moquehua: Open mixed formation, Torata, Weberbauer
7U12 (type, S. Weberbaurii). Chile.
|| Sphaeralcea Sandemanii (Sandw.) Macbr., comb. nov. Mal-
vastrum Sandemanii Sandw. Kew Bull. 263. 1950.
Perennial (?), the slender flexuose stems to 3 dm. high, abundantly
foliose and floriferous only above with some purplish branchlets
from the upper axils, glabrescent below, minutely stellate-pubescent
toward the tip; stipules filiform, about 1.5 mm. long; petioles to
1 cm. long; leaves subrotund-ovate (hederiform), 7-12 mm. long,
5-10 mm. wide, about medially 3-lobed, the largest intermediate
lobe ovate, acuminate, the lateral much smaller, deltoid, greenish
above, canescent beneath, stellate- tomentulose as calyces both sides;
flowers solitary or at tip of the axillary branchlets; pedicels glabrous,
2-3 cm. long, bent above the articulation, this 4-7 mm. below apex;
bractlets none; calyx tube 3 mm. long, 7 mm. broad, the deltoid
ovate acuminate and acute lobes 3.2-3.5 mm. long and wide; petals
rose-lilac, about 1 cm. long, nearly as wide, subtruncate; stamen
column pilose at base only, the head of anthers 4 mm. thick; carpels
(young) as styles glabrous, with 1 ascending ovule, obscurely trans-
versely rugulose, not aristate. — Because of the single ascending
ovule this must at present be placed in Malva^strum but in habit
and facies suggests Sida or even Anoda and the doubtful validity
508 Field Museum of Natural History— Botany, Vol. XIII
of distinguishing Sida and Malvastrum on this ovular character
has been expressed by Svenson, Amer. Journ. Bot. 33: 466. 1946.
However, Krapovickas thinks this is an aberrant Sphaeralcea, and
Hochreutiner has written me that he believes it would be clearer
if mentioned under Sphaeralcea, not on account of the disposition
of the ovule, which is variable, but on account of the rugulose
carpels. Since, as indicated under Malvastrum, these plants may
actually constitute a single group, I venture to transfer this species
to the older name and avoid the typographical awkwardness of
merely mentioning it.
Arequipa: Chala, 500 meters, (Sandeman Ifi20, type, Kew).
16. MALVASTRUM Gray
Tarasa Phil. Anal. Mus. Nac. Chile, Bot. 10. 1891. Urocarpi-
dium Ulbr. amend. Krap. Darwin. 10: 606-636. 1954, part.
Various in habit and endurance, foliage and inflorescence as
defined. Involucel ordinarily present. Anthers apical on the
filament tube. Ovary cells 5-many, the solitary ovule erect or
ascending; stigmas apical, capitate. Carpels uniseriate, indehiscent,
only apically dehiscent or completely dehiscent, incurved-rostrate
or long-aristate to muticous. — The lateral carpel walls are firm,
persisting or thin, soon disintegrating; this is the most difficult
genus of Tribe Malveae to define satisfactorily, having the solitary
erect or ascending ovule of Subtribe Malvinae and capitate stigmas
of Subtribe Sidinae, but being clearly affiliated with Subtribe
Abutilinae through the uniovulate species of Sphaeralcea; it is very
.probably of polyphyletic origin (Kearney, I.e. 119). The carpels
may split apically but not enough to release the seed (Hochreutiner).
The genus Sphaeralcea St. Hil. with which Malvastrum has been
merged (Jepson, Man. Fl. Pis. Calif. 632. 1925) is now restricted to
the species with the carpels comprised of an empty upper portion
and a fertile indehiscent part; apparently only two species have
been found in Peru with this fruit-character; cf. also Nototriche for
species with completely dehiscent carpels splitting into 2 valves.
For discussion see Kearney, Leafl. West. Bot. 5: 23. 1947; 6: 51. 1951.
However, at least in floristic work where simplification of generic
keys is so desirable, there seems no sound reason for maintaining
Tarasa as distinct, for its basic character — carpels completely
dehiscent, valves finally separating — is a common character de-
veloped to extreme degree, and that is variable, pointing to both
Malvastrum and Sphaeralcea.
Flora of Peru 509
See also Kearney, Leafl. West. Bot. 6: 51. 1951, and also I.e.
238-251, 1955, where he defines the genus to include only perennial
often somewhat woody species of subtropical America having a per-
sistent triphyllous involucre, yellow corolla and nearly indehiscent
dorsally smooth, laterally smooth or somewhat rugose carpels;
in Peru this definition would apply particularly to M. americanum,
M. coromandelianum, and close allies or variants. So delimited, the
known chromosome number is 6-18, Skovsted, Joum. Genetics 31:
263. 1935, in contrast to 15 for the annuals, centering around M.
peruvianum recently transferred to Urocarpidium by Krapovickas,
I.e.; compare Sphaeralcea St. Hil.; in view of the great variation of
characters, the difference in opinions between qualified students,
and the existence of species that admittedly are not surely assignable
to any other proposed segregates, it seems more than probable that
at least in floristic treatments one generic name (Sphaeralcea) will
suffice and sectional designations would then indicate clearly
apparent sp)ecific relationship. For convenience when the valid
name has not been made in Malvastrum the generic name Urocar-
pidium is used.
For the acaulescent group the descriptions have been drawn
largely from the account by Hill, Joum. Linn. Soc. Bot. 39: 216-230.
1909. He notes that while this species-group simulates in habit
Nototriche the indument is comprised of 2-3-forked trichomes
except in M. Richii and M. Weherhaueri and instead of volcanic ash
and barren slopes they ordinarily are associated with other plants
in argyllaceous soils; then, of course, they usually have the in-
volucral bractlets of Malvastrum, these sometimes obsolete or
promptly caducous (Af. nuhigenum, M. oriasirum), another example
of the character-continuity within the family, beautiful if at times
annoying! Hill based his key on carpel characters but as these are
seldom available or discernible it is largely academic; the size of
fully opened flowers appears to be constant, but there seem to be
too many species accepted on basis of pubescence and leaf-lobation
and shape, making my key also only suggestive.
Material referred in American herbaria to Malvastrum has been
mostly on loan to Antonio Krapovickas during the preparation of
this compilation.
The species particularly that have no involucel or the bractlets
caducous are M. nuhigenum, M. oriastrum. M. Sandemanii, omitted
here, is under Sphaeralcea.
510 Field Museum of Natural History— Botany, Vol. XIII
Plants acaulescent.
Leaves entire or subcrenate but only near apex; carpels apically
hirsute.
Leaves ovate or orbicular, obtuse; calyx lobes acute.
M. parnassiaefolium.
Leaves somewhat obovate, acute; calyx lobes acuminate.
M. alismatifolium.
Leaves crenate to lobate; carpels glabrous or dorsally hirsute,
rarely only apically.
Peduncles much longer than petioles; bractlets ovate.
M. Stuebelii.
Peduncles shorter than petioles or little longer; bractlets linear,
rarely lanceolate-ovate or obsolete.
Corollas conspicuous, about 2-3.5 cm. long; carpels dorsally
hirsute or glabrous (species characters not proved).
Carpels rostrate (Hill), stellate-hirsute; leaves obscurely or
not pinnate-lobulate.
Leaves tomentulose-stellate, mostly 3-lobed.
M. Weberbaueri.
Leaves hirsute or glabrous above, irregularly serrate-
lobulate M. Bakerianum.
Carpels erostrate (Hill); leaves usually pinnate-incised or
lobulate.
Ovary as carpels pubescent; petals, if purplish, in age
toward apex.
Leaves (some) subpinnate-lobulate . .M. Englerianum.
Leaves less regularly or less deeply subpinnate-incised.
M. Bakerianum, M. acaule.
Ovary as carpels glabrous; petals bluish-white, violet
basally M. Hauthalii.
Corollas small, rarely more than 12 mm. long; carpels hirsute
dorsally in M. oriastrum, M. Richii.
Plants glabrous unless stipules, bractlets, calyces.
Leaves orbicular, crenate M. rhizanthum.
Leaves ovate, lobulate or incised-crenate.
Peduncles as petioles short; leaves lobulate.
M. oriastrum.
Peduncles as petioles elongate; leaves incised.
M. nubigenum.
Flora of Peru 511
Plants obviously pubescent, especially the petioles and the
leaves, at least beneath.
Leaves coarsely serrate or incised.
Carpels glabrous, verruculose; leaves ovate-oblong.
M. nuhigenum.
Carpels densely hirsute, smooth; leaves rotund,
subcordate M. Richii.
Leaves crenate, cordate-rotund.
Carpels dorsally hirsute, smooth; petals white; pe-
duncles 1-2 cm. long M. crenatum.
Carpels apically puberulent, verruculose; petals purple;
peduncles 2-5 cm. long M. hetonicaefolium.
Plants caulescent.
Inflorescence a terminal compact or, early, ovoid-capitate spike.
M. americanum.
Inflorescence axillary and terminal, flowers sometimes solitary.
Shrubs, the indument close, malpighiaceous-strigose or finely
stellate-scabrous or -tomentulose; leaves crenate-serrate
(lobulate-incised, M. operculatum, M. HornschiLchanum).
Indument somewhat strigose-malpighiaceous; carpels aris-
tate M. coromandelianum.
Indument all or mostly more than 4-rayed trichomes; carpels
muticous, cusped, minutely rostrate or dorsally aristate.
Petals yellow; carpel awns various, not elongate-pilose-
plumose.
Indument canescent-tomentose; petals 6-7 nmi. long;
carpels mucronulate M. depressum.
Indument stellate-scabrous, scarcely tomentulose unless
younger parts.
Petals 10-12 mm. long ex char.; carpels minutely
rostrate M. scoparioides.
Petals about half as long; carpels muticous or cusped,
dorsally tubercled to aristate (species scarcely
distinct) Af. scabrum, M. scoparium.
Petals roseate; awns elongate, pilose-plumose.
Leaves oblong-lanceolate, crenulate M. Mandoni.
Leaves ovate, sometimes narrowly but more or less
lobulate or incised.
512 Field Museum of Natural History— Botany, Vol. XIII
Petals 5-10 mm. long; stamens capitate.
M. operculatum.
Petals about 15 mm. long; stamens in cylindrical head.
M. Hornschuchanum.
Annual (or biennial) herbs, the indument sparse or mostly
coarse, the leaves usually more or less lobed or incised
(crenate, M. Urbanianum).
Leaves not simply crenate.
Indument, at least mostly, finely stellulate.
Cal3rx trichomes sessile or not black-stiped; lateral leaf
lobes oblong-linear M. Pennellii.
Calyx trichomes in type dark-stiped; leaf -lobes rhombic-
ovate M. tenellum, M. congestiflorum.
Indument, at least mostly, setulose or hirsute-stellate.
Carpels entirely reticulate and with a small interior
endoglossum.
Leaves palmately parted U. chilense.
Leaves 5-lobed U. Mathewsii.
Carpels laterally smooth centrally except M. Shepardae,
without endoglossum.
Upper edge of carpels gibbous; leaves 5-lobed, mostly
simply strigose M. Shepardae.
Upper edge of carpels horizontal; leaves 3-5-lobed,
glabrous or nearly above or indument stellate.
Carpels larger than 3 mm., protuberous; leaves
3-lobed, glabrate above U. macrocarpum.
Carpels to 2.5 mm. large, lightly rugulose; leaves
3-5-lobed, stellate pubescent both sides.
M. peruvianum.
Leaves simply crenate, suborbicular M. Urbanianum.
Malvastrum acaule (Dombey) Gray, Bot. U. S. Expl. Exped.
150. 1854. Malva acaulis Dombey ex Cav. Diss. 2: 82, pi. 35,
fig. 2. 1786.
Acaulescent, the petioles (3^ cm. long), rosulate leaves beneath
and peduncles, these usually 2-flowered and 2-4 cm. long, more or
less strigose-hirsute; stipules membranous, lanceolate, acute, ciliate;
leaves subrotund or broadly ovate, truncate or subcuneate at base,
2.5-3.2 cm. long and broad, glabrous above (or pubescent in type?).
;^
Flora op Peru 518
the 5-7 lobes unequally and acutely dentate, the serrations ciliate
apically; bractlets 2 or 3, oblong-linear, acute, ciliate, &-9 mm. long;
calyx 7-8 mm. long, the ovate-lanceolate acute lobes 5 mm. long,
sparsely setose at base without, stellate-tomentose within; petals
broadly obovate, 2-2.5 cm. long, basally stellate-tomentose; stamen
head cylindrical; carpels about 20, densely stellate-tomentose dor-
sally. — Root fleshy, napiform, edible (Dombey). Description (as
other older acaulescent species) after Hill, Joum. Linn. Soc. Bot.
39. 1909. Petals viridine-yellow (Pennell). The carpels may prove
to be rostrate or the character variable; in this case the species
probably will include the forms or variants M. Bakerianum, M.
Englerianum, to either of which my 905 could be referred; it has
lavender flowers, leaves simply hirsute on both sides, minutely
rostrate carpels. Inasmuch as the original character describes the
entire plant as white-stellate, corolla yellow, three times as long as
cal3rx, the interpretation of the species may be incorrect, but prob-
ably (and conceivably as to pubescence) the description was in
error, since the common form in the Lima mountains seems to be
the plant interpreted by Hill as belonging here. Herrera 171 from
Valle de Paucartambo, not seen, ought to be M. Bakerianum as to
range, if that species is distinct. Type by Dombey from Cordillera
of central Peru without exact locality; also by Maclean.
Lima: Huamantanga (Mathews 597). Open rocky slopes, east
of Canta, 3,700-4,100 meters, Pennell U663 (det. Johnston). Near
Yauli, 905.
Malvastrum alismatifolium Schum. & Hieron. Bot. Jahrb.
21:318. 1895.
Habit of M. parnassiaefoUum and similar except for the obovate
leaves, these (petioles to 3 cm. long, sparsely ciliate) entire, acumi-
nate, glabrous, to 2.5 cm. long, 10-13 mm. wide; peduncles usually
1-flowered, densely hirsute below the flower, to 9 cm. long; bractlets
8, linear, acute, &-8 mm. long, scarcely 1 mm. wide, glabrous; calyx
9-13 mm. long, the lobes 6-9 mm. long, 2.5-3 mm. wide at base,
marginally strigose, tomentulose within; petals obtuse, claw ciliate,
1.5-2 cm. long; carpels 10, glabrous, sparsely setulose at tip only.
— Perhaps a variant of M. -parnassiaefolium but the leaves quite
entire, acute, and sepals acuminate; there is sometimes a 2-flowered
peduncle and the reticulate-rugulose carpels are glabrous except
for a tuft of trichomes at tip. F.M. Neg. 9306.
Amazonas: Near Centamal and CumuUca between Pacasmayo
and Moyobamba (Stuebel 39, type).
514 Field Museum of Natural History— Botany, Vol. XIII
Malvastrum americanum (L.) Torr. Mex. Bound. Surv. Bot.
38. 1858. Malva americana L. Sp. PI. 687. 1753, fide Fawcett &
Rendle. Malvastrum spicatum (L.) Gray, Mem. Amer. Acad. n. ser.
4: 22. 1849. Malva spicata L. Syst. Nat. ed. 10. 1146. 1759.
Annual or sometimes perennial herb, more or less densely
puberulent-tomentose, usually scabrous-stellate or the upper sur-
faces of the ovate leaves and the calyces tuberculate-hispid; stipules
lanceolate-filiform, 4-6 mm. long; flowers capitate or approximate
in terminal spikes, sessile, the lowest with small foliose bracts, these
mostly 2-cleft, calyx 5 mm. long; petals yellow, 6-8 mm. long;
carpels subrostrate or produced in inner angle, hispid dorsally. —
Often a meter or so high; leaves irregularly serrate-crenate, very
rarely obscurely 3-lobulate.
Junin: Sandy roadside, La Merced, 5293 (det. Hochreutiner) .
Widely distributed in warm and tropical regions.
Malvastrum Bakerianum Hill, Journ. Linn. Soc. Bot. 39: 228.
1909.
Depressed, leaves rosulate; petioles strigose-hirsute, 1.5-5 cm.
long; stipules linear-lanceolate, 1-1.5 cm. long; leaves ovate, rounded
or subcordate at base, 2-3 cm. long, 2-2.5 cm. wide, glabrous or
sparsely hirsute above in type, hirsute on nerves beneath, crenate-
serrate or obscurely lobulate; peduncles 1-3 cm. long, strigose;
bractlets persisting, linear, 1-3, 8-9 mm. long, ciliate; calyx 1 cm.
long, medially lobed, the ovate acute lobes nearly or quite glabrous
without, pubescent within; corolla at first white (or yellow?), 20-23
mm. long, petals broadly obovate, 12 mm. wide, tube 1.5 mm. long;
carpels reniform, dorsally stellate, 4.5 mm. long, beak 1 mm. long;
seeds reniform, 1.5 mm. long and broad (Hill from Pennell 134.17
with yellowish petals, changing to lavender); sometimes the leaves
as the calyces without and the pedicels densely strigose-hirsute,
var. strigosum Hill, Kew Bull. 160. 1935; or the leaves densely
pubescent on both sides with long rigid erect trichomes, forma
hirsutissimum R. E. Fr., Sv. Vet. Akad. Handl. ser. 3, 24, no. 2:
12. 1947. Weberbauer 7338, nearly glabrous except strigose petioles,
had brimstone-yellow flowers but lacks the stellate indument and
3-lobed leaves with acute teeth of M. Weberbaueri, but so deter-
mined by Ulbrich; without carpels its determination is uncertain.
Cuzco: Urcos, 3,600-4,800 meters (Stafford S32, var.). Near
Cuzco (Stafford 325, type, var. strigosum). Hills of Saxaihuamdn,
Herrera 529 (distr. as M. parnassiaefolium). Prov. Paruro, Vargas
Flora of Peru 515
8^9. — Puno: Sicuani district (Stafford 822.). Grassy meadows and
slopes, puna, 3,850 meters, Pennell 18^17 (type, f. hirsutissimum).
Sta Rosa (Stafford S80, S81, SSI A, SSlB). Argyllaceous fields,
Tarapoto (Hill 7j^, type). Pueara, open meadows (Weberhaicer J^S,
det. Hill). Vilcanota (Pentland, det. Hill). — Moquequa: Saylapa,
3,600 meters, Weherhauer 7888 (det. Ulbrich, M. Weherhamri),
"Cuntur-cupa" (Herrera).
Malvastrum betonicaefolium Hill, Joum. Linn. Soc. Bot. 39:
221. 1909. M. Purdiaei Gray, var. huantense Baker, Joum. Bot. 29:
171. 1891.
Acaulescent, the rosulate crenate cordate-subrotund leaves
pilose beneath especially on the nerves, 1.5-2.5 cm. long, nearly as
wide, the membranous lanceolate acute stipules 1.5 cm. long, the
petioles 3-7 cm. long, more or less pilose, and the peduncles 2-5
cm. long, densely strigose-hirsute with few-rayed trichomes particu-
larly toward apex; bractlets 2, membranous, glabrous, linear-ovate-
lanceolate, acute; calyx 8 mm. long, the acute lobes sparsely pilose
without, densely tomentose within; petals purple, 13 mm. long, half
as wide, fimbriate; stamens in an elongate head; carpels verruculose,
slightly tomentulose apically.
Huancavelica: Near Huanta (Pearce, t3rpe). — Ayacucho: Putis,
Prov. Huanta, Weberbauer 7589 (det. Ulbrich, M. nubigenum).
Urocarpidium chilense (Braun et Bouch^) Krap. Darwin. 10:
619. 1954. Malva chilensis Braun et Bouch^ Ind. Sem. Hort. Berlin
1. 1857. Malva scorpioides Turcz. Bull. Soc. Nat. Mosc. 36, pt. 1:
562. 1863. Malvastrum peruvianum (L.) Gray, var. scorpioides (Turcz.)
Baker, Joum. Bot. 29. 168. 1891. M. Hinkleyorum Johnst., Contr.
Gray Herb. 70: 73. 1924, fide Krapovickas.
Erect, glabrate or sparsely villous and setose above; petioles
1-2.5 mm. long, stipules narrowly lanceolate, obliquely acuminate
to 13 mm. long, 2-4 mm. wide, glabrous or sparsely ciliate; leaves
palmately 3-5-foliolate, 3-4.5 cm. long or larger, the leaflets more
or less dentate or lobate, often glabrous except for a bristle ter-
minating each tooth; peduncles 2-6 cm. long, the sessile or sub-
sessile flowers early unilaterally congested; calyx about 4 mm. long,
more or less hirsute and within minutely stellate, somewhat ac-
crescent in fruit, the ovate acute lobes about 2.5 nmi. long; bractlets
3, filiform, equaling the calyx; corolla white with purplish base or
violet; staminal tube glabrous or nearly, 1.5 mm. long, anthers about
10; carpels 15 or so to 1 mm. high, laterally reticulate and with
516 Field Museum of Natural History— Botany, Vol. XIII
a minute endoglossum within. — Johnston separated his plant from
M. peruvianum on the basis of the smaller paler flowers, more slender
larger stipules and divided or very deeply lobed leaves and (I.e. 85:
68. 1929) wrote: M. scorpioides Turcz. seems to be a phase of it.
Illustrated, Krapovickas, I.e. 617 (carpel), fig. 2; pi. 3 (M. scorpi-
oides).
Arequ'ipa: Chachani Mountain, 2,100 meters, (Hinkley ItS, type,
M. Hinkleyorum) . Ataquita, Goodspeed Exped. 1563 Jt. San Lazaro,
{Cerrillo 38). Yara, (Vargas 8013). Chile.
Malvastrum congestiflorum Johnst., Contr. Gray Herb. 70:
74. 1924.
Herbaceous sub-simple annual 1-several dm. high, the flexuose
stems deciduously stellate-tomentose, densely lanuginose toward the
apex; stipules lanceolate, about 5 mm. long; petioles stellate, 5-20
mm. long; leaves rhombic-ovate, acute, coarsely sinuate-dentate,
entire toward cuneate base, rarely palmately 3-lobed, the obovate
obtuse lobes coarsely few-dentate, or apparently sometimes all of the
leaves 3-lobed, and the lobes obtusely incised-dentate, in type 2-5
cm. long, 1.5-3 cm. broad, or to twice as large; flowers in dense
sessile terminal glomerules (type) or the glomerules in age shortly
pedunculate; calyx about 5 mm. long, tomentose, the ovate acute
lobes 3.5 mm. long; bractlets 3, linear, 3-4 mm. long; corolla pale,
about 4 mm. long, the obovate rounded lobes 2.5-3 mm. long;
stamen- tube sparsely long- villous, as the filiform styles; ovary
depressed, globose, densely stellate-tomentose. — Apparently related
to M. tarapacanum (Phil.) Baker of Chile, but distinguished by its
larger green, rather than tomentose foliage, pale corollas, and non-
aristate fruit (Johnston); my collections have 3-lobed leaves but
dark-stiped calyx trichomes. M. multicaule, lit., is apparently
a nomen nudum, even transferred without description from Malva
to Malvastrum, based on Lechler 178Jt. from Puno, not seen by me.
Arequipa: Rocky ravines, south slope of Chachani Mountain,
2,745 meters {Hinckley 37, type). Pampa de Arrieros, Pennell 13328
(det. Johnston, M. tenellum). — Cuzco: Ollantaitambo, Pennell 13656
(det. Ulbrich, M. multicaule). — Lima: Garden edge, Viso, 755 (det.
Johnston). — Puno: Chuquibambilla, Pennell 13387 (det. Johnston,
M. tenellum?).
Malvastrum coromandelianuin (L.) Garcke, Bonplandia 5:
295. 1857. Malva coromandeliana L. Sp. PI. 687. 1753.
\
Flora op Peru 517
Appressed strigose with single and bifurcate trichomes including
the calyces on their many angles; stipules lanceolate, acuminate,
7-9 mm. long; petioles 1-2 cm. long; leaves ovate to oblong, usually
3-6 cm. long; flowers early solitary, later on subsidiary branchlets,
the peduncles 1-3 mm. long; calyx and bractlets subequal, the
former to 7 mm. long in fruit; petals yellow, 8-9 mm. long; carpels
with 1 long apical spine and 2 medial dorsal cusps or short spines.
—Illustrated, Fawcett & Rendle, Fl. Jam. 5: 105.
Hudnuco: Near Hudnuco, SovJcup 2225 A. Warmer and tropical
areas.
Malvastnim crenatum Hill, Joum. Linn. Soc. Bot. 39: 227.
1909.
Depressed, the leaves rosulate; petioles 3-5 cm. long, densely
strigose-tomentose as leaf-margins and nerves beneath; stipules
lanceolate, glabrous, 3^ mm. long; leaves cordate-orbicular, about
1.5 cm. long, 2 cm. wide, the younger especially above strigose;
5-7-lobulate-crenate; peduncles 1-2 cm. long, densely ciliate; bract-
lets three, 4 mm. long, glabrous or ciliate; calyx 5-7 mm. long,
glabrous except ciliate-margined; corolla white, 8-12 mm. long,
petals 1 cm. wide, emarginate, the hirsute tube 2 mm. long; stamens
in obconic head; carpels rounded, dorsally hirsute. — Resembles
M. rhizanthum in leaf-form and flower-size but differs in pubescence
and ciliate calyic lobes (Hill). My collection was distributed as
Af. Richii, with which species it is easily confused; the leaves are not
so definitely lobed as in Weberbauer's collection (Fries). F.M.
Neg. 4311.
Junin: Near Yauli, 4,400 meters, Weherbatier S60, type. Cerro
de Pasco, 4,200 meters, S058 (det. Fries). Without locality (Mac-
lean).
Malvastnim depressum (Benth.) Svenson, Amer. Joum. Bot.
33: 465. 1946. Sida depressa Benth. Bot. Voy. Sulphur 69. 1844.
Af. dimorphum T. Howell, Proc. Calif. Acad. Sci. ser. 4. 21: 331.
1935, fide Svenson.
Densely canescent- and somewhat viscid-tomentose shrub (or
early subherbaceous) becoming sometimes a meter or two tall, the
orange-yellow flowers to 1.5 cm. in diameter; petioles about 2(-5)
cm. long; leaves ovate, cordate or broadly cuneate at base, crenate,
mostly 2-3.5 cm. long, about as wide, or the lower to twice as large;
primary pedicels 1-2 cm. long (Howell) the later nearly obsolete.
518 Field Museum of Natural History— Botany, Vol. XIII
solitary or mostly several on axillary branchlets; bractlets 3, filiform,
5 mm. long; calyx 1.5-2 cm. across, the ovate subacuminate lobes
5-10 mm. long, 4 mm. wide in fruit; petals subentire, barbate at
base; carpels about 10, pubescent, to 4 mm. long, with 2 prominent
beaks at outer angle and an obscure erect cusp at the inner angle,
the obliquely reniform seed smooth (after Howell). — Compare the
similar — perhaps scarcely distinct — M. scabrum, M. scoparium.
Illustrated, Svenson, I.e., pi. 15, fig. 5, opposite p. 465; pi. 16, fig. 6,
opposite p. 469.
Piura: Disintegrated granite, Amotape Hills (Haught & Svenson
11521). La Brea, Prov. Paita, Weherbauer 7760. — Cajamarca:
Prov. Cutervo, Stork & Horton 10208. Galapagos; Ecuador.
Malvastrum Englerianum Ulbr. Bot. Jahrb. 42: 115. 1908.
Depressed, acaulescent; stipules linear-lanceolate, 1-1.5 cm.
long or longer, persistently ciliate; petioles 2-4 cm. long; leaves
rosulate, erect, broadly ovate, truncate or subcordate at base,
5-7-pinnatifid or incised, rarely subentire, glabrous above, ciliate,
beneath stellate and simply pilose especially on the prominent
nerves, 1.5-2.5 cm. long, 12-20 mm. wide; peduncles pilose, 12-15
mm. long; calyx about 12 mm. long, medially parted, the ovate-
deltoid lobes acuminate, strigose, the linear-lanceolate involucral
hirsute-strigose segments 10-12 mm. long; petals white, 2.5-3 cm.
long, 12-15 mm. wide, truncate or emarginate, ciliate at base where
connate for 3-4 mm.; stamen tube 8 mm. long, efilamentous for
about 5 mm.; ovary densely pilose. — The divided leaves distinguish
it at once from M. acaule (Ulbrich); however, some of the leaves
of the type are shallowly lobulate. Perhaps my 905, from Yauli,
Junin, belongs here, with minutely rostrate stellate-hirsute carpels,
pale lavender flowers but the leaves are simply hirsute on both
sides. F.M. Neg. 9312.
Cajamarca: Punos, Chacabamba to Aguanuco, Raimondi (det.
Ulbrich). — Ancash: Tallenga to Piscapacha, 3,800 meters, Weber-
bauer 2888, type.
Malvastrum Hauthalii Ulbr. Bot. Jahrb. 42: 114. 1908.
Acaulescent from stout (to 2.5 cm. thick) rhizomes, the rosulate
leaves multilobed; stipules membranous, ciliate, lanceolate, 5-6
mm. long; petioles 1-2 cm. long; leaves broadly ovate or suborbicu-
lar, 12-20 mm. long and nearly as wide, the 6-9 lobes irregularly
pinnatifid or divided, above sparsely, beneath glabrate or in type
Flora of Peru 519
densely stellulate and pilose, the trichomes mostly simple; pe-
duncles 1-2 cm. long; involucral segments 2 or 3, filiform or narrowly
lanceolate or subulate, sub-glabrous except pilose margins, yellowish,
5-6 mm. long; calyx 9-10 mm. long, lobes &-6 mm. long, glabrate
in age; petals 20-23 mm. long, violet in herb., ciliate at base and
scarcely connate 1 mm.; stamen tube glabrous, efilamentose 6 mm.;
ovary glabrous. — Related to M. Englerianum (Ulbrich), but differs
in fruit characters (Hill). The Weberbauer specimen has nearly
glabrous, more cordate-orbicular leaves, but the young carpels are
glabrous and minutely muriculate; the petals are described by
Weberbauer as bluish white with a violet spot on the lower part.
F.M. Neg. 9315.
Cuzco: Puno near glaciers of the Auzengate, 4,500 meters,
Weberbauer 777S (det. Ulbrich). Bolivia.
Malvastrum Hornschuchanum (Walp.) Macbr., comb. nov.
Sphaeralcea Hornschuchana (Walp.) Baker, Joum. Bot. 31: 363.
1893. Malva Hornschuchiana Walp. Nov. Act. Acad. Leop.-Carol.
19. Suppl. 1: 303. 1843. Malvastrum Rusbyi Britton, Bull. Torrey
Club 16: 64. 1889, fide Krapovickas.
Branching shrub, the type 2.5-3 dm. high, the young branchlets
and leaves densely stellate-pubescent; petioles slender, 2-4 cm. long;
leaves ovate-reniform, crenate, 3-lobed, the terminal lobe longest;
peduncles axillary, 1-8 cm. long; flowers purplish, many, capitately
congested, 2-3 cm. wide; bractlets beautifully stellate-tomentose.
— After Britton, the type from La Paz, who allied the species to
M. capitatum Griseb. Goett. Abh. 19: 90. 1874. Actually it seems
to be a form or state of M. operculatum.
Cuzco: Pillahuata, Prov. Paucartambo (Herrera SS29). — Arequi-
pa: Tiabaya, Pennell 13090 (det. Johnston, M. Rusbyi). Bolivia.
Urocarpidium macrocarpum Krap. Darwin. 10: 624. 1954.
A more or less stellate-pilose annual (apparently) with 3 lobed
and serrate-crenate leaves exceeded slightly by few-flowered solitary
axillary spikes of roseate or purple flowers, the petals about 7 mm.
long; petioles to 1.5 cm. long, stipules triangular, 3 mm. long, half
as wide; bractlets 2 or 3, lanceolate, pilose as the campanulate
accrescent calyx without, this at anthesis 7 mm. long, the teeth
4.5 mm. long, 2 mm. wide at base; petals obovate, obliquely trun-
cate, slightly pilose basally as the staminal tube, this 3 mm. long
with about 12 reniform anthers; stigmas capitate, apically papillose,
520 Field Museum of Natural History— Botany, Vol. XIII
about 10; carpels scabrous, indehiscent, 3 mm. high, 3.5 mm. thick,
with conspicuous protuberances and irregularly rugose; seeds reni-
form. — Well marked by the large flowers and fruits, these notably
roughened (author). Illustrated, I.e., jig. 3, d, e, f (calyx, flower)
and pi. 5 (type); I.e. 617, fig. 2, j (carpel).
Lima: Canta (Pennell 1^89, type). Matucana, 92. Viso, Good-
speed Exped. 115^2.
Malvastrum Mandoni (Baker f.) Macbr., comb. nov. Tarasa
Mandoni (Baker f.) Kearney, Leafl. West. Bot. 5: 190. 1949. Sphaer-
alcea Mandoni Baker f ., Joum. Bot. 31 : 364. 1893.
Stem ligneous, virgate, to several meters high; stipules lanceo-
late, acuminate; petioles 1.5-2.5 cm. long; leaves oblong-lanceolate,
obscurely cordate at base, acute, 7-12 cm. long, pubescent on both
sides, paler beneath; flowers peduncled in the axils or subsessile and
aggregate toward the tips of the stems; sepals lanceolate, acuminate,
canescent-tomentose; petals obovate, pink-purple, 15-18 mm. long;
carpels including the 2 awns pubescent, the solitary seeds reniform
(Baker). — Loose-branched slender trees, the flowering twigs "weep-
ing," to 7 meters tall (Balls) ; but apparently ordinarily a shrub
flowering when only a few dm. high. F.M. Neg. 23738.
HuancaveUca: Mejorada, Stork & Norton 10899. — Ayacucho:
Huanta, Weherbauer 7512 (det. Ulbrich). — Cuzco: Machupicchu
2,400 meters, Herrera 1962 (det. Ulbrich); Mezia 8080 A (det:
Killip); Balls B6807. Bolivia.
I
Urocarpidium Mathewsii (Turcz.) Krap. Darwin. 10: 616.
1954. Malva Mathewsii Turcz. Bol. Soc. Nat. Mosc. 36, pt. 1: 563.
1863.
Annual similar to U. chilense but the leaves at least typically
merely 5-lobed and dentate-crenate, the stipules only to 2 mm.
broad and, especially the carpels 2 mm. high. — Krapovickas, I.e.
620, has noted that Pennell 131Jt9 and Weherbauer 7^.11 appear to be
intermediate in character; possibly this suggests that this group of
annual forms will prove to concern one variable genetic entity.
Illustrated, Krapovickas, I.e. (nutlet) fig. 2, (chromosomes) fig. 1.;
pi. 2 (type).
Cajamarca: Contumazd, Raimondi 7136, and others. — Ancash:
Lomas de Mongon. Goodspeed Exped. 9179; Lomas de La Chay,
Goodspeed Exped. 9215. — Lima: Lurin, 594^^. Chorrillos, 5862.
Valle del Rimac, Grant 7^28. Chancay, Stork & Vargas 9366.
Flora of Peru 621
Lomas de Pativilca, Goodspeed Ezped. 9227. Lomas de Dona Maria,
Goodspeed Exped. 9256. Barranca, Worth & Morrison 9116. —
Arequipa: MoUendo, Vargas 8^8; Cook & Gilbert 52. Chile.
Malvastrum nubigena (Walp.) Bak. f. Journ. Bot. 29: 172.
1891. Sida nubigena Walp. Nov. Act. Acad. Leop.-Carol. 19. Suppl.
1: 307. 1843. Malva nubigena Wedd. Chlor. And. 2: 276. 1857.
M. parnassiaefolia (Gray) Wedd. var. lohulata Wedd. I.e., 275.
Nototriche incisa Turcz. Bull. Soc. Nat. Mosc. 36, pt. 1: 568. 1863.
Acaulescent, the caudex ligneous, the petioles (1-7 cm. long),
peduncles (1-9 cm. long) and leaves glabrous or more or less strigose-
hirsute especially the younger leaves beneath on the nerves; stipules
to 12 mm. long; leaves ovate-oblong, cuneate to cordate at base,
1-3 cm. long, 9-25 mm. wide, 5-7-lobed, or irregularly incised-
crenate or lobulate; bractlets early caducous, 1-3, linear or filiform,
glabrous or ciliate, 3-6 mm. long; calyx 5-8 mm. long, the ovate
acute lobes 3-5 mm. long, a little pilose only within; petals violet,
to 2 cm. long; stamen head obovoid, brush-like; carpels about 10,
glabrous, verruculose. — Variable in leaf-form (cf. Hill & Burtt,
Lilloa 4: 279, fig. 1. 1939) and var. bipinnatifidum R. E. Fr. Sv.
Vet. Akad. Handl. ser. 2. 24, no. 2: 11, pi. 2, fig. 10. 1947, from
adjacent Bolivia. Synonymy after Hill.
Krapovickas, Darwin. 10: 609. 1954, observed that this species
is aberrant not only in the verruculose carpels but in having 10
instead of 6 diploid chromosomes.
Ayacucho: Choimacota Valley, Prov. Huanta, 3,700 meters,
Weberbatier 7589 (det. Ulbrich). — Puno: Tissaloma, 4,570 meters
(Meyen). Bolivia.
Malvastrum operculatum (Cav.) Hochr. Ann. Cons. Jard.
Bot. Geneve 20: 129. 1917. Malva operculata Cav. Diss. 2: 65,
pi. 35, fig. 1. 1786. Tarasa plumosa (Presl) Kearney, Leafl. West.
Bot. 5: 190. 1949. Malva plumosa Presl, Rel. Haenk. 2: 124. 1835.
Malvastrum plumosum (Presl) Gray, Bot. U. S. Expl. Exped. 147.
1854.
Entire plant canescent stellate; stipules linear-subulate, small;
petioles to 3 cm. long; leaves narrowly or broadly ovate to truncate
or cordate-suborbicular, obscurely or acutely 3-lobate, the middle
lobe twice as long as the lateral, all unequally incised-dentate to
repand-crenate, more or less 4 cm. long and wide; bractlets 3, linear;
corolla purplish, twice as long as calyx; stamen tube early villous;
522 Field Museum of Natural History— Botany, Vol. XIII
ovary villous; carpels about 9, tomentose and with 2 plumose awns
6 mm. long or twice as long as the carpel body, operculate. — There
seems to be only one species but some Peruvian specimens have been
referred to T. Rahmeri Phil. Anal. Mus. Nac. Chile, Bot. 10: pi. 1.
1891, which according to Reiche appears to differ only by its tri-
angular-oblong acute dentate leaves truncate at base, and possibly
separable as a variety. The type of M. plumosa was by Haenke
from "Peru and Chile." F.M. Neg. 23736.
Lima: Obrajillo (Wilkes' Exped.). Rio Chillon, Pennell 1^465
(det. Johnston). Without locality (Mathews 776, fide Gray).
Hoara, Domhey, type. Matucana, 263. — Arequipa: Chala, Worth
& Morrison 15699 (det. Johnston, Palaua sp.?). Yura, Schmidt.
Rio Chile, Munz 1551Jf.. Vincocaya, Rose 189Jt.8. Pampa de Ar-
rieros, Rose 18971. Lower slopes Mt. Misti, Sandeman 3735. Near
Arequipa, Pennell 13161 (det. Johnston); and Tingo, 131^2. Chile.
"Tarasa."
Malvastrum oriastrum (Wedd.) Baker, Journ. Bot. 29: 172.
1891. Malva oriastrum Wedd. Chlor. And. 2: 277. 1857.
Acaulescent, the caudex ligneous, glabrous unless stipules,
bractlets and calyx within; stipules membranous, lanceolate, acute,
1 cm. long, sparsely ciliate; petioles dilated at base, 2.5-3.5 cm.
long; leaves ovate, subcuneate at base, acute, 2.5-3 cm. long, 1.5-2
cm. wide, lightly to deeply 5-7-pinnate-lobed, the lobes unevenly
crenate-dentate; peduncles 1-2 cm. long; bractlets often obsolete or
caducous, subulate, ciliate, about 5 mm. long; calyx 9-10 mm. long,
the lobes 6 mm. long, acute, glabrous without, velvety tomentose
within; petals white, 15 mm. long, 10 mm. wide; stamen head
cylindrical; carpels hirsute at tip. — The abortion of the involucel
in some cases cannot be considered in any way to exclude this
species, so nearly related to other Malvastrums (Hill). F.M.
Neg. 35515.
Ayacucho: Mountains of Ayacucho (Pearce). Bolivia.
Malvastrum parnassiaefolium (Hook.) Gray, Bot. U. S. Expl.
Exped. 150. 1854. Sida parnassiaefolia Hook. Icon. 4: pi. 385. 1841.
Malva parnassiaefolia (Hook.) Wedd. Chlor. And. 2: 275. 1857.
Acaulescent from a fusiform root, the rosulate basally cordate-
ovate leaves entire or slightly crenate toward the obtuse tips,
glabrous or sparsely pilose beneath on the nerves, usually about
1.5 cm. long and nearly as wide; stipules membranous, glabrous.
I
Flora of Peru 523
lanceolate, acute, about 1 cm. long; petioles 3-5 cm. long, sometimes
sparsely setose, as the solitary peduncles, these to 7 cm. long;
bractlets 1 or 2, linear, 5 mm. long, glabrous; calyx 9-11 mm. long,
the lanceolate acute lobes 6-7 mm. long, sparsely setulose without,
densely setulose-tomentose within; petals violet or white, 13-18
mm. long, obtuse, stellate-tomentose below as stamen tube, the
anthers elongate-capitate; carpels 1-12, a little verruculose dorsally
and stellate pubescent especially toward the tip. — Marked, with
Af. alismatifolium, to which the Weberbauer collection was once
referred, by the entire or subentire leaves.
Cajamarca: Coymaloche Pass above Hualgayoc, 4,000 meters,
Weberbauer S99S; 271. Ecuador.
Malvastrum Pennellii Ulbr. Notizbl. Bot. Gart. Berlin 11:
524. 1932.
Sparsely branched glabrescent annual, the sparsely stellate
puberulent stems 2 dm. or more long; stipules narrowly lanceolate,
si)arsely fimbriate, green, 3-5 mm. long, 1-1.5 mm. wide; leaves
at the ends of the branches pedately 3-lobed or sub-5-lobed (petioles
1.5-3 cm. long), rather thin, sparsely stellate on both sides, bright
green, about 5 cm. long, 5-6 cm. wide, the middle lobe 1-1.5 cm.
wide at base, dilated to 2-2.5 cm. wide toward the tip, the oblong-
linear lateral lobes 8-12 mm. wide, all somewhat dentate; flowers
solitary in the leaf-axils or crowded at branchlet tips, some sub-
sessile on peduncles 7-12 mm. long, purple, 7-8 mm. broad; calyx
patelliform, yellowish, green-margined, sparsely stellate without, the
ovate lobes 4-5 mm. long, about as wide at base, little exceeded
by the glabrous mostly closed corolla; fruit nearly enclosed in the
little enlarged calyx, the glabrous compressed reticulate carpels
3-4 mm. long, 2 mm. wide, 1 mm. thick, the single seed triangular-
ovoid, 2 mm. long. — Easily recognizable by the large bright green
leaves with very regular linear-oblong lobes; allied to M. jacens
Wats, of Mexico, long-petioled, thinner more pubescent leaves
(Ulbrich). The name commemorates meritoriously Francis W.
Pennell, news of whose death relatively early (in 1952) I received
with great regret. Krapovickas has questioned the generic position
of this plant in a letter to Kearney. The Pennell specimens from
open sandy places were referred by Johnston to M. teneUum (Cav.)
Hieron. Bol. Acad. Nac. C6rdova 4: 15. 1881, of Argentina, perhaps
correctly.
Arequipa: Rock ledges, 2,600 meters, near Arequipa, Pennell
1S206, type. Near Arequipa, Pennell 1S170?
524 Field Museum of Natural History— Botany, Vol. XIII
Malvastrum peruvianum (L.) Gray, Bot. U. S. Expl. Exped.
146. 1854. Malva peruviana L. Sp. PI. 688. 1753. Malva limensis
L. Amoen. Acad. 4: 325. 1756, fide Johnston. M. limense (L.) Ball,
Journ. Linn. Soc. Bot. 22: 32. 1885. M. peruvianum (L.) Gray, var.
limense (L.) Baker f., Journ. Bot. 29: 168. 1891. Malva echinata
Presl, Rel. Haenk. 122. 1835? M. costata Presl and M. Haenkeana
Presl, I.e. 123?
More or less stellate-pubescent annual, 1-several dm. tall,
branching from near the base, the indument hirsute especially on
the peduncles and calyces of the axillary secund racemes of lavender
or rarely white flowers; stipules ovate-lanceolate-subulate, to about
6 mm. long; petioles 1-2.5 cm. long; leaves ovate to subrotund,
more or less clearly 3 (5)-lobate (sometimes medially), irregularly
serrate, subtruncate or subcordate, usually about 3-5 cm. long and
wide, sometimes twice as large, often glabrate especially above;
racemes ordinarily in all the upper axils and 2-4 cm. long, the slender
peduncles about as long; pedicels in fruit to 3 mm. long; bractlets
3, filiform, subequaling the calyx, this in fruit nearly 5 mm. long, the
corolla sometimes white with purplish base, 3-4 mm. long, little
exserted; calyx lobes ovate, acute or apiculate; carpels about 12,
dorsally rugulose, nearly glabrous, 2-4 mm. long. — Usually in loose
or sandy soils or disturbed fields as a weed. Perhaps too broadly
drawn but there seems to be variation without stabilization or con-
comitance in degree of leaf incision, size and color of flowers, size
and shape of stipules; my 92 seemed to be var. scorpioides, leaves
deeply lobed, now U. macrocarpum Krap. In view of the many
species already questionable here, it seems strange others could be
added, so presumably they must have characters. The species of
Presl, all based on collections by Haenke from mountains or valleys
of Peru, unknown to me, were separated by the author as follows:
M. echinata, like M. peruviana but much branched, decumbent,
leaves 3- instead of 5-lobed, broadly cuneate instead of cordate at
base, middle lobe little instead of larger (than lateral), flowers
axillary, solitary or in a capitate raceme, bractlets 2; M. costata,
affine M. limensis and M. echinata but very distinct especially in
the carpels, these 8, reniform, obtuse at both ends, glabrous, trans-
versely costate and impressed puncticulate; M. Haenkeana, affine
M. limensis but most diverse in tomentum, flowers and carpels, the
erect branches, leaf -nerves beneath and spikes tomentose; leaves
3-5-lobed, cordate; bractlets 3, setaceous; corolla twice as long;
carpels acute at both ends, tomentose, biaristate. This, ex char.,
Flora of Peru 525
seems to be distinct and possibly is an earlier name for M. eongesti-
florum or M. Pennellii, Hochreutiner has suggested that M. costata
may prove to be a species of Wissadula. Some of the following may
belong to segregated forms. Illustrated, Krapovickas, I.e. 617.
fig. 2 (carpel); pi. 7 (type).
Cajamarca: Below San Miguel, 2,000 meters, Weberbauer 188. —
Lima: Prov. Huarochiri, Goodspeed Exped. 11605 (det. Krapovickas).
Chicla (Ball). Canta, Pennell 1^89; 14591. San Buenaventura,
Pennell 1^550. Rio Blanco, Killip & Smith 21678 (det. Krap-
ovickas). Near Lima and Obrajillo, Wilkes' Exped.; Weberbauer 1j^;
169.— Junin: Tarma, Killip & Smith 21782 (det. Krapovickas).—
Hudnuco: Acomayo and Pillao, 2,100 meters, Woytkowski SU176;
Slt265 (det. Cuatrecasas). Near Hudnuco, ^JtO. — Huancavelica:
Yauli, 3,500 meters, Stork & Horton 10862 (det. Standley).— Cuzco:
Ollantaytambo, Herrera S409. Experiment Est. Kairu, Vargas 666;
669 (det. Standley). — Arequipa: Mejia {Gunther & Buchtien, det.
Bruns). Posco {Gunther & Buchtien 200A; 210; 211, det. Bruns).
Cachendo (Gunther & Buchiien 281; 21 U, det. Bruns). — Puno;
Salcedo, ScmJkwp ^9. Chile. "Rupfu" (Vargas).
Malvastrum rhizanthum Gray, Bot. U. S. Expl. Exped. 148.
1854. Malva rhizantha (Gray) Wedd. Chlor. And. 2: 276. 1857.
Acaulescent from a fusiform tuberous root, completely glabrous
(except caljrx); petioles mostly 3.5-4 cm. long; leaves rotund or
obovate-orbicular, mostly subcordate, about 18 mm. in diameter,
crenate or doubly serrate, coarser serrations few, the obtuse teeth
early bristle-pointed, 3-7-plinerved; peduncles 1-flowered, con-
gested, much shorter than petioles even in fruit; bractlets 3, linear,
not even ciliate, shorter than calyx, this 4-6 mm. long, the ovate
rather obtuse lobes pilose only within, the corolla 6-10 mm. long;
stamens in a globose head; fruit depressed, the rather membranous
carpels rounded, but the dorsal angles with 3^ soft long teeth;
seed reniform. — After Gray; according to Mathews the carpels are
verruculose. My collections were from rocky short-grass slopes.
Junin: Above Bafios, Wilkes Exped., type. Cerro de Pasco,
4,200 meters, 3057 (det. Fries); (Mathews 911; S9-4).— Lima: Rio
Blanco, 4,500 meters, 3025 (det. Fries). Viso, about 3,000 meters,
600. Yauii, Weberbauer 221.
Malvastrum Richii Gray, Bot. U. S. Expl. Exped. 149. 1854.
Acaulescent, hirsute all over even to the base of the corolla, with
stellate trichomes, the crowded rotund subcordate leaves and pe-
526 Field Museum of Natural History— Botany, Vol. XIII
duncles on the crown of a thick napiform root; petioles to 2.5 cm.
long; leaves about 12 mm. across, coarsely doubly serrate or incised,
strongly 5-nerved; peduncles umbellately congested, 6 or 8 mm.
long, 1-flowered, sometimes with 1 or 2 bractlets like the 2 or 3
glabrous or ciliate linear-setaceous ones of the involucel, these half
as long as the hirsute calyx; corolla slightly longer than the calyx,
this 4-6 mm. long, nearly glabrous within. — Like M. rhizanthum
and M. acaule but smaller in all parts and pubescent (Gray).
Junin: Baiios, Wilkes' Exped., type. San Jos^, 4,000 meters,
11^8 (det. Fries). Cerro de Pasco (Asplund 11735, det. Fries).
Malvastrum scabrum (Cav.) Gray, Bot. U. S. Expl. Exped.
147. 1854. Malva scabra Cav. Diss. 5: 281, pi. 138, jig. 1. 1788.
Shrub sometimes about a meter high, greenish but closely and
finely stellate pubescent including the calyces; petioles 0.5-2 cm.
long; leaves ovate, rather coarsely serrate, rounded-truncate at base,
subacute, usually about 3 cm. long, 2 cm. wide or twice as large;
flowers clustered in all the upper axils, subsessile; calyx somewhat
accrescent in fruit, finally about 1 cm. across, the lanceolate-ovate
acuminate lobes about 5 mm. long; flowers 5-7 mm. long, yellow;
carpels dorsally tubercled, apically muticous (or nearly). — Differs
perhaps too slightly from M. scoparium (Gray); and probably the
fruiting characters will be found to be inconstant. Since writing
this my idea that this should be included in M. scoparium has been
confirmed by Hochreutiner but the characterizations may stand
since both names are still in general use. F.M. Neg. 29768.
Food plant of Vanessa Caryi (Smjrth).
Lima: Dombey, cultivated, type. Obrajillo, Wilkes' Exped.
Chosica, 531. — La Libertad: Chicama Valley, common in neglected
fields, Smyth U9 (det. Killip).
Malvastrum scoparioides Ulbr. Notizbl. Bot. Gart. Berlin 11:
525. 1932.
Erectly branched, scabrous-stellate-tomentose shrub; stipules
about 5 mm. long; petioles 1-1.5 cm. long; leaves rhomboid-ovate,
2-4 cm. long, 2-3 cm. wide, coarsely crenate-serrate, little ciner-
escent, prominently nerved beneath; flowers solitary or congregated
at the ends of the branches, subsessile; bractlets 5-7 mm. long;
calyx patelliform, early 5 mm., later 8-10 mm. in diameter, lobes
subacute, 5 mm. long, 3-4 mm. wide at base; petals yellow, oblong-
ovate, 10-12 mm. long; stamen tube 6-7 mm. long, fimbriate at
Flora of Peru 527
base; fruit 2-2.5 mm. high, 8-10 mm. across, the 10-12 carpels
stellate-tomentose with 2 minute lateral beaks, 1 apical nearly
1 mm. long. — T3T>e nearly 3 meters tall. Belongs in the neighbor-
hood of M. scoparium (L'H^r.) Gray differing in size of flowers,
fruits and leaf-indentation (Ulbrich). F.M. Neg. 9310.
Ancash: Near brook below Pampa Ramos, 2,000 meters, (Weber-
bauer 3196, type).
Malvastrum scoparium (L'H^rit.) Gray, Bot. U. S. Expl.
Exped. 147. 1854. Malva scoparia L'H^rit., Stirp. Nov. 53, pi. 27.
1785.
Like M. scabrum but as to type differs in the somewhat larger
merely crenate scarcely dentate leaves canescent at least below and,
especially, in the erect beak at the incurved apex of the carpels
these also with dorsal awns (Gray). — Schuman, however, Fl. Bras.
12, pt. 3: 270. 1891 found no fundamental difference and Hoch-
reutiner has written me that he believes there is none and that this
is the earlier name, the dates as given originally and by Pritzel
having been rectified by Otto Kuntze. F.M. Negs. 7990; 9324.
Hudnuco: Near Hudnuco, Dombey, type. "Escoba cimarona"
(Dombey).
Malvastrum Shepardae Johnst. Contr. Gray Herb. 70: 75.
1924. Urocarpidium Shepardae (Johnst.) Krap. Darwin. 10: 621.
1954.
Annual herb, the many erect or decumbent viscid-villous stellate-
setose stems 2-6 cm. high; stipules scarious, ciliate, 3-4 mm. long,
scarcely 1 mm. wide; petioles 1-3 cm. long; leaves suborbicular or
broadly ovate, truncate or obtuse at base, 1-2.5 cm. wide, 1-2 cm.
long, broadly 3-lobate or irregularly dentate, setose; flowers axillary,
solitary from lower leaves or flowers many (Krapovickas), the
pedicels 2-5 mm. long; calyx 5 mm. long, setose, the oblong-lanceo-
late acute lobes about 3.5 mm. long, to 6 mm. in fruit; bractlets 2,
borne 0.5-0.9 mm. below the calyx; corolla purple, 5 mm. long or
exceeding the sepals by about 1 mm.; stamen tube (stamens about
10) and styles glabrous as the ovate strongly rugose carpels, these
2.5 mm. long. — Apparently allied to M. pygmaeum (Remy) Gray,
but differing ex char, in its acute calyx-lobes, pedicellate flowers,
glabrous fruit; perhaps comparable also to M. moUendoense Ulbr.
Named for Mrs. R. S. Shepard, who made a collection in the vicinity
of Lake Titicaca and Tacna during her missionary work about 1919.
528 Field Museum of Natural History— Botany, Vol. XIII
Krapovickas I.e. noted an endoglossum in carpels and chromosome
count of 20 as in the similar U. chilense. Illustrated, Krapovickas
I.e. 617, fig. 2 (carpel); pi. ^ (type).
Cajamarca: Chota, Raimondii 631 J^. — Hudnuco: near Huanuco,
20J^0. — Lima: Matucana, 93. — Cuzco: Ollantaytambo {Cook & Gil-
bert 644)' Urubamba, Vargas 7887. — Puno: in meadows Shepard
123, type). Bolivia; Argentina.
Malvastrum Stuebelii Hieron. Bot. Jahrb. 21: 318. 1895.
Herbaceous from fusiform root; petioles to 2.5 cm. long; leaves
ovate, acutish both ends, irregularly crenate-serrate, hirsute be-
neath especially on the pinnate nerves, otherwise glabrate, to 3.5
cm. long, 2 cm. wide; peduncles 1-flowered, hirsute, to 2 cm. long;
bractlets 3, ovate, acuminate, hirsute, 6-7 mm. long, 2.5-3 mm.
wide; calyx 10-14 mm. long, the hirsute lobes tomentose above;
petal claw ciliate, 2.5-3 cm. long, 17 mm. wide; carpels about 18,
densely hirsute. — Leaves almost if not quite glabrous above, crenate,
calyx densely strigose-tomentose, fruits conspicuously different from
M. parnassiaefolium (Hill). F.M. Neg. 9325.
Cajamarca: Monte Altura de Santa Ursula (Stuebel 38, type).
Malvastrum tenellum (Cav.) Hieron. Bol. Acad. Nac. Cordova
4: 15. 1881. Malva tenella Cav. Icon. PI. 5: 14. pi. U22, fig. 3. 1799.
A small annual (to 6 dm. high) with slender weak branching
pubescent stems and small subsessile flowers in clusters of 3 in the
upper axils; leaves 3-lobed, to 12 mm. broad and long, the petioles
somewhat longer; bractlets linear; petals light blue, slightly larger
than the calyx; carpels 9; seeds reniform. — Unknown to me unless
two or three specimens so determined by Johnston and in this work
referred to M. congestifiorum actually are referable to the species of
Cavanilles; it seems possible that only one species is concerned and
doubtful that the flowers are always only 3 in an axil.
Peru (see note above). Chile.
Malvastrum Urbanianum Ulbr. Bot. Jahrb. 42: 119. 1908.
Annual, the many erect or ascending stems becoming a dm. or
two long, sparsely stellate pubescent as the persisting subulate-
lanceolate stipules (2-3 cm. long); petioles 2-5 (-20) mm. long;
leaves rhomboid-semiorbicular, truncate or broadly cuneate at base,
rounded at apex, stellate on both sides and densely pilose beneath,
the nerves there prominent, the margins crenate; flowers roseate
Flora of Peru 529
white, the inflorescence subcapitate in the axils of the upper leaves;
calyx campanulate-tubular, nearly equaling the corolla, about 5
mm. long, the lanceolate-deltoid lobes dilated after anthesis, densely
villous especially toward the base; petals oblanceolate, subobtuse,
the narrowed base and margins pilose and stellate; stamen tube to
3 mm. long; ovary densely stellate; carpels 2.5 mm. long, 1.5 mm.
wide, dorsally pilose; seeds glabrous, foveolate-puncticulate, 1 by
1.5 mm. — Near M. tenellum (Cav.) Hieron. but distinguished by
the unparted leaves that suggest those of an Alchemilla (Ulbrich).
Dr. Kearney has kindly informed me of Krapovickas' intended
transfer of this species to Tarasa.
Puno: Above Cuyocuyo, 3,600 meters, Weherhauer 927, type.
Limbani, Sandia, Vargas IS 12 (det. Kearney).
Malvastrum Weberbaueri Ulbr. Bot. Jahrb. 42: 115. 1908.
Habit of M. Englerianum but leaves both sides densely appressed
tomentulose with minute stellulate trichomes, the lobes irregularly
incised-serrate, the teeth subapiculate, 2-3 cm. long, 1-3 cm. wide;
flowers many, on peduncles 1 cm. long; calyx 9-10 mm. long, the
indument minutely stellulate and simple trichomes intermixed, the
involucral segments 3, persisting, brownish, two lanceolate with
subulate tip, the third about 5 mm. long, subulate-filiform; petals
sulphur yellow, 18-20 mm. long, 10-11 mm. wide, connate about
2 mm., pilose below as the stamen tube, this 8 mm. long, antheri-
ferous for 4 mm.; fruit broadly conoid, 7 mm. broad, 5 mm. high,
densely tomentose with simple and stellate trichomes; seeds reddish,
glabrous, narrowly reniform. — Otherwise like the related species; the
overall indument is velvety but there are coarse strigose trichomes
on petioles, peduncles, calyces and often on the leaf-nerves beneath,
and the carpels are rostrate, the beak 1 mm. long (Hill). Cf. M.
Bakerianum for Weberbauer 7SS8 without fruit referred here by
Ulbrich. F.M. Neg. 9328.
Ancash : Above Acros, 3,500 meters, Weherhaiier 2760, type.
17. NOTOTRICHE Turcz.
Reference: A. W. Hill, Trans. Linn. Soc. ser. 2. Bot. 7: 201-266.
1909.
Characteristically little, or small depressed caespitose or pul-
vinate shrubs with ligneous subterranean caudex, rarely annuals, the
1-flowered peduncles more or less adnate as the stipules (these
rarely wanting) to the petioles. Leaves various, often pinnately or
530 Field Museum of Natural History— Botany, Vol. XIII
palmately lobed (rarely entire), incised or dissected, mostly basal,
the fusion of the stipules and the petioles forming a sort of vagina.
Involucel usually wanting. Calyx usually medially 5-lobed, often
stellate-tomentose, and with 5 glanduliferous nectaries at base.
Petals 5, obovate or oblong, more or less united. Ovary 5- to many-
celled, each cell with 1 ascending ovule. Styles capitate and stig-
matose apically. Fruits depressed, the numerous carpels 1-seeded,
cusped or beaked, at maturity splitting nearly to base (rarely
adhering to seed, Krapovickas), the seeds reniform, compressed. —
After Hill in the species descriptions at least in large part. Krapo-
vickas, Bol. Soc. Argent. Bot. 4: 107 et al has given basic chromo-
some number 5 and noted involucels in some annuals.
The name of Hill is so closely associated with this group that
his initials have not been used in the following compilation of his
studies. In Peru at least only acaulescent species of Malvastrum,
nearly always with an involucel, are likely to be confused with any
of these little alpine mallows.
Hill's introduction. I.e. 201-214, is interesting and informative;
the following key is based primarily on his, which however included
in 1909 only 62 species for the entire genus, fewer than known today
from Peru only; there are undetermined and apparently undescribed
species in herbaria, which it has been impractical for me to include.
Collections are still too limited for consideration of probable vari-
ability especially as regards characters of pubescence and probably
in some cases of leaf-dissection. The key therefore is only suggestive,
characters used for expediency or of necessity not being in all cases
my expression of their validity; as indicated by the monographer,
the main sections of the key, with present knowledge, merge. Hill's
listing of the species in Bot. Jahrb. 37: 559. 1906, with reference to
Baker, Syn. Malv. 40-43 for the original publication of each under
Malvastrum, has been accepted by me as valid transfer but in any
case, as usual in this work, the page reference to Hill's monograph
is cited.
Two names have been omitted: Sida acaulis Cav. Icon. 5: 13.
pi. U22. jig. 2, not Malvastrum acaule (Dombey) Gray, not identified
by Hill; M. Cavanillesii Gray, Bot. U. S. Expl. Exped. 156. 1854,
mentioned by Hill, I.e. 260 and from the short diagnosis essentially
a nomen nudum.
Annuals; flowers 2-3 (4) mm. long.
Leaves about orbicular, never deeply lobed; carpels minutely
birostrate.
Flora of Peru 531
Leaves crenate-dentate, loosely tomentose; carpels dehiscent.
N. piLsilla.
Leaves medially 3-lobed, stellate; carpels adhering to seed.
N. sarmentosa.
Leaves flabellate-reniform or suborbicular, then lobed to base;
carpels erostrate.
Leaves broadly 3-parted; carpels dehiscent N. nana.
Leaves 8-10-incised or -lobed N. pygmaea.
Perennials, the ligneous caudex often branched; flowers often larger.
Anthers globosely capitate; leaves usually subrotund, palmately
or flabellately parted the segments often similarly lobed or
incised, less frequently oblong to ovate and pinnate to bi-
pinnate; petals nearly always more or less coalescent at base
and rarely if ever longer than 2 cm. (see also N. lanata,
N. argylloides, anthers unknown). — See page 535.
Leaves more or less distinctly longer than wide, about oblong
or ovate, sometimes broadly, rarely broadly obovate or
subrotund but tending to pinnate or bipinnate or if
palmately or flabellately divided the segments (at least
shortly) pinnate or bipinnate (more or less clearly).
Leaves glabrous or glabrate at least beneath in age (see also
N. rugosa, N. flabelleta).
Leaf indument nearly or quite absent; petals 5-6 mm. long,
tube 1.5-2 mm. long N. salina.
Leaf indument rather tomentose; flowers (8) 10-20 mm.
long or longer, tube (3) 4-8 mm. long.
Calyx densely if finely pubescent; vagina ciliate to
sericeous.
Leaves diverse, fertile pinnate, sterile bipinnate.
N. longituha.
Leaves similar; corolla tube rarely 6 mm. long.
N. Mandoniana.
Calyx glabrous or nearly unless the lobes (several of the
following are similar).
Leaves suboblong or about oblong-lanceolate, rarely
linear.
Flowers 10-15 mm. long; carpels long-sericeous.
Leaves linear, 1-3-lobed; vagina ciliate.
N. armeriifolia.
532 Field Museum of Natural History— Botany, Vol. XIII
Leaves suboblong, subpinnate; vagina glabrous
or nearly N. longissima.
Flowers about 2 cm. long; carpels short-stellate
below; vagina ciliate N. longirostris.
Leaves ovate or about ovate; flowers about 2 cm. long
or longer; carpel trichomes all long.
Leaves somewhat bipinnate; vagina rarely glabrous;
petals about 2 cm. long, not red.
Vagina as carpels densely stellate-pilose.
N. anthemidifolia.
Vagina glabrous or ciliate.
Stipules eciliate or nearly; vagina glabrous;
carpels as leaf margins setulose.
N. aristata.
Stipules densely ciliate as vagina; carpels also
or these sericeous.
Leaf segments many N. sericea.
Leaf segments few N. acuminata.
Leaves obscurely or not bipinnate; vagina glabrous;
petals red, soon 3-4 cm. long.
Leaves palmately parted; calyx tomentose within.
N. purpurascens.
Leaves pinnatifid; calyx glabrous within.
N. Vargasii.
Leaves usually densely pubescent at least beneath.
Corolla crimson, 2-2.5 cm. long, tube obsolete; leaves
broadly obovate, often glabrate N. flahellata.
Corolla purple; leaves pubescent both sides.
N. porphyrantha.
Corolla not red, less than 2 cm. long; tube 3-5 mm. long;
leaves not obovate.
Leaves ovate-lanceolate, bipinnate; petals violet, 14
mm. long N. erinacea.
Leaves subrotund.
Indument tomentose; petals about 18 mm. long.
Indument lax; lobules subovate N. nigrescens.
Indument close; lobules suboblong . iV. artemisioides.
Indument minute; petals 7-8 mm. long. . . .N. rugosa.
Flora of Peru 533
Leaves suborbicular or reniform, rarely broadly obovate, pal-
mately or flabellately parted but the segments similarly
lobulate (more or less clearly) rarely unevenly; some
species, as observed by Hill, seemingly connect this with
some of the species of the previous group.
Leaves medially or submedially lobed, often broadly, lobes
few to (rarely) many, frequently with one to several
lateral or basal lobules, rarely many-lobulate apically.
Flowers about 1.5-2.5 cm. long.
Petals scarlet or red.
Leaves tomentose N. lanata, N. flabellata.
Leaves glabrous N. cupuliforme.
Petals never red.
Leaves glabrous or nearly on one side, unless the
lobules.
Leaf-lobes 3-5, medial, somewhat laterally lobulate;
indument sparse below.
N. sajamensis, N. condensata.
Leaf-lobes as lobules often many, latter narrow or
basally broad; indument often lacking below.
Lobes broad, laterally lobulate A^. ohcuneata.
Lobes not or little lobulate laterally, rather,
apically.
Lobules many; stipules narrow; indument soft.
N. ticsanica, N. sajamensis.
Lobules few; stipules oblong; indument in-
durate in age N. aretioides.
Leaves densely pubescent unless in age both sides.
Lobules dissimilar, partly lateral; indument short
or lax.
Primary lobes many N. famatinensis.
Primary lobes 3-5.
Indument minute, close N. obtusa.
Indument coarse, lax N. nigrescens.
Lobules similar, terminal; indument long, soft.
N. boriissica.
Flowers usually less than 10 mm. long.
Leaves densely pubescent both sides.
534 Field Museum of Natural History— Botany, Vol. XIII
Calyx about 8 mm. long; leaves often 6-9 mm. wide.
Stipules not developed N. estipidata.
Stipules well developed.
N. candaravica, N. holoserica.
Calyx 4-5 mm. long; leaves about 4 mm. wide.
Leaves suborbicular; stipules ovate .... AT. Azorella.
Leaves rhombic-obovate; stipules filiform.
N. congesta.
Leaves glabrous or glabrate beneath.
Flowers never red.
Leaf -lobes broad, with many minute lateral lobulae.
N. ulophylla.
Leaf-lobes about oblong, with few if any lateral
lobulae.
Lobes terminally multilobulate; flowers yellowish.
N. sajamensis.
Lobes few-lobulate or blades deeply 6-9-crenate;
flowers white or lilac.
Stipules glabrous or nearly; plants glutinous.
N. foetida.
Stipules stellate at least marginally.
Leaf lobules 5-9; vagina trichomes hirsute-
stellate N. ohcuneata, N. glacialis.
Leaf lobules 3-4; vagina trichomes subspinose
in age N. aretioides.
Flowers scarlet N. coccinea.
Leaves or (and) lobes basally or subbasally (often multi-)
parted (except N. Staff or dae), sometimes simply or the
3-9 lobes deeply bifid, trifid or more parted, pubescent
or rarely less so beneath (except in N. pedatiloba, gla-
brous beneath). N. sajamensis, N. horrusica, N. holo-
serica, N. nigrescens might be sought here.
Calyx lobes entire; primary leaf lobes 3-7.
Stipules broadly oblong, obtuse; lobules often very many.
Lobules numerous.
Leaves glabrous beneath; stipules broadly truncate,
glabrous A^. pedatiloba.
Leaves pubescent beneath as the obtuse stipules.
N. turritella.
Flora of Peru 585
Lobules about 12 N. Meyeni.
Stipules filiform to linear-lanceolate or linear-oblong.
Primary leaf -lobes 3, elongate, deeply 3-5-parted.
Corolla purple, 10-12 mm. long; tube short.
N. Orbignyana.
Corolla roseate, 17 mm. long; tube 6 mm. long.
A^. dissecta.
Primary leaf-lobes digituliform or 5-7.
Flowers 1 cm. long, white N. peUicea.
Flowers about 1.5 cm. long.
Lobules digituliform, elongate. . . .N. digitulifolia.
Lobules inflexed, short A^. alternata.
Calyx trifid; primary lobes 9, 5-lobulate N. Staff ordae.
Anthers disposed cylindrically in an oblong head, rarely slightly
long-obovoid but longer than thick; leaves in general ovate
or obovate, rarely linear, entire or subentire, rarely to sub-
rotund but the 3-7 segments tending to pinnate lobulation;
petals free or nearly except N. argentea, commonly longer
than 2 cm. See also N. flabellata.
Indument lacking or much less dense on leaves beneath.
Petals red, narrow; leaf indument velvety.
Leaf segments few, acute N, stenopetala.
Leaf segments many, obtuse A^. epileuca.
Petals never red, obovate; leaf indument rarely velvety.
Leaves linear, entire or shortly 1-2-lobed or fertile; simply
pinnate with few short lobes N. gracilens.
Leaves all much cleft or lobed, all wider than linear.
Flowers 3-3.5 cm. long at least typically.
Stipules tomentose; leaves lanate above. A^. Macleanii.
Stipules glabrous; leaves velvety above,
N. pichinchensis.
Flowers to 2 (2.5) cm. long.
Leaves longer than wide.
Indument tomentose; calyx pubescent.
Calyx lanate; leaves ovate-oblong, palmately
to subbipinnately parted A^. Matthewsii.
Cal3rx sparsely stellate; leaves oblong-lanceolate,
bipinnate N. Pearcei.
536 Field Museum of Natural History— Botany, Vol. XIII
Indument pulverulent; calyx tube glabrous.
N. argylloides.
Leaves suborbicular N. phyllanthos.
Indument dense, about equal on both surfaces.
Flowers about 2.5 cm. long.
Leaf segments multilobulate N. Herrerae, N. sulphurea.
Leaf segments 3-5- or few-parted.
Indument very short-tomentulose A^. Castelnaeana.
Indument long-sericeous N. argentea.
Flowers about 15-18 mm. long.
Indument tomentose.
Leaves trifid, lobes trifid and lobulate . . N. artemisioides.
Leaves 3-7-parted, lobes crisply multilobulate.
A^. pedicularifolia.
Indument lanate N. lanata.
Nototriche acuminata Hill, Trans. Linn. Soc. ser. 2. Bot. 7:
258. 1909.
Depressed, caespitose, the ashy sericeous rosulate leaves with
petioles 2-3 cm. long, the stipules adnate to below the middle and
with the membranous vagina 4-5 mm. wide, the free part linear-
lanceolate, acute, 5 mm. long, sericeo-ciliate above; leaves triangular
or cordate, 10-12 mm. long and broad, glabrous beneath, bipinnate,
the lowest segments 3-5-lobed, the upper ,3-lobed or entire, the
lobes obtuse; flowers from below the middle of the petioles; calyx
7-9 mm. long, glabrous or the triangular acuminate lobes with a few
sericeous trichomes; corolla rose-colored, 17-20 mm. long, the
obovate retuse petals coalescent into a tube 5-6 mm. long; stamens
in a globose head, the free filaments elongate; carpels 3 mm. long,
shortly beaked, dorsally ciliate.
Ayacucho: Andes of Ayacucho, 3,600-4,000 meters {Pearce,
type). — Puno: Cordillera de Puno (Lechler 1713).
Nototriche alternata Hill, Trans. Linn. Soc. ser. 2. Bot. 7: 241.
1909.
Pulvinate, the leaves borne as in the related N. turritella but the
petioles (8-10 mm. long) dilated to a broad flattened cuneate
expansion merging into the flabellate 7-parted leaf-blade, this 6-7
mm. wide, above densely below sparsely stellate tomentose, the
Flora of Peru 537
subentire or 3-5-parted segments 1.5-2.5 mm. long, the linear-
obtuse lobes inflexed and glabrous apically; flowers slightly above
the petiole base; calyx about 1 cm. long, tube subglabrous, subacute
lobes 4 mm. long, stellulate; corolla about 1.5 cm. long, obovate
petals to 5 mm. wide the basal tube 5 mm. long; carpels lignose,
8 mm. long, stellate-hirsute, the beaks 3 mm. long. — Remarkable
for the large carpels but mature ones unknown for N. sajamensis
and N. Meyeni, with which species, and N. obcuneata, it grows,
judging from the mixture of Stubel's types at Berlin (Hill).
Ayacucho: Near Cangrao, Prov. Huanta, Raimondi (det. Ul-
brich). — Tacna: Volcano Tacora, 4,200 meters (Stiibel, 108 packet 2,
type).
Nototriche anthemidifoHa (Remy) Hill, Bot. Jahrb. 37: 579.
1906; 254. Sida anthemidifoHa Remy, Ann. Sci. Nat. s^r. 3. 6: 356.
1846. Malvastrum anthemidifolium (Remy) Gray, Bot. U. S. Expl.
Exped. 152. 1854. N. discolor Turcz. Bull. Soc. Nat. Mosc. 36,
pt. 1: 567. 1863. A^. cheilanthifolia Turcz. I.e.
Depressed, the caudex branched, the rosulate leaves with
petioles 2 cm. long, stipules adnate below the middle, the mem-
branous free part 7 mm. long, filiform, acute, and as vagina glabrous
dorsally, densely setose-ciliate marginally; petioles sulcate above the
stipules, glabrate; leaves ovate or ovate-oblong, pinnate or bipinnate,
1-2 cm. long, stellate-tomentose above, subglabrous beneath, the
short lacinulae obtuse, entire or trifid, ciliate at tip; flowers from
below the middle of the petioles; calyx 6 mm. long, tube glabrous,
lobes sparsely stellate, acute; corolla rose-colored, 10-12 mm. long,
the broadly obovate petals truncate or retuse, the tube 3 mm. long;
carpels 8-12, 3.5-4 mm. long, sericeous pilose dorsally, the trichomes
6 mm. long, the beaks subulate 2 mm. long; seeds rugose (Weddell
specimens). F.M. Negs. 35502; 35504. Illustrated, Hill, I.e. pi. 27,
fig. 8 (plant); pi. SO, fig. 1 (flower and stipules), figs. SI, S2 (carpel
and seed).
Arequipa: Near Puno (Weddell jU46).— Tacna.: Tacora (Weddell;
Steinmann). To Argentina.
Nototriche aretioides (Gray) Hill, Bot. Jahrb. 37: 580. 1906;
231. Malvastrum aretioides Gray, Bot. U. S. Expl. Exped. 1: 153.
1854.
Depressed, caespitose, pulvinate, the caudex often much branch-
ed; stipules adnate to above the middle of the i>etioles (6-15 mm.
538 Field Museum of Natural History — Botany, Vol. XIII
long), the free part oblong, 3 mm. long, densely stellate, the broad
membranous vagina armed dorsally with subspinose stellate tri-
chomes; leaves semicircular, trifid, 3 mm. long, 8 mm. wide, densely
and finely stellate-tomentose above, glabrous beneath with 3^
obovate obtuse lobules; flowers subsessile, about middle of petiole;
calyx acutely lobed nearly to middle, 7 mm. long, lobes sparsely
tomentose, hirsute within and marginally; corolla blue-lilac, 1.5 cm.
long; petals broadly obovate, emarginate, connate for 4 mm.;
carpels 7-8, subulate-rostrate, dorsally long-sericeo-ciliate. — Illus-
trated, Hill, I.e. pi. 29, figs. 15, 16 (leaves).
Lima: Cosmopolia, 832 (det. Johnston). Casa Cancha (Picker-
ing, type). — Junin: Arapa Hacienda near Yauli, 4,400 meters,
Weberbauer 289; 221. Cerro de Pasco, 3063 (det. Ulbrich). Huaron,
1152 (det. Johnston). — Cuzco: Valle del Paucartambo, 3,700 meters
(Herrera 103). "Panti-thurpa" (Herrera).
Nototriche argentea Hill, Bot. Jahrb. 37: 585. 1906; 259.
Silvery even to the calyces with a sericeous tomentose indument,
depressed, caespitose; petioles 2-3.5 cm. long; stipules adnate to the
vagina 6 mm. long the free part subulate 8 mm. long, dorsally
glabrous; leaves broadly ovate or pentagonous, ternate or somewhat
5-parted, 2-2.5 cm. long, about 2 cm. wide, less densely sericeous
beneath, the laciniae of the bipinnatifid segments spathulate, crenate,
to 3 mm. long, 1-1.5 mm. wide; flowers inserted below the middle
of the petioles, subsessile; calyx 11-12 mm. long, the lobes narrowly
triangular, acute; corolla white or pale lilac, 2-2.5 cm. long, the
obovate retuse petals connate into a tube 3.5 mm. long; stamens
in an oblong head ; carpels dorsally sericeous, long-biaristate (imma-
ture).— F.M. Neg. 9339. Illustrated, Hill, I.e. pi. 29, fig. 1 (leaf and
flower).
Ancash: Above Huaraz, 4,400 meters, Weberbauer 3103, type;
226. Across to Chonta, Weberbauer 2777; 224. — Tacna: Candarave,
sandy tola heath, Weberbauer 7373 (det. Ulbrich).
Nototriche argylloides Hill, Trans. Linn. Soc. ser. 2. Bot. 7:
245. 1909.
Depressed, caespitose; leaves rosulate; petioles 2-4 cm. long,
stipules adnate above the middle, the free part membranous, linear-
lanceolate, acute, to 1 cm. long, glabrous or marginally very sparsely
stellate; leaves various, pentagonous to more or less ovate, palmately
lobed or parted, 10-18 mm. long, 8-12 mm. wide, the segments
Flora of Peru 539
multilobed or somewhat bipinnatifid, incised-crenate, often crisped,
the medial the largest, ashy puberulent-tomentulose above, medially
glabrous beneath; flowers about the middle of the petiole between
the stipules; calyx 6-9 mm. long, acute lobes 2-3 mm. long, tomentu-
lose both sides, the tube glabrous; corolla roseate or rose-violet,
18-20 mm. long, petals broadly obovate, the tube 5 mm. long;
carpels about ten, 7.5 mm. long, beaks 2 mm. long, densely stellate-
ciliate. — Petioles and calyces suffused with reddish-purple the latter
with conspicuous pink-purple veins. Illustrated, Hill, I.e. pi. 27,
figs. 6, 7 (plants); pi. SO, figs. 18, 27 (carpels).
Arequipa: Puno to Arequipa, Weddell ItS9U. Vincocaya, 4,376
meters {HiU 80, type; Copeland).
Nototriche arista ta Hill, Trans. Linn. Soc. ser. 2. Bot. 7:
255. 1909.
Depressed, the caudex branched, the rosulate leaves with petioles
2 cm. long, stipules adnate above the middle, the vagina 5 mm.
wide, the free part basally subulate, acute, 5 mm. long, all glabrous
except for the sparsely setose-ciliate margins; leaves more or less
ovate-elliptic, 10-18 mm. long, densely stellate-tomentose above,
glabrous below, palmately or subpinnately parted, the segments
rather crowded, entire or 3-7-cleft, the linear-oblong obtuse lacinulae
3-4 mm. long, ciliate apically; flowers at middle of petioles; calyx
10 mm. long, very sparsely ciliate without, the triangular acute
lobes stellate-ciliate within; corolla rose-violet, 16-18 mm. long, the
concave spathulate obovate petals coalescent into tube 4-5 mm.
long; stamens few, borne in a globose head; carpels 8-12, 8-9 mm.
long, dorsally setose-ciliate, the subulate beaks 4-5 mm. long. — Like
a large form of N. longirostris but the larger carpels setulose instead
of short-stellate, the longer beaks with stiff bristles (Hill). It seems
to be N, longissima except for the long softer pubescence of the
carpels, more ovate leaves. Illustrated, Hill, I.e. pi. SO, figs. 25, 26
(carpels).
Cuzco: Pacechac Valley near Urubamba, 4,500 meters {Hill 76;
77, type).
Nototriche armeriifolia Hill, Kew Bull. 160. 1935.
Depressed, caespitose, pulvinate, with ligneous branched sub-
terranean caudex; stipules adnate to the petiole and with it forming
a quasi vagina (3-3.5 mm. wide, 8-10 mm. long), the lanceolate free
part 1.5-2 mm. long, densely hirsute (dorsally glabrous) as the
540 Field Museum of Natural History— Botany, Vol. XIII
petioles with whitish stellate trichomes 3 mm. long; leaves linear,
1.5 cm. long, about 1 mm. wide, marginally incurved, with 1 or 2,
rarely 3 linear lobes about 2 mm. long, 5 (or so) mm. below the tip,
densely stellate velutinous-hirsute above, glabrous beneath; flowers
borne a little below the middle of the petioles; calyx nearly medially
5-lobed, 7.5 mm. long, nearly glabrous, the acute lobes with a few
long trichomes, within at base with 5 papillose glands and long-
pilose; corolla pale violet, 13-14 mm. long, the basally obovate-
cuneate petals 4 mm. wide, coalescent at base into tube 4 mm. long;
anthers many, globosely capitate; carpels 1.75 mm. long, beaks.
0.5-0.75 mm. long, dorsally covered with trichomes 3-4 mm. long.
Puno: Valley below Cunurana Range, Santa Rosa de Ayaviri:
to Sicuani (Stafford 362, type, Herb. Kew).
Nototriche artemisioides Hill, Bot. Jahrb. 37: 580. 1906; 228. ,
Depressed, caespitose, often pulvinate; petioles 1.5 cm. long, the
medially adnate stipules both sides as leaves and calyx lobes without
densely stellate-tomentose; free part of stipules linear, acute, 6-9;
mm. long; leaves rotund-cordate, trifid, 1 cm. long, 12 mm. wide,
the segments 3-lobed, the lobules 2-3-lacinulate; flowers subsessile,
borne below the middle of the petiole; calyx 12 mm. long, the linear-
lanceolate subacute lobes 8 mm. long; corolla less than 2 cm. long,
the violet obovate petals retuse, connate for 5 mm.; carpels 7 mm.
long, biaristate (beaks 3 mm. long), densely stellate-hirsute dorsally
and ventrally, the seeds sulcate dorsally. — Stamens relatively few
in a somewhat elongate head (Hill). N. Lobbii (Baker) Hill, 228,
pi. 28, jig. 8 (leaf), origin unknown, ex char, is similar but branching,
the primary leaf segments petiolate, more lacinulate, petals 2 mm.
connate. Illustrated, Hill I.e., pi. 28, fig. 6 (leaf with flower). F.M.
Neg. 9340.
Cajamarca: Grass steppes near Hualgayoc, 3,400 meters, Weber-
bauer U068, type; 271.
Nototriche Azorella Hill, Bot. Jahrb. 37: 582. 1906; 219.
Densely caespitose, pulvinate, with much branched ligneous
subterranean caudex; leaves numerous, congested, imbricate, densely
condensed in small cylinders; petioles 5 mm. long; stipules adnate
nearly to the leaf -blades, the resulting vagina membranous, 3-4 mm.
wide, the free part somewhat herbaceous, oblong, acute, 2 mm. long,
the face glabrous as the vagina but dorsally and marginally densely
stellate-ciliate or setose; leaves semicircular or orbicular, 5-7-
Flora of Peru 541
flabellately divided, 3 mm. long, 4 mm. wide, densely stellate-
tomentose both sides, the segments entire or 3-5-crenate-lobed the
medial one often 5-lobate; flowers below the middle of the petioles;
calyx 5 mm. long, subacute lobes 1 mm. long, stellate without,
glabrous within; corolla roseate-white, 7 mm. long, the petals nearly
truncate, coalescent into hirsute tube 2.5 mm. long; stamens about
14, the head exserted; pistils and carpels unknown. — With N.
congesta the two smallest perennial species; they both form hard
cushions, due to the density of the rather vertical caudex branches,
like those of the genus Azorella, sometimes 1 dm, across; the dense
cylinders are about 1 cm. across, only the upper leaves living, and
the center of each tiny rosette is occupied by a flower (Hill). Illus-
trated, Hill, I.e. pi. 27, fig. S (plant); pi. 28, fig. 15 (leaf and flower).
Puno: Poto to Ananca, Prov. Sandla, 4,600 meters, Weberbauer
957, type; 219. — Tacna: In small ravines in shelter of boulders,
5,100 meters, Volcan Tatora, Werdermann 1152; 1U77 (det. Ulbrich).
Chile.
Nototriche borussica (Meyen) Hill, Bot. Jahrb. 37: 579. 1906;
236. S>ida boriLSsica Meyen ex Walp. Nov. Act. Acad. Leop.-Carol.
19:Suppl. 1:308. 1843.
Depressed, caespitose, the silvery tomentose leaves hemispheri-
cally congested; stipules adnate to 2 mm. below the middle of the
petioles (these 1 cm. long) the subulate free part 3 mm. long; vagina
dorsally and petioles with sides minutely stellulate- tomentose;
leaves reniform, flabellately and deeply 5-7-parted, 3-4 mm. long,
7 mm. wide, segments 3-5-lobed, lobes cuneate, softly stellate both
sides; flowers from below middle of petioles; calyx 9 mm. long, ovate
lobes obtuse, velutinous stellate both sides; corolla about 1.5 cm.
long, white with dark violet or almost black stripes. — Meyen men-
tioned the species first in his Reise um die Erde 2: 31 and explained
his choice of name from the resemblance of the color of the flowers
to those of the flag of Prussia. Easily known by its long free petioles,
soft white tomentum, and cuneate leaf -lobes (Hill). F.M. Neg.
32637.
Arequipa: Vulcan El Misti, Alto de los Huesos, in black volcanic
ash, 3,800 meters (Meyen, type). — Tacna: Tacora, 4,670 meters,
Meyen.
Nototriche candaravica Ulbr. Notizbl. Bot. Gart. Berlin 11:
531. 1932.
542 Field Museum of Natural History— Botany, Vol. XIII
Pulvinate, the rhizome 4-6 mm. thick, the leaves very densely
rosulate at the tips of the erect branches; petioles 5-7 mm. long,
stipules (free part) oblong-lanceolate, about 3 mm. long, sparsely
villous; leaves cuneately suborbicular, deeply multilobulate, 2.5-3
mm. long, yellowish villous-stellate; flowers subsessile on pedicels
1.5 mm. long; calyx 7-8 mm. long; corolla violet, whitish toward
villous base, the tube 1.5-2 mm. long; petals oblong, nearly 8 mm.
long; stamen tube 5 mm. long; stamens in a globose head, the
filaments elongate. — Related to N. nigrescens with much larger
leaves and flowers; the polsters formed by the branches and leaves
are 1-2 cm. high, 1.5-3 cm. across, and the stamens are in an oblong-
cylindrical head (Ulbrich). It seems to be very near N. holoserica
but the leaves shorter in type (young).
Tacna: Tola heath, Candarave, 4,000 meters, Weberbauer 7372,
type.
Nototriche Castelnaeana (Wedd.) Hill, Bot. Jahrb. 37: 579.
1906; 257. Malvastrum Castelnaeanum Wedd. Chlor. And. 2: 283,
pi. 80A. 1857.
Caespitose, canescent-tomentose, the naked caudex simple or
sparsely branched; petioles little dilated; leaves pinnatisected, 1-2
cm. long, with 4-6 3-5-parted segments the lower the larger, the
lacinulae obtuse, plane, velvety stellate- tomentose both sides;
flowers sessile, between the setaceous stipules at or a little below the
middle of petioles; calyx campanulate, about 9 mm. long, softly
tomentose, the lobes triangular-lanceolate; petals about 2 cm. long,
scarcely emarginate; anthers in an oblong-cylindrical head. — Im-
perfectly known. F.M. Neg. 35503.
Cuzco: Cordillera de Cuzco, Castelnau, type. Without locality,
Maclean; Ruiz & Pavdn.
Nototriche coccinea Hill, Bot. Jahrb. 37: 583. 1906; 219.
Caespitose, pulvinate, the caudex densely branched; petioles
1-1.5 cm. long, the stipules adnate to above the middle, free part as
vagina membranous, linear, subacute, 4-5 mm. long, with the
petioles little tomentose, dorsally glabrous, marginally stellate-
pilose; leaves reniform, 5-lobate, 6 mm. long, 9 mm. wide, stellate-
tomentose above, nearly glabrous beneath, the lobes 3-5-lobulate,
the lobules broadly obovate-crenate; flowers below the middle of the
petioles; calyx 7-8 mm. long, tube glabrous, subovate acute lobes
stellate-pilose without; corolla scarlet, 1 cm. long, petals irregularly
Flora of Peru 543
emarginate, tube about 3 mm. long; carpels 8.5 mm. long, stellate-
hirsute dorsally, beaks 1 mm. long; seeds sulcate. — Branches slender,
forming a compact cushion, the root system much divided (Hill).
Illustrated, Hill, I.e. pi. SO, fig. 17 (carpel).
Ancash: Above Lake Yanganucos above Yungay, 4,600 meters,
Weherhauer S276, type; 226.
Nototriche condensata (Bak. f.) Hill, Bot. Jahrb. 37: 579.
1906; 224. Malvastrum condensatum Bak. f. Joum. Bot. 29: 363.
1891.
Depressed, caespitose, pulvinate, with stout branching caudex;
petioles to 2 cm. long; stipules adnate to about the middle of the
petiole, with it forming a membranous vagina 3-4 mm. broad, the
linear acute free part about 1 cm. long, as the petiole glabrous
dorsally, sparsely stellate marginally; leaf -blades broadly obcuneate
or flabelliform, more than medially 5-7-parted, to 1 cm. long, 10-12
mm. wide, densely ashy-stellate- tomentose above, sparsely so or
nearly glabrous beneath, the medial segments 3-5-lobed; flowers
borne a little below the middle of the petiole; calyx about 8-13 mm.
long, more or less densely stellate-tomentose, the subacute ovate
lobes tomentose marginally; corolla dark blue, 2.5 cm. long, the
obovate-oblong petals coalescent into a tube only 1-2 mm. long;
stamens forming a globose head, the free portions of the filaments
elongate; carpels 10-12, 7 mm. long, dorsally stellate below, the
beaks stellate-ciliate, 3 mm. long. — The leaf resembles the palm
of a hand, with finger-like lobes, some of which bear minute in-
curved lateral lobulae (Hill). Baker described the petals as only
8 mm. long, perhaps from not fully developed flowers, while Weddell
referred the type to N. phyllanthos. Conceivably the species when
better known may be found to include N. sajamensis, N. ticsanica
and A^. pulvinata Hill, Kew Bull. 18. 1928 of northern Chile, as it
does not seem probable that the differences in pubescence-distri-
bution, leaf-division and size described for these plants are really
specific factors.
Puno: Near Ayapata, Lechler 1972, type. Herb. Kew; same num-
ber, part. Herb. Paris.
Nototriche congesta Hill, Bot. Jahrb. 37: 582. 1906; 221.
Habit of the similar N. AzoreUa; petioles 6-7 mm. long; vagina
2-3 mm. wide; stipules filiform-subulate, acute, 2-3 mm. long,
glabrate as vagina except stellate dorsally and marginally; leaves
544 Field Museum of Natural History— Botany, Vol. XIII
more or less rhombic or broadly obovate, acute, 5-parteci medially,
about 4 mm. long and wide, somewhat stellate-tomentose both sides,
the middle segments shortly 3-lobate, the lateral inflexed; corolla
6 mm. long, petals more or less retuse, tube 1.5 mm. long. — Other-
wise like the related species; the leaves form a larger and less dense
mass; branches above covered with dead leaves, below only the
vaginae remain, these with bristle-like trichomes; large orange-
colored glandular nectaries at calyx base (Hill). Illustrated, Hill,
I.e. pi. 30, jig. 8 (glandular petals). F.M. Neg. 9344.
Puno: Suchez, Prov. Sandia, 4,500 meters, Weberhauer 1018,
type; 219.
Nototriche cupuliforme Krap. Bol. Soc. Argent. Bot. 5: 71.
1953.
Depressed, essentially glabrous, sometimes 4 cm. across, thej
ligneous caudex a cm. in diameter, and unique in flower-structure,
the wine red corolla with the segments borne below a cupuliform
extension of the corolla-tube; stipular vagina 1.5 cm. long, 2.5 mm.
wide at base, 1.5 mm. wide at apex, the triangular-lanceolate free;
portion 3-4 mm. long, about 1 mm. wide; leaves palmately nerved,
7-parted, the subequal subacute divisions sometimes uniciliate
at tip; flowers solitary at the free part of the stipules; calyx 5 mm.
long, teeth 2.5 mm. long, 2 mm. wide, with a few simple or stellate
trichomes at the margins; corolla glabrous, about 9 mm. long, the
tube 1.5 mm. long by 2 mm. wide, extended about 1 mm. above'
the base of the obovate petals; anthers and stigmas (6) globosely
congested; carpels 1-ovulate, glabrous or nearly. — The remarkable,
extension of the tubular union of the corolla segments is unique
(author). Illustrated, I.e. 69, fig. 6, c, d, f (leaf, corolla).
Puno: Pampa de Lacka, Prov. Carabaya, 4,360 meters, (Vargc
7119, type).
Nototriche digitulifolia Hill, Kew Bull. 127. 1948.
Depressed, caespitose, the polsters 2-4 cm. wide; caudex morel
than 1.5 dm. long; leaves crowded, white pubescent; petioles 12 mm.j
long, long-stellate above the stipules; stipules medially connate, the
resulting vagina membranous, only dorsally and marginally longi
stellate-tomentose, the free part linear-lanceolate, 8-9 mm. long;!
lobules 7, dissected to base into erect spathulate segments 1.3-5!
mm. long; flowers at middle of petioles; calyx 1 cm. long, the narrow
acute lobes strigose-stellate, the tube glabrous; corolla tube 3 mm.
Flora of Peru 545
long, stellate pilose, the obovate lobes 12 mm. long; anthers in
globose head; carpels 6 mm. long, beaks 2.75 mm. long, densely
stellate. — Allied to N. pedatiloba Hill but stipules narrower, leaf-
segments finger-form, calyx lobes longer, corolla tube shorter (Hill).
Illustrated, Hill, I.e. 126.
Puno: San Antonio de Esquilache, 4,500 meters (Stafford 765,
type).
Nototriche dissecta Hill, Trans. Linn. Soc. ser. 2. Bot. 7:
238. 1909.
Pulvinate, the hemispherical leaf-rosettes about 4 cm. across,
the petioles, these 1 cm. long, stipules and leaves lanately stellate-
tomentose; stipules adnate above the middle, the free part her-
baceous, linear, acute, 8-9 mm. long; leaf blades semicircular,
deeply trifid, 6 mm. long, about twice as wide, the segments equally
bitemate, 2-7-lobate, the linear-oblong lobes subacute; flowers
shortly pedicellate at the middle of the petioles; calyx lanate, 8-9
mm. long, the narrowly triangular lobes acute; corolla rose colored,
17 mm. long, the oblong-obovate subtruncate petals coalescent into
tube 6 mm. long. — The tomentum of long stiff bristles and the three
main segments of equal size are distinctive characters (Hill). Illus-
trated, Hill, I.e. p/. 28, fig. IS (leaf).
Apurfmac: Andahuaylas, 3,900-4,300 meters (Pearce, type).
Nototriche epileuca Hill, Bot. Jahrb. 37: 585. 1906; 252.
Depressed, caespitose, the laxly rosulate leaves white-covered;
petioles 3-4 mm. long, glabrous as the medially adnate stipules, their
free part linear, acute, 5 mm. long; leaves triangular, trifid, white-
tomentose above, glabrous beneath, segments pinnatifid, lobes
deeply 3-cleft, the somewhat involute laciniae crenate; flowers
subsessile; calyx 10-11 mm. long, lobes 6-7 mm. long, linear-
lanceolate, subacute, sparsely stellate puberulent without, tomentose
within; corolla blood-red, 2.5 cm. long, its obovate-oblong petals
retuse, adnate at base to staminal tube; carpels about 20, 8 mm. long,
each with 2 broad stellate beaks 4 mm. long, dorsally stellate. —
Petals practically free, stamens in an oblong head; resembles N.
stenopetala but the leaf-segments many and obtuse instead of few
and acute (Hill). Illustrated, Hill, I.e. pi. SO, fig. 28 (carpel).
F.M. Neg. 9345.
Ancash: Across to Chonta, 4,400 meters, Weberbauer 2801 ,
type; 224.
546 Field Museum of Natural History— Botany, Vol. XIII
Nototriche erinacea Hill, Kew Bull. 127. 1948.
Caudex stout, sparsely branched, crowned by rosettes 1 dm.
wide or wider, the leaves to 7 cm. long, densely stellate-tomentose
(drying flavescent); petioles 3-4 cm. long, free part of stipules
linear, acute, 1.5 cm. long, adnate part with vagina 2-3 cm. long,
stellate; leaf blades ovate-lanceolate, 2.5-3 cm. long, 2 cm. wide,
bipinnatifid, the primary segments 5 each side, the medial 2-2.8 cm.
long, bipinnate with crenate lobules, the terminal ones rounded, the
lateral oblongish, acute; flowers violet, subsessile among the stipules;
calyx 17 mm. long, tube 7 mm. long, lobes 1 cm. long, narrowly
triangular, stellate-tomentose without, strigose within; corolla tube
3 mm. long, sparsely pubescent or glabrous, petals narrowly obovate,
11 mm. long, 6 mm. wide; anthers in globose head; carpels 1 cm.
long, beaks 5 mm. long, dorsally sericeous with trichomes 5 mm.
long.— Similar to N. sulphurea but leaves and flowers smaller, some
calyx trichomes stoutly stipitate, corolla tube longer, petals not
tomentose, carpels long-rostrate (Hill). Illustrated, Hill, I.e.
Puno: San Antonio de Esquilache, 4,500 meters, stony slopes
(Stafford 758, type).
Nototriche estipulata Hill, Kew Bull. 128. 1948.
Caudex stout, to 1.5 dm. long, the densely crowded estipulata
leaves forming an oblong-ovoid head at the top; petioles about
1 cm. long, subglabrous above, margins and beneath stellate-pilose;
leaf blades 3 mm. long, 6 mm. wide, with about 9 short obtuse
fleshy lobules stellate-pilose both sides; flowers subsessile, 2 mm.
above base of petioles; calyx 8 mm. long, stellate-pilose, the tube
5-6 mm. long, lobes fleshy at tip; corolla tube stellate-pilose, 4.5
mm. long, suboblique obovate lobes 3-4 mm. long; anther head]
globose; carpels (immature) long-sericeous dorsally and on beaks, 1
stellate-pilose toward the base. — Affine N. famatinensis Hill of]
Argentina with stipules and especially N. compacta (Gay) Hill of!
Chile but leaves less lobate, corollas larger, indument longer and]
laxer (author). Illustrated, Hill, I.e.
Tacna: Volcan Tacora, 4,800 meters, Werdermann IWl, type.
Nototriche famatinensis [Hieron.] Hill, Bot. Jahrb. 37: 581. j
1906; 227.
Pulvinate from caudex, to 2 cm. thick; petioles more or less]
tomentose, 1-2 cm. long; stipules more than medially adnate, I
subulate free part to 1 cm. long, above and marginally stellate-]
Flora of Peru 547
pilose; leaves semicircular, flabellately to middle 7-11-lobate, 6 mm.
long, 11 mm. wide, above densely, beneath sparsely stellate-tomen-
tose, lobes entire to 9-lobulate, the medial 3-lobulate, the others
laterally 1-lobulate; flowers sessile at middle of petioles; calyx about
8 mm. long, the lobes narrowly ovate, acute, stellate-tomentose,
nearly glabrous within; corolla 7-9 mm. long, tube 2-3 mm. long;
carpels shortly birostrate, dorsally ciliate. — As elsewhere in this
work the author's name in brackets indicates that it was unpublished,
unless in synon)miy. Illustrated, Hill, I.e. pi. 28, fig. 7 (leaf).
F.M. Neg. 9346.
Tacna: Volcan Tacora, 4,800 meters, Werdermann lltOl (det.
Ulbrich). Western Argentina.
Nototriche flabellata (Wedd.) Hill, Trans. Linn. Soc. ser. 2.
Bot. 7: 222. 1909. Malvastrum flahellatum Wedd. Chlor. And. 2:
281. 1857.
Depressed, caespitose, velvety-tomentose; petioles 2-4 cm. long;
free part of vagina membranous, linear-subulate, acute, 1 cm. long,
densely floccose-stellate as the petioles; leaf blades broadly obovate-
cuneate to rounded, flabellately 5- or 7-parted, about 1.5 cm. long,
12-15 mm. wide, floccose-stellate both sides or glabrate beneath, the
lateral segments obovate-cuneate, 5-7, unequally incised-crenate,
the middle segment more or less trifid, the lobes incised-
crenate; flowers from nearly the middle of the petiole; calyx 18 mm.
long, the triangular-acute lobes glabrous within; corolla puniceous,
(1) 2-2.5 cm. long, the oblong, irregularly retuse petals 7 mm. wide,
scarcely coalescent; stamen column about 2.7 cm. long; carpels
9 mm. long, dorsally stellate, the beaks about 5 mm. long (Lechler
1972, in part), carpels nearly mature 7.5 mm. long, stellate below,
stellate ciliate above, the beaks about 2.5 mm. long (Weberbauer).
— The Herrera determination has been questioned in herb, by
Killip, perhaps with reason since D'Orbigny's specimen from La Paz
(perhaps the true type) was thought to have small flowers; this is
discussed by Hill; the anthers, too, as here defined are long-filamen-
tose and more oblong than globose in disposition. F.M. Neg. 23745
(Mandon). Illustrated, Hill, I.e., pi. 29, fig. 2 (leaf and calyx);
pi. 30, figs. 7, 8 (flowers), fig. lU (carpel).
Cuzco: Pachatusdn, Herrera 257 U, pt. (det. Ulbrich). — Puno:
Near Ayapata, Lechler 1972, pt. Poto to Anaca, Prov. Sandia,
4,400 meters, Weberbauer 961 . Adjacent Bolivia.
548 Field Museum of Natural History— Botany, Vol. XIII
Nototriche foetida Ulbrich, Notizbl. Bot. Gart. Berlin 11:
530. 1932.
Foetid glutinous rhizomatous perennial, the dense erect column-
ariform branches forming cushions 2.5-4 cm. high; stipules connate,
3-4 mm. long; petioles sparsely stellate, 5-7 mm. long; leaves
cuneately suborbicular, 2-3 mm. long, 3-5 mm. wide, crenately
incised, densely white tomentose above, sparsely stellulate or sub-
glabrous beneath; flowers white, subsessile at tips of branches;
calyx 8-9 mm. long, the lanceolate-oblong obtusish lobes 2.5-3 mm.
long, sparsely stellulate, otherwise glabrous; petals obovate, ciliate
toward base, about 8 mm. long; stamen tube cylindrical, 4-5 mm.
long, anthers globosely capitate. — Well-marked by the odor and
glutinous condition; habit similar to N. obcuneata with parted
leaves, reddish flowers, and to N. glacialis, the flowers opening pale
lilac (Ulbrich).
Tacna: Tola heath, Candarave, 4,500 meters, Weberhauer 7361,
type.
Nototriche glacialis Ulbr. Bot. Jahrb. 54: Beibl. 117: 62. 1916.
Similar to N. obcuneata (fide author) but with much slenderer
longer rhizomes; petioles about 8 mm. long, the free part of stipules
4-5 mm. long, obtuse, sparsely appressed stellate; leaves broadly
obovate, 4-6 mm. long, 6-9 mm. wide, the flabellate lobes some-
times with a few minute stellate trichomes beneath, 6-9-crenate;
flowers borne above the middle of the petioles, pale lilac, finally
white, 9-10 mm. long; calyx nearly 3 mm. long, sparsely and mi-
nutely stellate, glabrous within; petals basally fimbriate, clawed,
somewhat retuse; anthers subglobosely capitate, not exserted; ovary
stellate-tomentose below, penicillate apically with long sericeous
trichomes. — The slender rhizomes are a dm. or two long, the leaves
olive-green beneath, the flowers small; type from the base of glacier
Cuspicocha, in stone-drift.
Junin: Above Hacienda Acopulca, 4,900 meters, {Weberbauer,
6523, type).
Nototriche gracilens Kilhp & Macbr., sp. nov.
N. armeriifoliae similis sed floribus majoribus et imprimis
antheris in cylindricam columnam instructis facile distinguenda;
petiohs circa 1 cm. longis; vagina vix 1.5 mm. lata glabra ut stipulae
parte libera subulato-lineari 3-5 mm. longa; lamina 2-2.5 cm.
longa, 1-2 mm. lata integra vel breviter 1-2-lobulato vel laminae
Flora of Peru 549
folii fertilis interdum simpliciter pinnatilobata 6-7 mm. lata, lobis
circa 3-4 mm. longis, supra dense stellato-tomentosa, subtus glabra;
flores subsessiles in petiolo inter stipulas producti; calyx 1 cm.
longus, lobis triangularibus acutis solum intus tomentosis; corolla
2-2.5 cm. longa, petala in tubum vix 2 mm. leviter connata fere
10 mm. lata. — The type in the National Herbarium bearing the
inscription "probably new," E. P. Killip; I have shared in its
publication with my friend's kind permission.
Huancayo: Laguna Huacracocha, 5,000 meters, Soukup S60U,
type.
Nototriche Herrerae Ulbrich ex Hill, Kew Bull. 129. 1948.
Resembles N. pedicularifolia but the leaves 5.5-7 cm. long,
densely tomentose, the flowers much larger; stipules about 17 mm.
long, linear, stellate-tomentose, the adnate base and vagina 2-3.5
cm. long; petioles 3.7 cm. long, free part 1.7 cm. long; leaf blades
ovate, 2.5-3 cm. long, 2.3 cm. wide, 5-lobed, the lobes pinnately
multilobulate; flowers between the stipules; calyx tube 7 mm. long,
lobes 1 cm. long, acute, stellate-tomentose; corolla large, drying
purple with yellowish base, tube 2 mm. long, pubescent, lobes
obovate, emarginate, 2.3 cm. long, pubescent within at base; head
of anthers long-obovoid ; carpels long-sericeous. — Seems, ex char., to
be N. sulphurea. Illustrated, Hill, I.e.
Cuzco: {Herrera S679c, type, Herb. Kew).
Nototriche holoserica Hill, Kew Bull. 248. 1927.
Pulvinate; petioles 6-8 mm. long; stipules adnate above the
middle, the free part filiform or subulate, to 5 mm. long; leaves
lanate, stellate-tomentose both sides, semicircular or reniform, about
6 mm. long, 9 mm. wide, multilobulate, the lobes 1.5-2.5 mm. long;
flowers shortly pedicellate; calyx 8-9 mm. long, the lobes lanate;
corolla blue, about 1 cm. long, the tube 4 mm. long, hirsute without;
carpels beaked, stellate-tomentose below. — Like N. compacta (Gay)
Hill of Chile but sericeous lanate, the leaves multilobulate (Hill).
Cuzco: Valle del Paucartambo, 3,800 meters, (Herrera 2S90, fide
Herrera). "Turpai" (Herrera). Chile.
Nototriche lanata Hill, Trans. Linn. Soc. ser. 2. Bot. 7: 229.
1909.
Habit of N. flabellata, but the leaves much more deeply parted,
the tomentum more sericeous; leaves rather laxly rosulate; petioles
550 Field Museum of Natural History— Botany, Vol. XIII
1.5-2 mm. long; stipules adnate above the middle, the membranous
vagina 2-3 mm. broad, the free part subulate, 5-7 mm. long, sparsely
stellate or glabrous except for the lanate-stellate margin; leaf -blades
almost rotund, 12 mm. long and broad, flabellately 5- or 7-9-parted,
above sparsely, beneath densely lanate-stellate, the larger segments
cuneate or ovate-cuneate and pinnatifid, the lobes and smaller
lateral segments entire, 3-lobulate or incised, with short congested
linear obtuse lacinulae; flowers from the middle of the petioles;
calyx about 9 mm. long, lanate, the acute lobes tomentose within;
corolla rose- violet, 18 mm. long, the broadly obovate retuse petals
coalescent into a tube 5 mm. long; carpels about 8, nearly mature
6 mm. long, the broadly subulate beaks 2 mm. long, shortly stellate-
pilose. — Type from northeast side of Lake Titicaca. Seems, as
suggested by Hill, more or less allied to N. pedicularifolia, but the
anthers not described. Illustrated, Hill, I.e. pi. 29, fig. 18 (leaf).
Puno (probably) . Adjacent Bolivia.
Nototriche longirostris (Wedd.) Hill, Bot. Jahrb. 37: 579.
1906; 252. Malvastrum longirostre Wedd. Chlor. And. 2: 281. 1857.
Depressed, rarely branched, the small plants usually consisting
of a single leaf -rosette; petioles to 2 cm. long, the stipules adnate to
above the middle, the vagina 4-5 mm. wide, more or less stellate
hirsute above, the triangular acutish free part to 5 mm. long, sub-
glabrous beneath, the margins setose-ciliate; leaves oblong-lanceo-
late, 1-1.5 cm. long; rather laxly stellate- tomentose above (or in age
glabrate), glabrescent beneath, bipinnate with 3-4 segments each
side, these as the terminal lobe 3-30-parted, the oblong or linear
lacinulae often ciliate apically; flowers below the middle of the
petioles; calyx campanulate, 8-9 mm. long, tube glabrous, the acute
lobes rather setose-stellate, marginally and within tomentose;
corolla drying white-violet, 1.5-2 cm. long, the obovate petals
scarcely emarginate, their tube 4 mm. long; stamens in a globose
head; carpels 6-8, stellate-tomentose below, setulose-ciliolate above,
about 7 mm. long, beaks 3 mm. long; seeds sulcate. — The specimens
as determined may be found to include segregates which are perhaps
only variants, however, when carpels are available, their pubescence
may distinguish it from N. longissima and N. aristata. Illustrated,
Solms, Bot. Zeit. 65: 120, pi. 2, figs. 5, 12; Hill, I.e. pi 30, figs. 22, 23
(carpel and seed); Weberbauer, 202 (plant). F.M. Neg. 23748.
Junln: Morococha, Raimondi (det. Ulbrich). La Oroya, Kalen-
born 117 (det. Killip). — Huancayo: Huanacocha, Soukup 3613 (det.
Flora of Peru 561
Killip, N. longissima). — Ayacucho: Mt. Razuhuillca, Weberbauer
7Jt92 (det. Ulbrich, N. Mandoniana). — Cuzco: Valle del Paucar-
tambo, 4,400 meters, Herrera 1012; 2322 (det. Macbride, N. Man-
doniana?). Andes of Cuzco, Castelnau, type; Gay. "Turpa"
(Herrera). Bolivia.
Nototriche longissima Hill, Bot. Jahrb. 37: 586. 1906; 257.
Caespitose, the caudex little if at all branched, sometimes a dm.
long or longer, the oblongish pinnatifid leaves laxly rosulate, petioles
to 3.5 cm. long, glabrous as the linear-oblong acute free part of the
stipules or these ciliate, 6 mm. long; leaves about 1.5 cm. long,
stellate-hirsute above, glabrous below, the segments subentire or
3-cleft, the linear lobes about 4 mm. long or little ciliate at the
subacute tips; calyx above the middle of the petioles, about 8 mm.
long, the acute lobes glabrous but ciliate within; corolla violet, 1.5
cm. long, the retuse p>etals connate into tube 4 mm. long; carpels
about 10, dorsally ciliate, with 2 ciliate beaks or awns 4 mm. long.
— Weherhauer 7771 is not typical as it has sparsely ciliate vaginas,
corollas 18-20 mm. long; N. pseudoglabra Hill of southern Bolivia,
to which it is referred, is soon glabrous except for the densely long-
pilose vaginas, much smaller flowers. Allied to A^. Pearcei, with
larger flowers. Illustrated, Hill, I.e. pi. 27, figs. 9, 10 (plants);
pi. SO, fig. 2k (carpel). F.M. Neg. 9349.
Ancash: Ocros to Chonta, 4,400 meters, Weherhauer 2785, type;
224.— Lima: Stony slopes, Rio Blanco, 3052 (det. Ulbrich, N.
longirostris). — Junin: In deep grass of swales, Cerro de Pasco, 3085
(det. Ulbrich). Laguna Pomacocha, Ochoa 251 (det. Killip); ^87.
— Apurimac: Sorococha Pass, 4,400 meters, Chincheros to Anda-
huaylos. West 3731 (det. Johnston, N. Pearcei).— Cuzco: Near
Auzanyate Glaciers, Weherbaiier 7771 (det. Ulbrich, N. pseudoglabra).
Nototriche longituba Burtt & Hill, Kew Bull. 130. 1948.
Rosettes 3-4 cm. broad; leaves about 2 cm. long, stipules nar-
rowly linear, the free part 5.5-8 mm. long, with petioles and vagina
(5-5.5 mm. long) densely and long-sericeous; fertile leaves simply
pinnate-lobed, the others bipinnatifid, primary lobes five, 3-4 mm.
long, 1.5-4 mm. wide, stellate- tomentose above, early sericeous
beneath, finally glabrous, the lobules setose-apiculate; calyx 4 mm.
long, lobes narrowly lanceolate, 4 mm. long, finely stellulate; corolla
white, about 15 mm. long, the tube 6-8 mm. long, glabrous; petals
obovate, truncate, 4 mm. long; anthers about 12; immature carpels
552 Field Museum of Natural History— Botany, Vol. XIII
beaked, very long-sericeous. — Distinctive by the long corolla tube;
allied to N. sericea Hill but blade of fertile leaves reduced, pubescence
stellate. Illustrated, Burtt & Hill, I.e. 131.
Puno: In turf, San Antonio de Esquilache, (A. Howell Williams,
type).
Nototriche Macleanii (Gray) Hill, Bot. Jahrb. 37: 579. 1906;.
230. Malvastrum Macleani Gray, Bot. U. S. Expl. Exped. 1: 152.
1854.
Resembles N. aretioides, but the blue-violet corollas 3-3.5 cm.
long; leaves imbricate, ashy-green; petioles 1.5-2 cm. long; stipules
adnate above the middle, the free part of the vagina linear-spathu-
late, 5-6 mm. long, lanate-stellate at the tips above, more or less
glabrous below; leaf-blades flabelUform, palmately divided, 5-6
mm. long, 10-12 mm. wide, lanate-stellate above, subglabrous
beneath, the segments deeply 3-lobed or the medial rarely 5-lobed,
the lobes obovate-crenate; flowers sessile, nearly at the middle of
the petioles; calyx 1.5 cm. long, the acute linear-lanceolate lobes
9 mm. long, sparsely stellate-pilose without; corolla blue- violet,
3-3.5 cm. long, the petals obcuneate, retuse, scarcely coalescent;
stamens in a cylindrical column, the free filaments elongate; carpels
immature, but apparently birostrate, dorsally densely stellate-
tomentose. — Illustrated, Hill, I.e., pi. 29, fig. 13 (leaf); Weberbauer,
202 (plant).
Junin: Arapa, near Yauli, 4,600 meters, Weberbauer 382; 221.
Above La Oroya, 4,300 meters, Weberbauer 1705; 223. Without
locality {Maclean, type).
Nototriche Mandoniana (Wedd.) Hill, Bot. Jahrb. 37: 579.
1906; 253. Malvastrum Mandonianum Wedd. Chlor. And. 2: 282.
1857.
Depressed, the rosulate leaves with petioles 2-2.5 cm. long,
stipules adnate to the middle the membranous free part linear-
lanceolate, acute, 1 cm. long, as the petioles and vagina glabrous
dorsally, setose-stellate marginally; leaves oblong-obovate or
elliptic-lanceolate, bipinnate, pinnate or sub-pinnate, 1.5-2.5 cm.
long, stellate-tomentose above, glabrous below or early sparsely,
stellate, lacinulae unequal, linear-oblong; flowers from middle of
petioles; calyx 8-10 mm. long, lobes ovate, acute, finely stellate-
tomentose without; corolla rose- violet, 1.5-2 cm. long, the obovate-
oblong petals retuse, coalescent into tube 5 mm. long; carpels about
Flora of Peru 553
9. — Differs from N. anthemidifolia especially in the velvety pu-
bescent cal3nc and the larger flowers (Hill). F.M. Neg. 9351.
Huancavelica: Cordillera Huaytara, 4,100 meters (Pearce).
Cordillera de Pachatusdn, Herrera 257 U, pt.— Puno: Near Poto,
Weherhauer 1002b; 219. Bolivian boundary, Weherbauer 1019.
Bolivia.
Nototriche MatthewsH Hill, Trans. Linn. Soc. ser. 2. Bot.
7: 248. 1909.
Depressed, caespitose, often pulvinate; petioles 1 cm. long;
stipules adnate to above the middle, the free part herbaceous,
subulate, 4 mm. long, tomentose above, glabrous dorsally, stellate-
pilose marginally; leaves ovate-oblong, deeply palmately parted,
rarely trifid or somewhat pinnatifid, 1.5 cm. long, lanate-sericeous
stellate-tomentose above, glabrous beneath, the lowest segments
entire or none, intermediate 7-8 mm. long, trifid or entire, medial
segment 3-5-lobed, lobes all linear-oblong, obtuse, margins involute;
flowers borne medially; calyic 11 mm. long, lobes acute, 7 mm. long,
lanate-stellate both sides; corolla in herb, blue, 2-2.5 cm. long,
petals obovate-oblong, retuse, hirsute at clawed base, nearly free;
stamens in oblong head, free filaments very short; carpels 10, about
4 mm. long, stellate-tomentose, the beaks about 2 mm. long, apically
ciliate. — Pl*obably the Wilkes' U. S. Expl. Exped. specimens from
Casa Canchi above Lima, referred doubtfully to Malvaslrum pin-
natum (Cav.) Baker, belong here as possibly a specimen by Gay
from Cuzco (Hill) ; the Cavanilles' type from Chimborazo, Ecuador,
reportedly had yellow flowers; nevertheless it may prove to be the
same but see Hill's discussions. Petals white, the tips pale lavender
(Grant). Illustrated, Hill, I.e. pi. 29, fig. 11 (leaf). F.M. Neg.
32639.
Lima: Casa Concha, WiUces (det. Gr^y, M. pinnatum?). Casa-
palca, 8S1 (det. Johnston). — ^Junfn: Cerro de Pasco, Matthews 682,
type. Cobracancha Valley, near Cerro de Pasco, 4,200 meters,
Grant 7509. Huar6n, 11S8. La Oroya, Kalenbom 175 (det. Killip,
N. pinnata).
Nototriche Meyeni [Solms] Ulbr. Bot. Jahrb. 42: 120. 1908;
239.
Depressed, pulvinate, the caudex densely branched, the rosulate
leaves congested hemispherically; petioles 6-7 mm. long; stipules
(free part) oval-oblong, 2-3 mm. wide, obtuse and fimbriate, sub-
554 Field Museum of Natural History — Botany, Vol. XIII
glabrous both sides; leaves reniform, stellate- villous both sides,
4 mm. long, 8 mm. wide, parted or trifid, the narrowly cuneate
segments obtuse; flowers about 12 mm. long, below the middle of
the petioles; calyx 6-7 mm. long, its lanceolate-deltoid stellate lobes
2-2.5 mm. long, the yellowish tube glabrous; petals apparently
white marked with blue-black, 7-8 mm. long, 4-5 mm. wide, obtuse,
the tube 6-7 mm. long; anthers many in a globose head; carpels 9,
stellate, including the 2 short beaks (Hill). — Species was confused
with N. borussica (Meyen) Hill and N. sajamensis (Hieron.) Hill,
the latter with filiform stipules and the former with free petioles and
soft white tomentum, and was recognized in fruit by Count Solms
as distinct; only much more material and study will prove the
constancy of these characters. Illustrated, Hill, I.e. pi. 28, fig. 9
(leaf); pi. 30, fig. 12 (carpel). F.M. Neg. 9352.
Puno: Altos de Toledo, 4,700 meters, Meyen, type. — Huancave-
lica: Quispiriza, Raimondi (det. Ulbrich). Without locality, (Lohh).
Nototriche nana Hill, Kew Bull. 19. 1928.
Prostrate caespitose annual, the axillary branches 1-3 cm. long;
petioles 0.5-1.5 cm. long; stipules linear-lanceolate, 2-3 mm. long,
adnate to the petiole below forming a short vagina; leaves herbace-
ous, rosulate, greenish cinereous, minutely and sparsely stellate-
tomentose as the branches, triangular-semicircular, 5 mm. long, 7
mm. wide, 3-parted to base, palmately veined, the rotund-obcuneate
segments trifid or multilobed, the lobes obovate, obtuse or subacute;
flowers sessile near petiole bases; calyx 3 mm. long, lobes acute,
2 mm. long, sparsely or scarcely hirsute; corolla 2.5 mm. long;
stamens in a globose head; carpels 1.75 mm. long, erostrate, minutely
stellate dorsally. — Near N. pusilla Hill but leaves basally 3-parted,
carpels erostrate (Hill) ; an involucel of 3 filiform bractlets is usually
present; petals in corolla tube as little wings (similarly in N. sar-
mentosa, N. pusilla) ; anthers 5; carpels 8 (Krapovickas) . Illustrated,
Hill, I.e. 20; Krapovickas, Bol. Soc. Argent. Bot. 4: 109b (carpel),
110c (leaf).
Tacna: Ancara, 4,300 meters, Volcan Tacora, Werdermann 1121
(type, N. nana)', also 1122 (det. Ulbrich). Chile.
Nototriche nigrescens Hill, Bot. Jahrb. 37: 580. 1906; 223.
Caespitose, often pulvinate; stipules adnate to middle of petioles,
these often 2-2.5 cm. long, the membranous narrowly free part
about 8 mm. long, more or less stellate- tomentose; leaves var5ang ;
Flora of Peru 565
from ovate to semicircular, flabellately 3-5-parted, about 8 mm.
long, above densely, below in age more sparsely stellate, the seg-
ments 3-5-lobed, the medial the larger, the crenate obovate lobules
obtuse; flowers subsessile; calyx lobes narrowly oblong, obtuse,
about 12 mm. long, densely stellate tomentose without; corolla
blue-lilac or violet, 2 cm. long, the broadly obovate petals retuse,
basally 3 mm. connate; carpels (not mature) 3-5 mm. long, shortly
birostrate, dorsally densely stellate, the beaks ciliate. — The numer-
ous stamens are in a globose head, the filaments rather long, the
anthers dark purple-green; suggest A^. flabellata in lobation, the
leaves only sparsely stellate below (Hill), but as to type and 7881
this refers only to old leaves; indument especially of vagina is stellate-
hirsute. F.M. Neg. 9354.
Junln: Hacienda Arapa near Yauli, 4,400 meters, Weherbauer
S81, type; 221. — Cuzco: Above Marcapata, 4,800 meters, Weher-
bauer 7881 (det. Ulbrich). Paucartambo Valley, Herrera 1052 (det.
Killip). "Huikkuna-thurpa" (Herrera).
Nototriche obcuneata (Bak. f.) Hill, Bot. Jahrb. 37: 579. 1906;
225. Malvastrum obcuneatum Bak. f. Joum. Bot. 29: 363. 1891.
M. lobulatum Wedd. Bull. Soc. Bot. Fr. 12: 82. 1865?
Depressed, pulvinate; petioles 1-2 cm. long; stipules adnate
above the middle of the petiole, the membranous free part linear,
acuminate, about 7 mm. long, glabrous except the stellate margins;
leaf-blades semicircular or reniform, flabellately 9-13-parted, 4-6
(8) mm. long, 8-9 mm. wide, densely stellate- tomentose above,
nearly glabrous beneath, the segments 3-7-lobed, the obovate or
spathulate inflexed lobes unequally 5-crenate or -incised, about 11
mm. long; flowers at about middle of petioles; calyx 6 mm. long, the
teeth tomentose; corolla apparently violet, to 2 cm. long, the
obovate or obcuneate petals retuse, coalescent into a tube 5 mm.
long; carpels 8-10, 7 mm. long, the beaks about 3 mm. long, dorsally
stellate-hirsute. — Flowers blue (Baker) ; at first rose, becoming white
(Weberbauer). As original description of Baker gives petals as only
8 mm. long the Bolivian plant may be distinct from the Peruvian
collections referred here, former correctly N. lobulata (Wedd.)
Macbr., comb. nov. Illustrated, Hill, I.e., pi. 28, fig. 2 (leaf). F.M.
Neg. 23750.
Lima: Casapalca, 8SS (det. Ulbrich). — Puno: Poto, Prov. Sandia,
4,500 meters, Weberbauer 983. Ananca, 4,700 meters, Weberbauer
102S.—Tacnsi: Volcan Tacora, Werdermann IJtOO (det. Ulbrich).
Northern Bolivia.
556 Field Museum of Natural History — Botany, Vol. XIII
Nototriche obtusa Hill, Bot. Jahrb. 37: 583. 1906; 220.
Caespitose, often pulvinate; petioles, these 7-8 mm. long, and
stipules, these medially adnate, the vagina membranous, the her-
baceous linear obtuse free part 3-4 mm. long, stellate-pilose only
dorsally and marginally, the trichomes not at all interlocked; leaves
reniform, flabellately or palmately dissected, 4-5 mm. long, 9-10
mm. wide, above densely, below laxly stellate- tomentose, the seg-
ments 3-lobed or entire, obovate-oblong; flowers below middle of
petioles; calyx lobes narrowly oblong, obtuse, 1 cm. long, stellate-
tomentose without; corolla violet, 1.5-2 cm. long, the obliquely
obovate or basally cuneate petals coalescent into a tube 6 mm. long;
stamens globosely capitate. — This species shows fairly close re-
semblance to N. artemisioides (Hill), and also to N. famatinensis.
Illustrated, Hill, I.e. pi. 30, figs. 2, U (calyx with nectar glands).
F.M. Neg. 9355.
Ancash: Above Piscapaccha, 4,500 meters, Weberhauer 2897,
type; 224.
Nototriche Orbignyana (Wedd.) Hill, Bot. Jahrb. 37: 579.
1906; 237. Mcdvastrum Orbignyanum Wedd. Chlor. And. 2: 279.
1857. M. Copelandii Hieron. Bot. Jahrb. 20: Beibl. 49: 43. 1895,
fide Hill.
Pulvinate; leaves densely congested, lanate stellate above,
sparsely tomentose beneath; petioles 12-15 mm. long; stipules
medially adnate, free part filiform, 10-12 mm. long, sparsely stellate-
tomentose as vagina; leaf blades flabellate, unequally trifid, 6-8 mm.
long, 12-14 mm. wide, lateral segments 4-parted, the lacinulae
sometimes 3-4-lobed, smaller medial segment 3-5-lobed, the lacinu-
lae linear, obtuse, 2-4 mm. long; calyx 8 mm. long, tube sparsely
stellate, lobes about 5 mm. long, acute, glabrous within; corolla
10-12 mm. long, purple or deep blue, the retuse broadly obovate
petals coalescent into a tube less than 2 mm. long; stamens in
a globose head; carpels sericeous-pilose, 2.5 mm. long, beaks about
1 mm. long (not quite mature). — In N. sajamensis the leaves are less
deeply divided and the corolla tube is 5 mm. long (Hill) . F.M. Negs.
35506; 9356 (M. Copelandii).
Arequipa: Vincocaya, 4,376 meters, (Copeland, type, M. Cope-
landii). Bolivia.
Nototriche Pearcei (Bak. f.) Hill, Bot. Jahrb. 37: 579. 1906;
255. Malvastrum Pearcei Bak. f. Joum. Bot. 29: 364. 1891.
I
Flora op Peru 657
«
Depressed, the caudex branched, the rosulate leaves with petioles
to 7 cm. long, the stipules adnate to above the middle, the free part
herbaceous, linear, acute, 8 mm. long, puberulent-stellate above,
glabrous beneath, the vagina and petioles more or less stellate-
pilose only marginally; leaves oblong-lanceolate, bipinnate, 1-2 cm.
long, sparsely stellate-tomentose above, glabrous below, the seg-
ments entire or trifid, apically ciliate; flowers shortly pedicellate at
the middle of the petioles; calyx 12 mm. long, the triangular, acute
lobes 4-5 mm. long, without sparsely stellate, within densely tomen-
tose; corolla rose-colored, 2-2.5 cm. long, the obovate retuse petals
1 cm. wide, coalescent into a tube 2 mm. long; stamens in an oblong
head, the free filaments elongate; carpels with conspicuous beaks,
dorsally long-ciliate.
Apurimac: Andsihuaylas, (Pearce, type). — Ayacucho: Huanta,
(Pearce).
Nototriche pedatiloba Hill, Trans. Linn. Soc. ser. 2. Bot. 7:
240. 1909.
Depressed, caespitose, forming small cushions with ligneous,
branched caudex; leaves more or less imbricate, grayish-green;
petioles about 8 mm. long; vagina membranous, 6 mm. wide, the
free part 2 mm. long, 3 mm. wide, truncate, dorsally glabrous,
marginally stellate-fimbriate; leaf -blades semicircular or reniform,
flabellately about 11-parted, 3 mm. long, 7 mm. wide, ashy stellate-
pubescent above, glabrescent below, the segments deeply 4-5-lobed,
the 50 or more lobules fleshy, linear-obovate, obtuse, 1.5 mm. long;
flowers from near the base, shortly peduncled; calyx campanulate,
somewhat inflated, reddish purple, 1 cm. long, the obtuse pubescent
lobes 3 mm. long, the tube glabrous; corolla violet, 16-18 mm. long,
the retuse petals connate into a tube 5 mm. long; carpels 6-8, 5-6
mm. long, rostrate, stellate-tomentose, the beak stellate-ciliate,
about 2 mm. long. — The deeply and multilobed leaves point to
a relationship with N. sajamensis and N. Meyeni; the erect fleshy
lobulae, which occur in little groups of 4 or 5, are really the lobes
of the leaf divided almost to their bases, each leaf thus consisting of
about 50 lobulae borne on a small pedate expansion; the plant is
unique by its broadly truncate, membranous stipules and the
finally inflated calyx (Hill). The var. appendiciUata Burtt, Kew
Bull. 132. 1948 is marked by 2 appendages arising from the surface
of the vagina, either linear, membranous, 3-4 mm. long, margins
pilose or reduced to glabrous teeth 0.5 mm. long. The author (I.e.)
discusses the possible character of these appendages, suggesting that
558 Field Museum of Natural History— Botany, Vol. XIII
the phenomenon of diplophylly may be concerned. Illustrated,
Hill, I.e. pi. 27, fig. ^ (plant) ; pi. 28, fig. 12 (leaf) ; pi. 30, fig. 5 (calyx) ;
pL 30, figs. 10, 20 (fruits); Burtt, I.e. (var. appendiculata) .
Arequipa: Vincocaya, 4,376 meters, carpeting the ground with
N. argylloides, near summit of Arequipa-Puna railway (Hill 78,
type). — Puno: Minas de San Antonio, Sandeman 3920 A; 3931 (var.).
San Antonio de Esquilache (Stafford 705 A, type, var.).
Nototriche pedicularifolia (Meyen) Hill, Bot. Jahrb. 37: 579.
1906; 246. Sida pedicularifolia Meyen ex Walp. Nov. Act. Acad.
Leop.-Carol. 19: Suppl. 1: 308. 1843. N. incana Turcz. Bull. Soc.
Nat. Mosc. 36, pt. 1: 567. 1863, fide Hill.
Habit of the related N. argylloides, but all vegetative parts,
except glabrate stipules, densely ashy stellate- tomentose; petioles
2-3 cm. long; stipules adnate medially, free part linear, acuminate,
1 cm. long; leaves cordate or pentagonous, 2 cm. long, 1.5-2 cm.
wide, palmately 3-7-parted, the segments subpinnatifid, incised
crenate, lacinulae crisped, obtuse, the largest medial segment bi-
pinnatifid the lateral smaller, simpler; flowers borne medially; calyx
8 mm. long, the ovate lobes subacute; corolla violet, 15 mm. long;
petals linear- or cuneate-oblong, 3 mm. wide, retuse, scarcely coa-
lescent; stamens in oblong head; carpels about 12, rounded, 2 mm.
long, dorsally pilose with long silky pure white trichomes. — M.
pulverulenta Burtt & Hill, Kew Bull. 135. 1948, of Sajama, Bolivia,
may occur in Peru. It is of interest as the anther column is 3 mm.
long, 2.5 mm. wide, and in this character approaches species with
globosely borne anthers, the petals coalescent 5 mm., pilose at
base, broadly obovate, carpels beaked. Illustrated, Hill, I.e. pi. 27,
fig. 5 (plant); pi. 30, fig. 16 (carpel). F.M. Neg. 9357.
Arequipa: Vincocaya (Copeland). — Puno: Near Pisacoma, Meyen,
type. Ananca, 4,700 meters, Weberbauer 1023, part. — Tacna:
Tacora, Weddell; Meyen. Bolivia; Chile.
Nototriche pellicea Hill, Kew Bull. 133. 1948.
Depressed, caespitose, pulvinate, hirsute; petioles 4 mm. long;
vagina 2.5-3 mm. broad; stipules adnate about medially, the free
part linear-lanceolate, acute, 4 mm. long, 1 mm. wide, only dorsally
densely stellate-tomentose as the oblong-orbiculate leaves, these
4.5 mm. long, 6 mm. wide, deeply 5- or 7-parted, the segments ■
digitately lobulate with 30-50 narrowly linear-oblanceolate lobules;
flowers from near base of petioles; calyx cylindric-campanulate,
Flora op Peru 559
7-7.5 mm. long, tube glabrous, lobes stellate-hirsute both sides, to
3.5 mm. long, acute; corolla white, 1 cm. long, the petals with
purple lines without, the hirsute tube 3 mm. long; carpels 3.75 mm.
long, beaks 1.25 mm. long, stellate-setose. — Like N. turritella Hill
but with narrow longer leaf lobes, stipules larger, trichomes longer,
flowers white-purple, carpel beaks larger (Hill). Illustrated, Hill,
I.e. 134.
Type found only on barren exposed places above copper veins,
and ash from plants tested quantitatively contained 0.77 per cent
copper.
Puno: Santa Lucia, 4,570 meters, (Sharpe 168, type). "Turp."
Nototriche phyllanthos (Cav.) Hill, Trans. Linn. Soc. ser. 2.
Bot. 7: 232. 1909. Sida phyllanthos Cav. Diss. 5: 276, pL 127. 1788.
Sida saxifraga Humb. & Bonpl. PI. Aequin. 2: 116. 1813, fide Hill.
F*ulvinate, about 3 cm. across; petioles about 8 mm. long, the
stipules adnate to middle or rarely nearly to leaf blade, the resulting
vagina 3.5-4 mm. wide, free part subacute, subglabrous, margins
stellate, 3-4 mm. long; leaves rotund-reniform, 3-parted, 8-10 mm.
long, 10-13 mm. wide, the segments 3-5-parted or -lobed, the ovate-
oblong lacinulae rarely lobulate, velvety pubescent above, vernicose,
glabrous or very sparsely stellulate beneath; flowers about from
middle of petioles; calyx stellate, 1 cm. long; corolla 18-20 mm.
long, violet, the petals scarcely coalescent basally; stamens in a
cylindrical column; carpels about 9 mm. long, stellate-ciliate, the
beaks 4 mm. long. — After Hill who, however, drew the description
from Ecuadorean plants, the type, from Peru (maybe from Guaya-
quil region) unknown to him. Hill describes at length the identity
and relationship. F.M. Neg. 35507 (S. saxifraga).
Peru (probably) : Dombey, type. Ecuador.
Nototriche pichinchensis (Humb. & Bonpl.) Hill, Bot. Jahrb.
37: 579. 1906; 231. Sida pichinchensis Humb. & Bonpl. PI. Aequin.
2: 115. 1813. Malvastrum pichinchense (Humb. & Bonpl.) Gray, Bot.
U. S. Expl. Exped. 152. 1854.
Resembles closely N. phyllanthos, at least as interpreted, but
tjrpically larger, sometimes 6-8 cm. across; petioles 2-4, or in var.
less than 1 cm. long; stipules subulate, the free part 7-10 mm. long;
leaves in type 18 mm. long, 22 mm. wide, divided as in A^. phyl-
lanthos or in var. 1 cm. long, 16 mm. wide, the 3 segments trilobed
and each again trilobed; calyx 16 mm. long or in var. 10-13 mm.
560 Field Museum of Natural History— Botany, Vol. XIII
long; corolla about 3 cm. long, in var. 2 cm. long. — Seems via the
variant (var. angusta Hill of Ecuador) to pass into N. phyllanthos.
However since Ulbrich (Notizbl. Bot. Gart. Berlin 11: 527. 1932) has
retained the species, his identification without more material may be
followed. Illustrated, Humb. & Bonpl., I.e., pi. 116.
Cuzco: Valle del Paucartambo, 3,600 meters, Herrera 1816 (det.
Ulbrich). Ecuador.
Nototriche porphyrantha Ulbr. Bot. Jahrb. 54: Beibl. 117:
62. 1916.
Similar to but apparently well distinct from N. flabellata; vagina
2-3 mm. wide, stipules (free part) subulate-linear, about 1 cm. long,
marginally stellate; leaf blades greenish, rather lightly and finely
floccose-stellate above, subtomentose beneath, 22-25 mm. long,
18-25 mm. wide, trifid, the medial lobe cuneate-obovate, the lateral
obliquely obovate, 2-3-cleft, the lobes all incised-crenate; flowers
shortly pedicellate; calyx 18 mm. long, lobes ovate-lanceolate,
glabrous within; corolla purple, about 3 cm. long; stamen 3.5 cm.
long; anthers in globose-ovoid head; carpels long- villous, the villous
beaks 6-7 mm. long.— F.M. Neg. 9358.
Cuzco: On rocks, Pisac to Paucartambo, 4,000 meters, Weber-
bauer 6917, type.
Nototriche purpurascens Hill, Trans. Linn. Soc. ser. 2. Bot.
7: 249. 1909.
Depressed, the stout branched caudex to 1.5 cm. across, the rosu-
late leaves with petioles 1.5-4 cm. long, stipules adnate nearly to the
middle, the vagina 3 mm. wide, the free part narrowly triangular,
acute, 5 mm. long, glabrous; leaves broadly ovate or pentagonous,
palmately parted, 2 cm. long, 2-2.5 cm. wide, densely stellate-
tomentose above, glabrous beneath, the lowest segments subentire,
7-10 mm. long, the intermediate pinnate with 6-10 lacinulae, medial
segment pinnate with larger and smaller lobes alternating, all with
strongly inflexed margins; flowers borne about medially; calyx 2 cm.
long, the lobes broadly ovate, acute, 1 cm. long, glabrous without,
tomentose within, purplish; corolla copper-colored, 3-3.5 cm. long,
the obovate petals truncate or retuse, their tube 3-4 mm. long;
stamens in a globular head, subsessile, few, dark purple; carpels
densely pilose dorsally, shortly birostrate, about 4 mm. long (scarcely
mature). — The corolla was described as 22 mm. long, but probably
Flora of Peru 661
not fully grown, as the Herrera plant has some similarly unde-
veloped flowers. F.M. Neg. 23781.
Cuzco: Valle del Paucartambo, 3,700 meters, Herrera lOSU.
Bolivia.
Nototriche pusilla Hill, Bot. Jahrb. 37: 587. 1906; 218.
Caespitose-pulvinate annual, the leaves in a terminal rosette and
with short spreading horizontal branches terminating in rosettes of
leaves and flowers; f)etioles 5 mm. long; stipules of leaves without
flowers little adnate, of those with flowers adnate medially forming
membranous vagina, the filiform acute free part 2-3 mm. long, laxly
stellate-tomentose dorsally and marginally; leaves more or less
cordate or orbicular, crenate or somewhat dentate, 5 mm. long,
6 mm. wide, loosely stellate-tomentose both sides; flowers medial,
sessile; calyx lobes 4-4.5 mm. long, sparsely tomentose both sides;
corolla white, 3-4 mm. long; carpels 1.5 mm. long, minutely biro-
strate, dorsally stellate-hirsute. — The dense tomentum is somewhat
shaggy (Hill); flowers have no involucel; petals inserted as in N.
nana and N. sarmentosa; anthers 5; carpels dehiscent, about 10
(Krapovickas, Bol. Soc. Argent. Bot. 4: 111. 1951). Illustrated,
Hill, I.e., pi. 29, figs. 20, 21 (plants); pi. 30, fig. 29 (carpel). F.M.
Negs. 9360; 35508.
Ancash: Near Chonta, 4,400 meters, Weberbauer 2788; 224.
Without locality, Weddell, type (Herb. Paris). Bolivia; Argentina.
Nototriche pygmaea (Remy) Hill, Trans. Linn. Soc. ser. 2. Bot.
7: 218. 1909. Sida pygmaea Remy, Ann. Sci. Nat. s^r. 3, 8: 238.
1847. Malva pygmaea (Remy) Wedd. Chlor. And. 2: 274. 1857.
Malvasirum pygmaeum (Remy) Gray, Bot. U. S. Expl. Exped. 150.
1854.
Small annual about 4 cm. long, the caudex shorter than 1 cm.;
petioles 4 mm. long; stipules medially adnate, the subulate free part
3 mm. long, dorsally stellate tomentose; leaves reniform or sub-
flabelliform, about 2 mm. long, 3 mm. broad, incised-crenate or
lobate, the lobes 8-10, densely stellate-tomentose both sides; flowers
borne medially, pedicellate; calyx about 4 mm. long, triangular
lobes densely tomentose; flowers white or with dark center, 3 mm.
long, obovate petals with 1 mm. long tube; carpels 7, minutely
stellate at rounded tip, 1 mm. long. — Type by D'Orbigny near the
Laguna de Potosi, Bolivia, due east of Tacna (Hill). Illustrated,
562 Field Museum of Natural History— Botany, Vol. XIII ]
Hill, I.e., pi 29, figs. 22, 2^ (plant and flower) ; pi. 30, fig. 30 (carpel). \
F.M. Negs. 35509. ■,
Moquehua: East of Carumas, 4,500 meters, Weberbauer 7359 i
(det. Johnston). Bolivia. ]
Nototriche rugosa (Phil.) Hill, Bot. Jahrb. 37: 579. 1906; 245. |
Malvastrum pedicularifolium (Meyen) Gray, var. rugosum Phil, ex 1
Reiche, Fl. Chile 1: 235. 1896. |
Depressed, often pulvinate, the rosulate leaves ashy pulverulent- ij
stellate except medially beneath, the petioles 2-5 cm. long, the t
stipules adnate about medially, the free part membranous, subulate, |
to 5 mm. long, as petioles glabrous except ciliate marginally; leaves |
palmately 5-parted or digitately lobed, about 1 cm. long and broad, >
the pinnatifid lobes incised crenate, crisped; flowers at about the ;|
middle of the petioles; calyx 6-7 mm. long, tube glabrous, lobes ,
ovate, obtuse, 2-2.5 mm. long, pulverulent; corolla white or blue, f
7-8 mm. long; petals obovate, the tube 3 mm. long; stamens few in
globose head.— F.M. Neg. 9362. •
Tacna: Volcan Tacora, 4,400 meters, Werdermann 1398 (det. ■
Ulbrich). Northern Chile. "P
Nototriche sajatnensis (Hieron.) Hill, Bot. Jahrb. 37: 579. f
1906; 243. Malvastrum sajamense Hieron. Bot. Jahrb. 21: 319. |
1895. I
Depressed, pulvinate; leaves rosulate and more or less imbricate '^
at the tips of the branches, more or less densely stellate-tomentose |
or beneath subglabrous; petioles 6-8 mm. long, the stipules adnate ^
to above the middle forming a vagina 4 mm. wide, the free part |
linear-filiform, about 5 mm. long, 1 mm. wide, the entire organ |
glabrous or margins stellate-ciliate; leaf blades reniform, flabel- f;
lately 9-11-parted, 4-5 mm. long, 10-12 mm. wide, the segments f
deeply trifid, the laciniae deeply bifid or trifid with obtuse lacinulae, ;i
1.5-2 mm. long; flowers basal, pedicellate; calyx 8-9 mm. long, the f
lobes obtusish, 3 mm. long, stellate-hirsute, the tube subglabrate; |
corolla yellowish-white (drying dark purple), 10-14 mm. long, ovate |
petals acutish, tube about 5 mm. long; carpels (immature) stellate- 1
ciliate, the beaks 1.5 mm. long. — Appears to be allied with N. [>
Orbignyana and N. alternata (Hill). But compare N. condensata. |
Illustrated, Solms, Bot, Zeit. 65: 120. pi. 2, figs. 1, 3, 14; Hill, I.e. |
pi. 28, fig. U (leaf). F.M. Negs. 9364; 9365. |
Tacna: Volcan Tacora, Stilbel (Steinmann). Bolivia.
Flora of Peru 563
Nototrlche salina Burtt & Hill, Kew Bull. 135. 1948.
Caudex often branched toward apex, bearing rosettes 2-4 cm. in
diameter; free part of stipules membranous, 4-5 mm. long, 1.5-2.5
mm. wide, triangular-lanceolate, acute, adnate for 5 mm., forming
glabrous vagina 4.5-5 mm. broad; leaves ovate-cordate, with 5
primary lobes disposed palmately, pinnately lobulate with obtuse
segments, sparsely stellate only above if at all; calyx 5 mm. long,
lobes 1.5 mm. long, glabrous or nearly so without, pilose within
marginally and toward the apex, basal nectaries semicircular;
corolla dark purple, tube glabrous, 1.5-2 mm. long, petals obovate,
emarginate, about 5 mm. long, glabrous; anthers in globose head;
carpels 4-4.5 mm. long, beaks 1.5 mm. long, dorsally long-sericeous.
— Resembles N. aristcUa Hill, N. longirostris (Wedd.) Hill and N.
rugosa (Phil.) Hill of Chile; it differs from the first two by the
glabrous or subglabrous leaves and from the last also by the ros-
trate carpels (Hill). Illustrated, Hill, I.e. 137.
Arequipa: In turf, sand and volcanic ash. Borax Lake, Salinas
Pichu Pichu, (Stafford 1325, type).
Nototriche sarmentosa Hill, Kew Bull. 20. 1928. N. Weder-
mannii Ulbr. Notizbl. Bot. Gart. Berlin 11 : 529. 1932.
Prostrate annual sarmentose herb, the axillary branches 4-7
cm. long, sparsely and laxly stellate as the petioles, these 8-12 mm.
long, and the linear-lanceolate half adnate stipules; leaves rosulate,
semicircular, herbaceous, medially (more or less) trilobed, 4-5 mm.
long, (4) 7-13 mm. wide, minutely velvety stellate, medial lobe
sub- trilobu late, the lobules themselves 4-7-lobulate, the ultimate
crenate; flowers sessile in the middle of the petioles; calyx 5 mm.
long, medially 5-lobed, the lobes acute; corolla about 2.5 mm. long,
tube 0.5 mm. long; stamens few; carpels 1.75 mm. long, apically
stellulate. — Allied to N. nana Hill but coarser with larger leaves
medially trifid, and carpels minutely rostrate (Hill), adhering to
seeds; flowers often with an involucel of 2 filiform bractlets; corolla
tube persists as a disk (Krapovickas, Bol. Soc. Argent. Bot. 4: 108.
1951). Illustrated, Hill, I.e. F.M. Neg. 23751.
Tacna: Ancara, Volcan Tatora, 4,300 meters, Werdermann 1123,
type (also type of N. Werdermannii). Argentina.
Nototriche sericea Hill, Trans. Linn. Soc. ser. 2. Bot. 7: 258.
1909.
Depressed, the caudex branched, the rosulate leaves with petioles
to 8 cm. long, the stipules adnate about to the middle and with
664 Field Museum of Natural History— Botany, Vol. XIII
vagina 2 mm. wide, the membranous subulate filiform free part
1 cm. long, densely as vagina above and marginally sericeous ciliate,
glabrous dorsally; leaves triangular-ovate, palmately parted, the
segments pinnate or bipinnate, above more or less densely, beneath
very sparsely sericeous, the lobes entire or 3-5-cleft, the numerous
ultimate lacinulae narrowly linear, obtuse; flowers borne medially,
pedicellate; calyx 8-9 mm. long, lobes acute or acuminate, 3-4 mm.
long, without and marginally sparsely ciliate, the tube glabrous;
corolla blue- violet, 14-17 mm. long, the obovate-cuneate retuse
petals coalescent into a tube 4-5 mm. long; stamens borne globosely,
the free filaments long; carpels about 13, 4 mm. long, densely
sericeous, the beaks 2 mm. long. — Distinguished by its silky gray
appearance and the many linear segments of the leaves, and the
minute anthers; trichomes are sessile with 2-4 long arms (Hill).
Illustrated, Hill, pi 29, figs. 9, 10 (leaf, flower).
Puno: Crucero Alto, 4,470 meters (Hill 82, type).
Nototriche Staffordiae Burtt & Hill, Kew Bull. 136. 1948.
Globosely rosulate plants 2.5-4 cm. in diameter and unique by
the character of the trifid calyx-lobes; caudex mostly simple; free
part of stipules linear, acute, 7 mm. long, adnate with vagina 5 mm.
long, free part of petiole 3 mm. long, stellate-pilose (as stipules);
leaves 4-5 mm. long, 10 mm. wide, 9-dissected more than medially,
these divisions 5-lobed, the lobules linear; calyx campanulate, stel-
late-pilose, 8 mm. long, the lobes trifid more than medially, the
linear divisions 2.5 mm. long; corolla tube 1 mm. long, glabrous;
petals obovate, 6.5 mm. long, 4.5 mm. wide, glandular-pubescent
medially and ciliate toward base, strongly veined, greenish striped
with brown; anthers in a globose head; carpels (immature) densely
stellate-pilose, shortly and obtusely rostrate. — Corolla tube adheres
subpersistently to the calyx; the corolla disarticulates above the
base leaving a rim apparent as a flap if calyx is dissected before
anthesis. This interesting species commemorates fittingly the intel-
ligent collecting of Miss P. Stafford. Illustrated, Burtt & Hill,
I.e. 137.
Puno: Sandy slopes just below moraine, 4,570 meters, San
Antonio de Esquilache (Stafford 1263).
Nototriche stenopetala (Gray) Hill, Bot. Jahrb. 37: 579. 1906;
251. Mcdvastrum stenopetcUum Gray, Bot. U. S. Expl. Exped. 155.
1854.
I
Flora of Peru 565
Depressed, pulvinate; petioles 12 mm. long, the stipules adnate
nearly to the middle, the vagina 3 mm. wide, membranous and gla-
brous as the subulate free part, this 3 mm. long; leaves triangular
or cordate, 10 mm. long, 12-13 mm. wide, white stellate velvety
above, glabrous beneath, trifid, the more ample middle lobe pinnate,
segments trifid or entire, the lobes 3-5 mm. long, linear-oblong,
little inflexed, often ciliate; flowers from below the middle of the
j)etioles; calyx 9 mm. long, lobes narrowly triangular, subacuminate,
sparsely stellate without, densely white tomentose within; corolla
scarlet, 17 mm. long, the narrowly spathulate petals entirely or
essentially free; stamens few in cylindrical head; carpels about 12,
hirsute. — My si)ecimens from wet grassy pond-slope, petals scarlet
above, white below, red-veined. Illustrated, Hill, I.e. pi. 29, fig. 17
(leaf).
Lima: Casa Cancha, Pickering, type. Without locality {Mac-
lean). — Junin: La Oroya, Kaknborn 116 (det. Killip). Hacienda
Arapa near Yauli, Weherhauer SJ^. Cerro de Pasco, S061 .
Nototriche sulphurea Hill, Bot. Jahrb. 37: 583. 1906; 248.
Robust, caespitose, the caudex to 2.5 cm. in diameter; leaves
laxly rosulate, densely stellate-tomentose with long sulphur-colored
trichomes, this indument also covering the petioles (4 cm.), medially
adnate filiform acute stipules (12-15 mm.) and ovate acute calyx
lobes (11 mm.) without; leaves broadly ovate-cordate or penta-
gonous, palmately parted or pinnatifid, 2-2.5 cm. long and nearly
as wide, the oblong segments pinnatifid, the obovate laciniae entire
or incised-crenate; flowers medial, sessile; corolla pale blue, 2-3.5
cm. long, the obcuneate petals retuse, tomentose at base, scarcely
coalescent into short tube; carpels 15-20, 5 mm. long, beaks 1 mm.
long, dorsally sericeous. — Stamens in a more or less oblong head
(Hill). The dense villous pubescence and much divided leaves are
striking characters. Illustrated, Hill, I.e. pi. 29, figs. 8, U (leaves).
F.M. Neg. 9366.
Cuzco: Puna above Marcapata, Weherhauer 7882; Herrera 296
(both det. Ulbrich). Prov. Canas, Herrera 932 (det. Standley,
A^. porphyrantha) . — Puno: Poto to Ananca, Prov. Sandia, 4,700
meters, Weherhauer 968, type; 219. Andes of Pebechucho {Pearce).
"Thurpa" (Herrera).
Nototriche ticsanica Ulbrich, Notizbl. Bot. Gart. Berlin 11:
531. 1932.
566 Field Museum of Natural History— Botany, Vol. XIII
Pulvinate, the densely rosulate leaves or branches 2 cm. long;
petioles 12-15 mm. long, the stipules adnate to above the middle,
free part 5 mm. long, triangular lanceolate, only marginally stellate-
villous; leaves obovate-orbicular, 7-8 mm. long, 8-10 mm. wide,
flabellately multiparted, the lobes obtuse, above densely, beneath
glabrate or early sparsely tomentose; flowers purple, sessile; calyx
about 8 mm. long, sparsely stellate, the lobes nearly 3 mm. long,
densely villous; petals about 13 mm. long, oblong-obovate; stamen
tube cylindric, 1 cm. long, the filaments 1.5-2 mm. long, in a globose
head. — Petals white within, purple without except below (Weber-
bauer). Allied to N. coccinea with smaller leaves divided less,
scarlet somewhat larger flowers (Ulbrich). However, compare
N. condensata and note.
Moquehua: Carumas near Volcano Ticsani, 4,400 meters, Weber-
bauer 7320, type.
Nototriche turritella Hill, Trans. Linn. Soc. ser. 2. Bot. 7;
241. 1909.
Caespitose, the ultimate branches densely clothed with leaf-
remains and verticillate floriferous and sterile leaves, erect and
turret-like, 2-6 cm. high, the pale violet flowers with deep purple
centers sessile in the crowded leaf -rosettes at their tips; petioles
4-6 mm. long; stipules nearly completely adnate, forming vagina
3 mm. wide, the linear-oblong free part obtuse, 3 mm. long, the
entire structure glabrous except the stellate- tomentose margins;
leaves broadly cuneate or cuneate-reniform, obscurely trifid, 4-5
mm. long, 8 mm. wide, early densely stellate both sides, small
medial segment 9-11-parted, the lateral segments 3-5-lobed, the
lobes multilobulate, these ultimate lacinulae in 4's or 5's, the tips
soon glabrous, their upper margins inflexed; calyx tubular cam-
panulate, 9 mm. long, lobes 3.5 mm. long, densely stellate except
the obtuse fleshy tips; corolla 13-15 mm. long, more or less rotate,
petals broadly obovate, scarcely retuse, with tube 5 mm. long;
carpels about 8, usually only 1 maturing, 4 mm. long, stellate-pilose,
beaks about 1 mm. long. — In black volcanic ash slopes of El Misti
with a vertical range of about 1,000 meters, conspicuous and beauti-
ful when in full flower around the middle of March (Hill). Illus-
trated, Hill, I.e. 202 (habit photograph); pi. 27, figs. 1, 2 (plants);
pi. 28, figs. 11 (leaf and flower), 16 (leaf -blade) ; pi. 30, figs. 6 (open
flower), 13 (carpel).
Arequipa: Volcano El Misti, 4,000-5,000 meters, (Hill, type);
Solon I. Bailey. Alto de los Huesos, 3,700 meters, Weberbauer 1U21.
Flora of Peru 567
— Tacna: Volcan Tacora, 4,500 meters, Werdermann IJ^U (det.
Ulbrich).
Nototriche ulophylla (Gray) Hill, Bot. Jahrb. 37: 579. 1906;
224. Malvastrum ulophyllum Gray, Bot. U. S. Expl. Exped. 1: 150.
1854.
Similar and closely allied to A^. obcuneata; petioles to 12 mm.
long; stipules adnate to about the middle of the petiole, the free
part broadly linear subacute, about 6 mm. long; leaves reniform-
rotund, 7-11-parted flabellately to the middle, 5-6 mm. long, 8-10
mm. wide, the 3-7-lobed segments extremely crowded, the lobulae
obovate, involute and crisped; calyx 6 mm. long, the obtuse lobes
very sparsely stellate; corolla 6-7 mm. long; carpels 8, about 1.5
mm. long, but immature, minutely rostrate, densely sericeous-
hirsute dorsally, the trichomes 3-4 mm. long. — Differs from the
related species in having longer lateral or secondary lobulae, which
are often again lobed, and the entire leaf has a very crisped appear-
ance; it also differs in the smaller flowers (Hill). Illustrated, Hill,
I.e., pi. 28, fig. 1 (leaf).
Lima: Alpamarca, near the snow line, Pickering, type. Casa-
palca, 8S3 (det. Johnston). — Huancavelica: Pisco to Ayacucho,
4,800 meters, Weherhauer SUUS.
Nototriche Vargasii Krap. Bol. Soc. Argent. Bot. 5: 73. 1953.
Depressed perennial, the ligneous subterranean caudex about
a cm. in diameter; stipules medially connate forming a glabrous
vagina, to 2 cm. long, 2.5-3 mm. wide, the linear-lanceolate free
portion to 1 cm. long, 2 mm. wide, sparsely ciliate apically; leaves
unequally pinnatifid, about 5 cm. long, 4-5 cm. wide, the smaller
lanceolate lower divisions opposite, entire, the narrower longest
medial pair more or less pinnately lobed, 2-2.5 cm. long, the remain-
ing upper divisions much reduced and subentire, all stellulate-
tomentulose above, glabrous beneath, some lobes terminally ciliate;
flowers solitary; caljrx campanulate, about 2 cm. long, glabrous
except slightly stellate in the 5 nerves above and puberulent on the
edges of the broadly triangular (7 mm. long) teeth; nectaries about
4 mm. long and broad, connate at base; corolla red, about 3.5 cm.
long, the tube 7-8 mm. long, 6 mm. in diameter at base, the obovate
petals obliquely truncate, about 12 mm. wide, pilose at base between
the plaits and the upper part of the tube; anthers in a globose head;
stigmas capitate, 17; carpels (immature) rigidly stellate and aristate.
— Distinguished from all related species by the large red flowers and
568 Field Museum of Natural History — Botany, Vol. XIII
deeply pinnatifid leaves (Krapovickas). Illustrated, I.e. jig. 7 (leaf,
flower).
Cuzco: Hacienda Poqquera, Prov. Espinar, 4,500 meters, (Vargas
10582, type).
18. GAYA HBK.
Like Sida except that the membranous inflated carpels converge
apically, dehiscing dorsally in 2 valves and developing within an
incurved erect appendage (endoglossum) originating near base of
dorsal wall, later separating, often apparently attached to ventral
wall and more or less enclosing seed; 2-3 ligaments hold each carpel
to columella until maturity. — Kearney, Amer. Midi. Nat. 46: 123,
quotes Hochreutiner's observations that explain the development of
the endoglossum and the ligaments, previously misinterpreted.
Students are referred to the full account of this and other similarly
fascinating studies by Hochreutiner, Ann. Cons. Jard. Bot. Geneve
21: 347-387. 1920. The several Peruvian forms are not clearly
distinct and a revision of the genus is very much needed.
Leaves mostly 7-10 cm. long, glabrate above in age.
Flowers in 2's or 3's in the upper axils G. triflora.
Flowers all solitary G. calyptrata.
Leaves mostly or all 3-5 cm. long, densely tomentose even in age on
both sides G. peruviana, G. jaenensis, G. Weherbaueri.
Gaya calyptrata (Cav.) HBK. ex Schum. in Mart. Fl. Bras. 12,
pt. 3: 351. 1891. Sida calyptrata Cav. Diss. 2: 57. 1786. S. nutans
L'H^r. Stirp. Nov. 5: 119 bis, pL 57 bis. 1789. S. disticha Cav. Icon.
5: 12, pi lt22. 1799.
Slender-stemmed shrub, the branching stems glabrous below,
minutely stellate-tomentulose toward the tips as the membranous
cordate-ovate acuminate coarsely serrate leaves of these glabrate in
age above, commonly 7-10 cm. long, about 6 or 7 cm. wide at base;
peduncles typically capillary, solitary, often reflexing, 2 or more
cm. long, 5 cm. or longer in fruit; calyx lobes ovate, acuminate, about
5 mm. long; petals 10-12 mm. long; fruits depressed, puberulent,
about 1 cm. across the slightly reticulate obtuse carpels with parch-
ment-like walls. — Quite possibly should be drawn to include G.
Weherbaueri Ulbr. (which see) but the original specimen is the large-
leaved plant with light pubescence illustrated by L'H^ritier, I.e.
F.M. Negs. 29777; 29781 (5. disticha).
Flora of Peru 569
Lima: Sands near Lima, Domhey, type. Without locality, Ruiz
& Pavdn (type, S. wutans). — Cuzco: Chacra de Caiia, Prov. Anta,
Vargas 145. To Mexico?
Gaya jaenensis Ulbr. Notizbl. Bot. Gart. Berlin 11: 539. 1932.
Few-branched cinereous-velutinous pubescent shrub, the close
indument yellowish in the leaves and calyces; stipules 3 mm. long,
promptly caducous; petioles 2-5 mm. long; leaves oblong-lanceolate,
2-3 cm. long, 3-15 mm. wide, serrate, nerves prominent only be-
neath; peduncles 1.5-2 cm. long, articulate 5-6 mm. below the calyx,
this patelliform, nearly medially 5-lobed, the lobes about 5 mm.
long, 3 mm. wide at base, 1-nerved; corolla orange, 2 mm. long,
glabrous, as the basally dilated half as long stamen- tube; styles 10;
fruit depressed-globose, 8-10 mm. broad, the 10-12 1-seeded carpels
tomentose, 5 mm. high, obtuse, the pergamentaceous endocarp
hamulose-rugulose, the seed pubescent with appressed and spreading
trichomes. — Type 1 meter high. Resembles and allied to G. her-
mannioides HBK. of Mexico but leaves and flowers different in size;
the fruit however is similar; the seeds are disclosed by the dorsal
dehiscence but are retained by the parchment-textured wing-like
walls in the hamulately roughened dark endocarp, which probably
serves as an aid to germination by water-absorption (Ulbrich).
Cajamarca: Between Shumba and Ja^n, Weberbauer 6189, t)rpe.
Gaya peruviana Ulbr. Notizbl. Bot. Gart. Berlin 11: 540. 1932.
Branches few, acutely diverging, the younger as the lanceolate-
subulate stipules and petioles sparsely pilose, these both to 3 mm.
long; leaves lanceolate, tnmcate or subcordate at base, very acute
at tip, entire or sometimes indistinctly serrate, 1.5-3 cm. long, 3-10
mm. wide, sparsely stellulate or pilose with simple trichomes above,
tomentose beneath, the nerves there prominent; peduncles 1.5-2
cm. long, articulate, 5-8 mm. below the calyx, this tomentose, cleft
more than medially, the acute lobes 3-4 mm. long, 2 mm. wide at
base; flowers yellow, 8-10 mm. long, glabrous as all parts except
ovary; fruit glabrescent, the 10-12 carpels 4 mm. high, 2-2.5 mm.
wide, obtusish, the triangular seeds pilose only at the angles. —
Tjrpe about 1 meter tall. Related to G. aurea St. Hil. with longer
petioles, larger flowers and fruits (Ulbrich).
Piura: Shrubby grass steppes, 1,500 meters, (Weberbatier 6363,
type).
570 Field Museum of Natural History— Botany, Vol. XIII
Gaya triflora Hochr. Ann. Cons. Jard. Bot. Geneve 20: 141.
1917.
Upper stems herbaceous, velutinous-tomentulose (toward tips
slightly glandular) as the petioles (2-11 cm. long), leaves beneath
and peduncles, these 3, or in fruit 6 cm. long; leaves cordate-ovate,
acuminate, irregularly crenate-serrate, sometimes sub-trilobed, 7-15
cm. long, 4-8 cm. wide, in age above sparsely pubescent but velvety
to touch, palmately many-nerved from base; flowers (2) 3 in each
axil; calyx pubescent without, pilose within, the ovate lobes minutely
acuminate, about 4 mm. long, 3.5 mm. wide at base, little accrescent
in fruit; petals suborbicular, entirely glabrous, about 11 mm. long;
stamen column scarcely 4 mm. long, glabrous but with a setose line
around it at insertion with petals; fruit to 12 mm. across, the sparsely
pilose carpels crescent-appendaged within and with large wings
transversely 7-8-nerved and marginally dentate; seeds pilose except
laterally nearly glabrous. — Remarkable in the 3 flowers in each
axil (sometimes only 2), glabrous petals, line of trichomes encircling
stamen column (author). The type was referred by Baker to G.
subbiloba HBK. of Colombia, the flowers solitary and scarcely to be
expected in Peru. F.M. Negs. 9404 (Dahlem spec); 23691.
Amazonas: Chachapoyas, Mathews. Without locality, Mathews
3236, type.
Gaya Weberbaueri Ulbr. in Notizbl. Bot. Gart. Berlin 11:
541. 1932.
Yellowish velutinous-pubescent even to the calyces; stipules
subulate, 2 mm. long; petioles 1.5-4 cm. long; leaves oblong-oval,
finely serrulate, 2-4.5 cm. long, 2-3 cm. wide, nerves prominent
beneath; peduncles 1.5-3.5 cm. long, articulate 5-10 mm. below the
subcampanulate calyx, this parted nearly medially, the acuminate
lobes 2.5 mm. long, 2 mm. wide at base; corolla orange, campanulate,
about 10 mm. long, the obovate suborbicular petals sparsely fim-
briate; ovary pilose; fruit semi-globose, 8-10 mm. broad, the 12 or
more triangular-oblong carpels 4-5 mm. high, 2-2.5 mm. wide,
dehiscing toward apex, yellowish as the rugulose endocarp, the 2
mm. long seeds minutely pilose only on angles. — Resembles G.
calyptrata (Cav.) HBK. with shorter petioles, larger flowers, smaller
fruits (Ulbrich). Apparently should be drawn to include G. jaenen-
sis and G. peruviana (this not seen), but all three species ex char,
similar to each other and to G. aurea St. Hil. Fl. Bras. Merid. 1:
193. 1827, from which species it may be noteworthy. Ulbrich gives, ■
Flora of Peru 571
for G. peruviana, only comparative diflFerences commonly meaning-
less as indicators of genetic distinction; and G. aurea St, Hil. seems
to have little to distinguish it from G. kermannioides HBK. and
G. canescens HBK. Nov. Gen. & Sp. 5: 268, 269. 1822, both included
by Hochreutiner in G. calyptrata, the former as a small-leaved
variety; the problem however is beyond the scope of this work; it
was Ulbrich's since he was proposing new names.
Cajamarca: Above Las Huertas, 1,100 meters, Weberbauer 7118,
type. — Hudnuco: Dry slopes, Hudnuco, 24-47 (distributed as G.
calyptrata). — Junin: Tarma, 1008 (distr. as G. calyptrata); Killip &
Smith 21798. La Oroya, Kalenborn 130.
19. CRISTARIA Cav.
Reference: Hochreutiner, Ann. Cons. Jard. Hot. Geneve 21:
347-357. 1920.
Prostrate ascending often tomentose herbs with dissected or
angulately lobed leaves, the flowers axillary or in terminal racemes.
Bractlets none. Anthers clustered at apex of filament-tube. Ovary-
cells many, 1-ovuled ; style branches filiform, truncately or capitately
stigmatose. Carpels in a carpocrater (cup formed by expansion of
carpel and columella bases), uniseriate, usually produced at maturity
into a pair of erect-conniving wings, the cells closed even in fruit or
more or less opening bilaterally, the solitary seed pendulous or
apically affixed horizontally. — Hochreutiner, Ann. Cons. Jard. Bot.
Geneve 21 : 405-428. 1920, has considered the presence (to absence)
of the apical carpel-wings as of secondary importance, the presence
of a carpocrater being primary (Hochreutiner, I.e. 348-357); this
structure, "aptly so designated by Hochreutiner and described by
him in detail" (Kearney, Amer. Midi. Nat. 46: 95. 1951) is most
typically and best developed in this genus where expanded separated
bases of carpels become fused with the enlarged base of columella
or receptacle to form a cupula around the lower part of the fruit.
This student anticipated others by several years in observing,
naming and describing this interesting development.
Indument hirsute; petals 8-10 mm. long C. divaricata.
Indument stellate; petals 5 mm. long, or 12-17 mm. long.
Loosely branched; leaves dissected C. formulosa.
Closely branched; leaves often trifid, lobes pinnate. .C. multifida.
Cristaria divaricata Phil. Anal. Univ. Chile 82: 312. 1892
(1893).
572 Field Museum of Natural History— Botany, Vol. XIII
Much branched, hirsute, the trichomes in part simple; leaves
suborbicular, trifid to base, the segments incised, with narrowly
linear obtuse lobes; peduncles 2 in each axil, longer than the leaves;
sepals densely villous, 4 mm. long; petals 8-10 mm. long. — The
leaves are sometimes 9-lobulate; var. hirsuta Phil., entire plant
white-hirsute. The Peruvian collection agrees with Reiche's de-
scription of PhiHppi's plant except 4 dm. high, petals 15 mm. long,
pedicels mostly solitary (Bruns). It seems probable that the Peru-
vian specimen is referable to C. formulosa Johnst., or that the latter
is synonymous with the present species.
Arequipa: Lomas, Mexia; (Guenther & Buchtien 218, det. Bruns).
Chile.
Cristaria formulosa Johnst. Contr. Gray Herb. 85: 74. 1929.
Laxly ascending annual, the slender stipitately glanduliferous or
sparsely stellate pubescent stems to about 5 dm. long, internodes
2-8 cm. long; stipules lanceolate, entire; petioles as long or longer
than leaves, these dissected, often 3-5-foliolate, gradually reduced
above, often pinnate with more or less remote lobules, the lower
broadly ovate, often cordate or reniform, 3-7 cm. long, nearly as
wide, lightly stellate pubescent; pedicels 1-3 cm. long; calyx 6 mm.
long, the lobes broadly lanceolate; petals roseate, 12-17 mm. long;
carpels 20-25, glabrate, wings oblong-ovate, 1.5-2 cm. long; seeds
about 1.2 mm. high, 0.8 mm. wide. — Distinct by its loosely branched
habit, dissected leaves (Johnston). Determinations by author.
Arequipa: Sand dunes of Mollendo, Mexia OJf.183. Silty fiats
below conglomerate bench near Mollendo, Worth & Morrison 15727.
Chile.
Cristaria multifida Cav. Icon. 5: 11. Obs. 1799.
Erect, rarely 2 meters high, but the stems and branches arcuate-
flexuose, sometimes forming small clumps, lightly pubescent with
minute stellate trichomes or densely only on the calyces, these about
3 mm. long, nearly twice as long in fruit, the rotund ovate lobes
acute; leaves broadly ovate to elliptic, often trifid, the lobes more
or less deeply pinnate with short obtuse lobules; pedicels slender,
soon 1.5-2.5 cm. long; flowers lilac or whitish; petals about 5 mm.
long; fruits 7 mm. across, topped conspicuously at maturity by the
papery carpel-wings.— F.M. Negs. 29765; 29767.
La Libertad: Ditch along railroad, Chicama Valley, Smyth 54-
— Lima: Above Chosica, 1,700 meters, Weberbauer 5318; 144.
J Flora of Peru 678
Lurin, on dryer sands toward the sea, 5938. Near Lima, Dombey,
type. — Arequipa: Tingo, 2,100 meters, CockereU; Pennell 181 18.
MoUendo, Hitchcock 2285^. Yura, CockereU. — Moquehua: In hills
near Moquehua, Weherhauer 7JUj^. Mt. Estuquina, 1,600 meters,
Weberbauer TJM-
SIDA L.
Abutilothamnus Ulbrich, Notizbl. Bot. Gart. Berlin 6: 316.
1915, fide Hochreutiner.
Herbs or shrubs, often half-shrubs, the indument tomentose or
hispid, the leaves usually serrate. Calyx bractlets none (at least in
Peru) the calyx usually angled, the lobes erect or connivent over the
fruit, sometimes much-accrescent. Ovary 5-many-celled, the cells
1-ovuled; stigmas apical. Carpels usually with 2 beaks, awns, cusps
or bristles, sometimes greatly reduced or lacking, indehiscent,
apically slightly 2-valved or sometimes opening irregularly at base,
with no inner appendage, the seed pendulous or affixed horizontally,
without envelope. — My indebtedness to Fawcett and Rendle's
scholarly account of the species of Jamaica (Fawcett and Rendle,
Fl. Jam. 5: 107-120. 1926) will be apparent as many of the widely
distributed species are also in Peru; there is great need for a general
revision. Most of the well-known species listed now from Peru are
illustrated in Rodrigo's excellent account of the species of Argentina,
Rev. Mus. La Plata n. ser. 6, Bot. : 81-212. 1944; the plates show habit,
calyces, carpels and petals. For convenience the key is designed
(it is hoped) for determination of the more common Peruvian species;
even so, many are scarcely distinguishable especially when immature.
I acknowledge my indebtedness to Fries, Sv. Vet. Akad. Handl. ser.
3. 24, no. 2: 18-19. 1947, for the s)mon3rmy and key to the species
centering in S. oligandra; with accumulation of collections the species
may be found to be fewer as some at least of the characters used as
diagnostic are known to vary for better known forms, and except for
Fries' publication I would have merged them into two or three
(rather; to be regretted— B.P.G.H.).
Since this was written the species known also from Panama, the
West Indies and North America have been keyed by Kearney,
Leafl. West. Bot. 7: 138-150. 1954.
Leaves not lobed or obscurely and angulately, rarely subhastately.
Leaves 3 or more times longer than wide, or not at all cordate.
Margins of linear leaves quite entire S. linifolia.
574 Field Museum of Natural History— Botany, Vol. XIII
Margins of leaves at least minutely crenate-serrate.
Prostrate, in age forming mats S. ciliaris.
Erect, more or less ligneous toward base.
Peduncles even in fruit rarely twice as long as calyces.
Leaves canescent puberulent at least beneath.
Flowers 2-2.5 cm. wide, rarely white or yellowish.
S. Weberhaueri.
Flowers much smaller, usually yellow or white.
Carpels usually 6 (5-10), the beaks retrorsely hispid.
S. salviaefolia.
Carpels always 5, the beaks puberulent . . S. spinosa.
Leaves usually soon glabrate; carpels 7-12, the beaks
puberulent .S^. glomerata, S. acuta.
Peduncles all or mostly soon more than twice as long as
calyces.
Leaves subhastately lobulate; pedicels filiform.
*S^. Ruizii.
Leaves never subhastate.
Carpels always 5.
Petals white; carpels opening below S. alba.
Petals yellow; carpels opening above. . . .S. spinosa.
Carpels 7-14.
Leaves usually glabrate; petals yellow or white;
awns short; stamen tube mostly papillose.
S. acuta.
Leaves usually tomentulose beneath; petals yellow;
awns about as long as body; stamen tube
pilose S. rhombifolia.
Leaves about twice as long as wide, broadly ovate or subrotund,
more or less cordate (unless S. acuminata, S. cordifolia, S.
grewiifolia) .
Leaves quite entire S. grewiifolia.
Leaves serrate-margined
Prostrate-ascending, herbaceous or suffrutescent.
Calyx after an thesis obviously accrescent; petals exserted.
S. macrodon.
Calyx not at all accrescent; petals little exserted.
S. veronicaefolia.
Erect, more or less ligneous, at least at base.
Flora of Peru 575
Flowers sessile or shortly pedicelled, rarely solitary.
Indument mostly hispid; carj)els 5 S. urens.
Indument tomentose; carpels 7-12.
Carpels long-awned S. cordifolia.
Carpels awnless S. acuminata.
Flowers long-pedicelled, mostly or all solitary.
Petals 4-5 mm. long; leaves pubescent both sides.
S. paniculata.
Petals 1 cm. long or longer; leaves glabrate above.
S. chachapoyensis.
Leaves at least medially lobed.
Stamen tube glabrous; anthers as carpels 5.
Carpel awns 4-6 mm. long.
Leaf-lobes lanceolate-ovate, acute . S. lomageiton, S. oligandra.
Leaf-lobes obtuse, suboblong, not narrowed at base.
S. patuliloba.
Carpel awns about 14 mm. long S. lomana.
Stamen tube pubescent, anthers 10-20; carpels 7-9.
Leaves lobed deeper than medially, the lobes narrowed to base;
anthers 10 or 20 S. jatrophioides.
Leaves lightly lobed, the lobes ovate; anthers 10 or 15.
S. lomageiton, S. palmata.
Sida acuminata DC. Prodr. 1: 462. 1824.
Erect, branched, ligneous below, to about a meter high, the
younger parts fulvous stellate tomentose especially the leaves be-
neath and the calyces; stipules linear, deciduous, about 1 cm. long;
petioles 3-10 mm. long; leaves elliptic-lanceolate, somewhat cordate
at base, acute or subobtuse, 3-6 cm. long, coarsely serrate; flowers
1-several in the axils, the pedicels usually short; calyx 4-5 mm. long,
the tube not angled; carpels 7-12, not awned, dorsally rounded and
stellate pubescent, reticulate on the inner sides. — The Lima specimen
is the West Indies form (Ulbrich); it seems probable that the single
Peruvian record was a casual introduction (surely — H.).
Lima: Near Lima in 1831, Meyer. Northern South America;
West Indies.
Sida acuta Burm. Fl. Ind. 147. 1768. S. carpinifolia L. f.
Suppl. 307. 1781, fide Hochreutiner.
576 Field Museum of Natural History— Botany, Vol. XIII
Erect-ascending or subprostrate, branching, the usually minute
indument of younger stems, petioles (4-5 mm. long), peduncles and
calyces stellate and simple; stipules 1-1.5 cm. long, sublanceolate
to linear; leaves oblong-lanceolate to ovate, the larger oblique,
rounded to cuneate at base, 1.5-7 (9) cm. long, irregularly serrate,
mostly 3-nerved, early pubescent as petioles, later glabrate; flowers
at first solitary, later with a flowering branch and subsessile, be-
coming more or less capitately or umbellately disposed; calyx 6-8
mm. long, 10-nerved, the lobes acuminate, exceeded by the yellow
petals; carpels 7-12, 3-4 mm. long, laterally reticulate, dorsally
ridged, apically shortly rostrate and puberulous; seeds puberulent
only at hilum. — The var. hispida Schum. has longer indument, in
part. Determinations by Standley. Illustrated, Rev. Mus. de La
Plata, n. ser. 6, Bot. 7: 156. F.M. Negs. 7558; 9368 (var.).
A weedy shrub used for brooms, often a meter or so high, with
orange, rarely white, flowers.
Junin: Peren^ Bridge, Killip & Smith 25315. Puerto Yessup,
Killip & Smith 2633^.— Loreto: Iquitos, Williams 7968. Florida,
Klug 2088.— Uu^nuco: Tingo Maria, Allard 20566.— Cuzco: In
hedges, Urubamba Canyon, Vargas 11063. Santa Ana, Cook &
Gilbert 14^34^. — Puno: (Lechler 2398, type, var. hispida). Old and
New World Tropics. "Jocuchuchupa" (Cook & Gilbert).
Sida alba L. Sp. PI. ed. 2. 960. 1763. S. angustifolia Mill. Card.
Diet. ed. 8. 1768.
Finely stellulate pubescent, becoming glabrate on the leaves
beneath, also more or less tomentulose; stipules 3-7 mm. long,
subulate; petioles to 2.5 cm. long, mostly much shorter; leaves
broadly ovate to narrowly ovate-lanceolate, serrate, 5-nerved, mostly
2-4 cm. long; peduncles about 1 cm. long; early flowers solitary, the
later approximate or short-racemose often followed by a floriferous
branchlet; calyx of S. spinosa; petals white; carpels 5, puberulent
including the 2 short beaks, 2 mm. long, whitish, membranous, ,
irregularly dehiscing below; seeds trigonous, about 2 mm. long.—
This is probably a variety of S. spinosa L. (B.P.G.H.), which is also
my impression.
Peru (probably). Tropical America; Africa; India.
Sida chachapoyensis Baker, Journ. Bot. 30: 324. 1892.
Ligneous, erect, the petioles, these 2-2.5 cm. long, and slender
medially articulate peduncles furfuraceous pubescent; leaves oblong
Flora of Peru 577
or ovate, cordate or subcordate at base, acuminate, 6-10 cm. long,
nearly glabrous above, pubescent beneath; flowers axillary but
paniculate in the upper axils; petals bilobed, nearly 13 mm. long,
much longer than the calyx; carpels 5. — Section uncertain (Baker).
Amazonas: Sesuya, Prov. Chachapoyas, (Mathews S0U9, type).
Sida ciliaris L. Syst. Nat. ed. 10. 1145. 1759.
Prostrate perennial, the spreading 1-several dm. long stems
somewhat ligneous toward the base, glabrate, the young branches
more or less pubescent with stellate or forked trichomes, the oblong-
elliptic to obovate leaves often with simple pubescence above,
usually about 1-1.5 cm. long, the petioles not longer; stipules ciliate,
linear-lanceolate; peduncles adnate to the petiole of the foliose bract,
terminal, with usually several (-8) umbelliform flowers; calyx hir-
sute, 4-5 mm. long or longer, the red petals 6-7 mm. long; carpels
commonly 5-6 (-8 in t5rpe), about 2 mm. long, tuberculate-spiny;
seeds appressed puberulent, about 1.5 mm. in diameter. — The leaves
are serrate only above the middle. Collected in adjacent western
Ecuador. Illustrated, Rev. Mus. La Plata n. ser. 6, Bot. 96. F.M.
Neg. 19684 (var.).
Peru (probably). To Mexico, Paraguay and the West Indies.
Sida cordifolia L. Sp. PI. 684. 1753.
Herb or half-shrub commonly a meter or more high, softly
canescent tomentose occasionally also with some longer indument,
finally glabrate below; stipules filiform, 5 mm. long or longer; upper
petioles usually about half as long as the leaves, these usually sub-
cordate-ovate, obtuse or acute, mostly 3-6 cm. long, serrate, about
equally tomentose both sides; flowers in terminal and axillary dense
racemes or corymbs, rarely solitary in the axils, the articulate
peduncles often exceeding the petioles; calyx 6-7 mm. long; petals
about 1 cm. long, yellow, orange or red-veined; carpels 7-12, 3-4
mm. long, reticulate, dehiscing at apex, this usually provided with
2 long spine-like retrorsely hispid awns; seeds glabrous except at
hilum, about 2 mm. long. — Determinations by Standley. In spite
of the name leaves are rarely cordate, usually subcordate or rounded
(B.P.G.H.). Illustrated, Mart. Fl. Bras. 12, pt. 3: pi 62.
The fiber which the plant contains is said to be so excellent that
its cultivation could be exploited.
San Martin: Tarapoto, Williams 5S97; 67S6. Juanjul, Kliig
S928 (det. Standley).— Junin: La Merced, 5^5.— Loreto: Yuri-
578 Field Museum of Natural History— Botany, Vol. XIII
maguas, Poeppig 2076; Williams 3919; 3996; UJ^6; J^3I^7; 78J^6.
Balsapuerto, Klug 2876. Rio Paranapura, Klug 3928. — Cuzco:
Prov. Anta, Vargas lk7. Prov. Convencion, Weherbauer 79J^8.
Widely distributed in warm regions. "Caballo usa," "sinchi-
pichana" (Williams).
Sida glomerata Cav. Diss. 1: 18, pi. 2. 1785.
Erect, branching, ligneous below, the younger stems and petioles,
these 5-7 mm. long, puberulent; stipules persisting, lanceolate,
acuminate, glabrous but ciliate, 8-10 mm. long; leaves lanceolate
subrhombic, narrowed both ends or truncate to subcordate at base,
usually 3-nerved; flowers terminal and axillary, sessile or nearly,
often globosely capitate mixed with stipular bracts; calyx hispid on
the 5 angles and 10 nerves, 5-7 mm. long, the lobes acuminate;
petals yellow or white; carpels 5, glabrous, reticulate, cuspidate,
2 mm. long, the seeds puberulent only near the hilum. — Type,
Jussieu, without locality, Peru. Probably, practically, a variety of
S. acuta (B.P.G.H.). Determinations by Standley.
In Peru as elsewhere commonly used as a broom; the muci-
laginous sap is reported to serve as soap.
San Martin: Chazuta, Klug 4-137. — Junin: La Merced, Killip &
Smith 23555 (det. Killip). — Loreto: Mishuyacu, clearing, Klug 210.
Yurimaguas, Williams 3966; Jt363. Rio Itaya, Williams 3235.
Rio Nanay, Williams 332. Iquitos, Williams 1306; 1362; 35U6.
Tropical America. "Sinchi-pichana" (Klug).
Sida grewiifolia (Ulbr.) Hochr. comb. nov. in herb. Ahutilo-
thamnus grewiifolius Ulbr., Notizbl. Bot. Gart. Berlin 6: 316. 1915.
Branches few, strict, terete, then stout, the younger as the peti-
oles, these mostly 1-1.5 cm. long, leaves beneath and calyces brown-
ish puberulent-tomentose; stipules linear-lanceolate, about 1 cm.
long, promptly caducous; leaves entire, oblong-ovate, those of the
primary branches 1.5-2 dm. long, 6-10 cm. wide or larger, of the
axillary flowering branchlets to 8 cm. long, 2-3 cm. wide, the 3-5
palmate nerves prominent both sides; peduncles 4-10 mm. long,
articulate 1-3 mm. below the calyx, this 4 mm. long, its lobes
broadly ovate or suborbicular, half as long; petals oblong-obovate,
spreading, 6-7 mm. long, barbate at base, adnate to very short
stamen tube about half its length; styles filiform, about 15, stigmas
depressed-capitate; fruit black, the 15-20 uniseriate acute carpels
separating from axis, dehiscent dorsally, nearly 1.5 mm. high.
Flora of Peru 679
hispid-tomentx)se, the single subglobose seed nearly 1 mm. thick,
villous. — Type a shrub or tree 3-10 meters tall; fruit similar to that
of Abutilon but the solitary pendulous ovule allies the genus to Sida
(author). I cannot understand why this is not considered a Sida;
here is no true argument for a new genus; there is a lot of variation
in the organization of ovules, often inside of one flower (B.P.G.H.).
Rio Acre: Monte Alegre, Seringal San Francisco, Uk 9589, type.
Sida jatrophioides L'H^r. Stirp. Nov. 5: 117, pi. 56. 1789.
S. rupo Ulbr. Bot. Jahrb. 54: Beibl. 117: 75. 1916, fide Clement,
in herb.
Slender, sparsely branched annual, only the upper stems, petioles
(4-10 cm. long), and lax (in fruit) panicle branches densely setulose
and somewhat stellate, or glabrescent except the calyces; leaves few,
subbasally or more than medially 5-7-lobed, suborbicular, the lower
to about 1 dm. wide, the uppermost more or less reduced, early,
especially beneath, sparsely stellulate and setulose, the lobes some-
what narrowed toward base, irregularly serrate and often incised,
acute or subobtuse; pedicels in fruit several mm. to 10 or more long;
calyx about 5 mm. long, somewhat accrescent, villous-setulose, the
ovate-lanceolate lobes acuminate; corolla purple, subcylindrical,
the glabrous obtuse petals 6-7 mm.(?) long; stamen tube rather
densely villous-stellate; anthers 20 or 10 (S. rupo); ovary glabrous
as styles, these 2.5 mm. connate; carpels 5-8, rugulose, with 2 apically
retuse hispidulous awns about 5 mm. long, the single obliquely
ovoid seed nearly 2.5 mm. long (after Ulbrich as to fruit). — The
awns are often broken off or deciduous. Ulbrich separated his species
on the basis of the villous stamen tube; however, according to the
characterization of the t3rpe the tube is villous; in any event the
character above is of questionable specific value. R. E. Fries, how-
ever, maintains the species; the latter's S. decandra, Sv. Vet. Akad.
Handl. ser. 3. 24, no. 4: 16. 1947, from Chimborazo has the long
carpel awns of S. lomana but the 10 anthers of S. rupo; the con-
stancy of these characters remains to be proved. — Illustrated, Cav.
Diss. 5: pi. 131, fig. S (as S. palmata), fide Fries; Jacq., Icon. Rar.
3: pi. 51,7; L'H^ritier, I.e., pi. 56. F.M. Negs. 21604; 29786; 9802
(both as S. palmata) ; 9381.
Lima: Chancay, Dombey, type. San Bartolom^, (Asplund 10810,
det. Fries). San Augustfn Lomas, Weherhauer 52^7. — lea: Pampano
to Huaytara, 1,700 meters, Weherhauer 5392 (tjrpe, S. rupo). —
Moquehua: Toward Torata, Weherhauer 7^27 (det. Ulbrich, S. rupo).
Lomas de Molenca, (Espoto 21, det. Ulbrich, S. rupo).
580 Field Museum of Natural History— Botany, Vol. XIII
Sida linifolia Juss. ex Cav. Diss. 1: 14, pi 2, fig. 1. 1785.
Erect herb or becoming ligneous below, rarely a meter high, with
linear, quite entire leaves, the flowers mostly in dense terminal
corjrmbs, the rather sparse appressed indument in part simple;
petioles as the linear stipules about 4-7 mm. long; leaves 5-15 cm.
long or longer, only a few mm. wide; calyx hirsute, the lobes acute,
about 5 mm. long, the petals to nearly twice as long, white with
purple bases, purple or yellow; carpels 7-9, glabrous, obscurely
comute, 2.5 mm. long; seeds glabrous, scarcely 2 mm. long. —
Determinations by Standley. Illustrated, Mart. Fl. Bras. 12, pt. 3:
pi 57.
San Martin : Tarapoto, Williams 5^29; 58^7; 5901 . San Roque,
Williams 7279; 7493. Lamas, Williams 6^5^. Without locality,
according to Cavanilles, Jussieu. Tropical America and Africa.
Sida lomageiton Ulbr. Bot. Jahrb. 54: Beibl. 117: 73. 1916.
Annual; stipules to 2 mm. long, caducous, hirsute; petioles about
2 cm. long, hirsute and tomentose; leaves about 5 cm. long and
wide, nearly medially 3-lobed, irregularly serrate, the ovate lobes
acute, tomentose both sides; flowers more or less paniculate at the
ends of short slender axillary branchlets; pedicels 2-5 mm. long,
elongating to 1 cm. or more after an thesis; calyx about 6 mm.
long, villous, subglabrous within, lobes ovate-lanceolate, 3 mm. long;
petals purple, stellate puberulent toward obtuse apex; stamen tube
covered below, 3 mm. long, glabrous as styles, these nearly 0.5 mm.
connate, and depressed-capitate stigmas; carpels 5, clavate-pyri-
form, 4.5 mm. long, rugulose, glabrous as the 4 mm. long awns and
the large solitary seed except at the hilum (after Ulbrich). — Not
seen by Fries, apparently referred by Clement (in herb.) to S.
jatrophioides, but suggested by Fries as possibly a part of S. oli-
gandra; however, it closely resembles S. palmata vegetatively and
Hochreutiner asks: is it not that species? That is my impression
but apparently Johnston and Clement don't think so. F.M. Neg.
9383.
Lima: Lomas, San Augustin, Weberbauer 52^0, type. Ambar to
Huacho, 1,200 meters, Stork 111^70 (det. Johnston). Santa Clara,
Rose 18619 (det. Clement). — Arequipa: Posco, Cook & Gilbert J^O
(det. Ulbrich).
Sida lomana Bruns, Mitt. Inst. Allgem. Bot. Hamb. 58. 1929.
Sparsely branched annual with indument of stellate and simple
trichomes, the latter most abundant on the petioles (4-10 cm. long),
Flora of Peru 581
stems above and calyx lobes; stipules 5 mm. long; leaves cordate,
deeply 5-lobed, the ovate lobes irregularly subcrenate-serrate (or
obscurely), scarcely acute, sparsely pubescent with simple trichomes
both sides; flowers many, in dense panicles in the upper axils, the
I>edicels to 2 cm. long (5 cm. in fruit); calyx campanulate, 4 mm.
long (8 mm. in fruit); the lobes 3 mm. long, hispid; corolla 6 mm.
long, stellate puberulent apically; stamen tube 5 mm. long, glabrous
as ovary, the 5 styles 4 mm. long; carpels obliquely pyriform, 5 mm.
long, glabrous, rugose, with 2 spreading awns 14 mm. long, these
retrorsely flavescent pilose except toward base; seeds 2.5 mm. long.
— Nearly S. oligandra as suggested by the author, but distinct by
the long awns. Illustrated, Bruns, I.e. 59. F.M. Neg. 20925.
Arequipa: Lomas, Paseo, (Guenther & Buchtien 193, type; 19j^b).
Mejia, (Guenther & Buchtien 19 U; 19Uo).
Sida macrodon DC. Prodr. 1: 464. 1824. S. physaloides Presl,
Rel. Haenk. 2: 105. 1835, fide Schumann. S. intermedia St. Hil.
Fl. Bras. Merid. 1 : 188, pi. 36. 1827, fide Schumann.
Small, prostrate, suffrutescent, the many simple branches a dm.
or two long, early puberulent tomentulose and with some longer
spreading trichomes, glabrate in age; stipules 3-4 mm. long; petioles
2-2.5 cm. long; leaves subrotund or broadly ovate, cordate at base,
obtuse, rather coarsely serrate crenate, stellate pubescent especially
beneath, usually 1.5-2 (4) cm. long and about as wide; peduncles
2-2.5 cm. long, solitary or binate, slender; calyx subpentagonous,
canescent, soon 6-8 mm. long, 1.5 cm. long or longer in fruit; petals
8-12 mm. long, pilose at base, roseate or purple; stamen tube 3-4
mm. long, pilose; carpels glabrous, 2.5 mm. long, muticous, the
epicarp fragile, tardily dehiscent; seeds smooth, glabrous. — After
Schumann. Carpels said to be only 5 in the Haenke type, 6-11 in
the plant of DeCandolle according to R. E. Fries, Sv. Vet. Akad.
Handl. ser. 2. 42, no. 12: 36. 1908, but otherwise ex char, there
seems to be only one species; DeCandolle wrote "10" and Hoch-
reutiner has kindly looked at the type and noted about 8 visible,
so the number 5 for the Haenke plant could easily be an error; of
course, if preferred, the name of Presl may be used for the plant
of Peru until more material shows the character constant or vari-
able. F.M. Negs. 7995; 32641 (S. physaloides).
Hudnuco: Near Hudnuco, (Haenke, type, S. physaloides). South-
eastern South America.
582 Field Museum of Natural History— Botany, Vol. XIII
Sida oligandra K. Schum. in Mart. Fl. Bras. 12, pt. 3: 321. 1891.
Annual or enduring and lignescent toward base, sometimes
a meter high or higher, glabrate below, shortly stellate and simply-
hispid above or the leaves merely stellate pubescent especially
beneath; stipules fihform, caducous, 5-7 mm. long; petioles 3-5 (or
lower to 8) cm. long; leaves more than medially 3-5-lobed, the upper
3-6 cm. long and broad, the lobes about lanceolate, attenuate to
base, acutish; panicles ample, the pilose peduncles 1.5-3 cm. long;
calyx campanulate, subtomentose, 5-6 (-8 in fruit) mm. long, the
lanceolate lobes acute; petals 5-6 mm. long, marginally stellate
above; stamen tube glabrous, anthers 5; carpels trigonous, dorsally
muriculate, 3^ mm. long, the pilose awns 6 mm. long, the tuber-
culate seeds 2 mm. long.— F.M. Neg. 9388.
Cajamarca: Cascos, Cerro de Catache, Raimondi (det. Ulbrich).
— Cuzco: Galea, Raimondi (det. Ulbrich). Ecuador; Bolivia.
Sida palmata Cav. Diss. 1: 20, pi. 3. 1785, not 5: 274, pi. 131.
1788, fide Fries. S. ricinoides L'H^r. Stirp. Nov. 5: 115, pi. 55. 1789.
Annual or more enduring and lignescent at base, to a meter high
or higher, stellate-tomentose above; stipules filiform, caducous;
petioles about as long as the leaves, these to a dm. or so long, some-
what broader, glabrous or nearly above, more or less stellate-
tomentose beneath, angulately 5-7 lobed, the lobes ovate, serrate;
panicles ample, the peduncles to 1 cm. long or longer in fruit; calyx
stellate-hirsute with some long simple trichomes above, 5-7 mm.
long including the lanceolate deltoid acute lobes, these equaled by
the obovate rounded red-violet petals, accrescent in fruit; stamen
tube hirsute toward the little enlarged base, 3-4 mm. long; anthers
10 or 15; ovule pendent; carpels 7, semiorbicular, glabrous, rugose,
sulcate dorsally, 3 mm. long, the slender rigid retrorsely setulose
awns 4-5 mm. long. — After Fries as to flowers and fruits, based on
Asplund Ecuadorean collections. I doubt if S. lomageiton is separ-
able. As observed by Fries, earlier students have shown that the
date of L'H^ritier's work was actually 1789, not 1785 as printed.
F.M. Negs. 29785; 7998.
Lima: Near Lima, Domhey, type (also of S. ricinoides). — Caja-
marca: Cascas, Raimondi (det. Ulbrich). Ecuador.
Sida paniculata L. Syst. Nat. ed. 10. 1145. 1759. S. floribunda
HBK. Nov. Gen. & Sp. 5: 258. 1822, fide Fawcett & Rendle.
Ligneous below, early yellowish stellate-tomentose, often about
a meter high; stipules filiform, 5-8 mm. long; upper leaves shortly ;
Flora of Peru 588
petioled, to about 5 cm. long, ovate, cordate, more or less acuminate,
unevenly serrate, somewhat puberulent stellate above, tomentulose
beneath; first flowers solitary, the later disposed on branchlets in
each axil forming in age ample terminal leafy panicles, the i>edicels
filiform, elongate; calyx 2.5-3 mm. long, the lobes subacute; petals
dark purple or maroon, 4-5 mm. long; cari>els five, 2.5-3.5 mm. long,
dorsally stellulate, acute or minutely cusped or biaristate; seeds
sparsely appressed puberulent, 1.5 mm. long. The var. rufescens
Baker, Joum. Bot. 30: 295. 1892, refers to the leaf -color of a variant.
Illustrated, HBK., Nov. Gen. & Sp. 5: pi. U7S (as S. floribunda);
Rev. Mus. de La Plata n. ser. 6, Bot.: 120. F.M. Neg. 9801 (S.
floribunda).
Common food plant of Dysdercus (the "cotton-stainer") and
scourge of the cotton grower (Smyth).
Libertad: Trujillo, KiUip & Smith 21520 (det. Killip). Chicama
Valley, Smyth ItS; 15. — Lima: San Lorenzo near Callao, Gaudichaud
2Jt; Andersson. Near Lima, Wilkes' Exped. Chosica, 508. — Cuzco:
Rio Vilcanota, Mexia 80U9 (det. Killip). Guillabamba, Marin 16 U8.
West Indies; Mexico; South America; Africa. "Escoba."
Sida patuHloba R. E. Fries, Sv. Vet. Akad. Handl. ser. 3. 24
no. 2:\b,pl.l, figs. U-6. 1947.
Erect annual, the new branchlets densely pubescent with short
simple trichomes and some longer (1-1.5 mm. long) spreading setae;
stipules linear-subulate, 2-4 mm. long; petioles slender, rigid, to
4 cm. long, shorter above; leaves 3-8 cm. long and little wider,
openly cordate at base, more than medially triparted, the oblong
lobes obtuse, the lateral of the larger leaves with 1 lobule, little if at
all narrowed to base, coarsely crenate-serrate, leixly and shortly
stellate-hirsute above with some decumbent simple trichomes inter-
mixed, more densely stellate beneath; flowers in ample leafy ter-
minal panicles, the pedicels to 2 cm. long or longer in fruit, densely
hirsute as calyces, these 3-4 mm. long, to 9 mm. long in fruit, the
narrow lobes acute; petals red, 4 mm. long, stellate-hirsute without
at apex; stamen tube glabrous, 1.5-2 mm. long; anthers, styles and
carpels 5, the latter 3 mm. long, dorsally tuberculate, glabrous
except the retrorsely spinulose rigid yellowish awns, these 4-5 mm.
long. — Distinguished by author from S. lomageiton as indicated in
key and from S. lomana further in shape of leaves and smaller fruit.
It (as S. lomageiton) may prove to be a variant of S. oligandra.
Determinations by Clement.
584 Field Museum of Natural History — Botany, Vol. XIII
Lima: Shale slope, Matucana, 161 (det. Johnston, S. oUgandra).
Matucana, in rocks, 102 (det. Johnston, S. lomageiton, leaf -form);
(Asplund 112^2, type). Canta, Pennell 1^597, and rocky slopes
near Viscos, 1W4- (both det. Johnston, S. lomageiton).
. Sida rhombifolia L. Sp. PI. 684. 1753. I
Ordinarily much branched, somewhat ligneous below, minutely
stellate puberulent above; stipules linear, as long or longer than the
petioles (3-5 mm.) ; leaves oblong to lanceolate or rhomboid, crenate
to entire at base, 3-nerved, tomentose beneath, crenulate toward the
acute or obtuse tip, usually 5-6 cm. long; peduncles (1) 2.5-3 (4)
cm. long, articulate above the middle, mostly solitary, rarely on
a secondary branchlet or corymbose at tip; calyx 6-7 mm. long,
10-angled, the lobes acute; petals yellow or with red-purplish base,
exceeding calyx, retuse; carpels (7) 10-14, the 1 or 2 awns (some-
times reduced) glabrous or puberulent; seeds brown, subcaudate,
puberulent only at hilum. — A very variable cosmopolitan weed
(Hochreutiner). Three variants in Peru are of little interest: var. j
surinamensis (Miq.) Schum. in Mart. Fl. Bras. 12, pt. 3: 329. 1891,"
var. guazumifolia (Klotzsch) Schum., I.e., and var. Poeppigiana
Schum., I.e. Determinations by Standley except as noted. Illus-
trated, Amer. Journ. Bot. 33: 468, fig. 3; Mart. Fl. Bras. 12, pt. 3:
pi. 63. F.M. Negs. 7985 (var.); 9396 (var.).
Contains much cellulose and as S. cordifolia, said to be som^
times cultivated.
Cajamarca: Tambillo and Cascas, Prov. Cutervo, Raimondi (both^
var. guazumifolia, det. Ulbrich). — San Martin: Boqueron, Allard
22072 (det. Lyman Smith). San Roque, Williams 7333. Zepelacio,
Klug 3606. Tarapoto, Woytkowski 35100 (det. Cuatrecasas). —
Hudnuco: Pozuzo, 4557 (det. Blake). Mito, 1562. Near Hudnuco,
Sawada 66; Kanehira 272 (det. Johnston). Chinchao, Mexia OJi.149
(det. Johnston). — Junln: La Merced, Killip & Smith 23787. Puerto
Yessup, river thickets, Killip & Smith 2683U (det. Ulbrich). — Lima:
Chosica, Mexia 0Jf.09Jf. (det. Johnston); Soukup 2052. Matucana,
383. Rio Rimac Valley, Ball. — Loreto: Yurimaguas, Williams 38^3;
HU; H81; 5106. Rio Nanay, Williams 253; 570. Near Iquitos,
Williams 1303; 1317; Klug J!t35; 1328; Williams 1356 (var. surina-
mensis, det. Ulbrich). Pro- and Caballo-cocha, Williams 1968;
20366 (both det. Ulbrich, var. surinamensis). — Cuzco: Guillabamba,
Soukup 201 . Machupicchu, Vargas 527. — Puno : Aconeque, Metcalf
30577 (det. Leonard). Near Puno, Soukup 461. All warmer
regions.
Flora of Peru 585
Sida Ruizii Ulbr. Bot. Jahrb. 42: 122. 1908.
Erect terete branches early sub-scabrous with minute stellate
trichomes; stipules subulate; petioles 0.5-2 cm. long; leaves lanceo-
late-rhomboid, sometimes subhastate, acute, 2.5-4.5 cm. long, about
1.5-2 cm. wide, unequally and coarsely repand-dentate, densely
tomentose both sides with appressed stellate trichomes; peduncles
slender, 2-4 cm. long, articulate 10-13 mm. below the stellate-
glandular calyx; petals broadly ovate, obtuse, truncate or sub-
emarginate, 6-7 mm. long, 4-5 mm. wide, barbulate at base; stamen
tube 3 mm. long, filaments about 2 mm. long; carpels 5, broadly
ovoid, 3 ntmi. high, with beaks 0.5 mm. long, dorsally densely,
laterally microscopically stellulate, the central axis 3-3.5 mm. high.
— Type a scrap without data but probably from Peru and related to
S. chapadensis K. Schum. of Brazil (Ulbrich). F.M. Neg. 9393.
Lima: Chancay, Ruiz (& Pav6n), type.
Sida salviaefolia Presl, Rel. Haenk. 2: 110. 1835.
Erect perennial herb or flowering as an annual, the stems and
leaves especially beneath tomentulose including the calyces; stipules
filiform; petioles about 1 cm. long or the upper shorter; leaves oblong
or ovate-oblong to lanceolate, subcuneate at base, unevenly serrate,
often about 3.5 cm. long, 1-1.5 cm. wide, reduced above; flowers
solitary or 2 or crowded toward the tips of the stems; peduncles
somewhat longer than petioles; calyx 5-6 mm. long, the lobes acute;
petals yellowish or white with red veins or red at base, not or scarcely
exceeding calyx; carpels about 2-2.5 mm. long (without beaks),
typically 6 (5-8), glabrous or dorsally puberulent toward the re-
trorsely hispidulous awns, these about as long or shorter than the
muricate body of the carpels. — Carpels 5-8, nearly black, with 2
retrorse barbs of varying length terminating the beaks; lateral ribs
form spinose projections (Svenson, Am. Joum. Bot. 33: 466-467.
1946), who referred the Peruvian plant with some doubt to S.
campestris Benth. of Ecuador, with about ten carpels and simple
trichomes on the stems. Actually it seems to be as similar to the
Presl plant, and the name of that, being earlier, is used here. Fur-
thermore, Fries, Sv. Vet. Akad. Handl. ser. 3. 24, no. 2: 13. 1947,
placed here Asplund 562S from Salinas, Ecuador, which in all
probability is the same, noting however the carpel awns as 0.5-0.75
mm. instead of 2-2.5 nmi. long as in type; moreover, Svenson
observed variations in awn-length. This presumably is var. sub-
mutica J. T. Howell, Leafl. West. Bot. 6: 169. 1952, of the Galapagos,
586 Field Museum of Natural History — Botany, Vol. XIII
the carpel awns 1 mm. long or at most half as long as the body of
the carpel. I follow Baker f. who considers it a variety of S. spinosa,
which is again a cosmopolitan weed with innumerable variations —
B.P.G.H. This would be my preference but in this work I try to
present problems rather than to impose decisions. Illustrated,
Svenson, I.e. 408, fig. 1 (as S. campestris).
Piura: Quebrada Mogollon, Amotape Hills, (Haught & Svenson
11521 A). Ecuador; Galapagos Islands.
Sida spinosa L. Sp. PI. 683. 1753. S. angustijolia Lam. Encycl.
1: 4. 1783, fide Fawcett & Rendle.
Stellulate puberulent-tomentose to glabrate (in age) perennial
herb usually less than 1 meter high, the yellow flowers solitary in
the axils or on a subsidiary branchlet, less frequently corymbose;
stipules subulate, 5-9 mm. long; petioles to 2 cm. long, often with
1-2 tubercles or small spines just below the base; leaves ovate-
elliptic to narrowly lanceolate, usually truncate or rounded at base,
softly tomentose beneath, crenate-serrate, 1-3 (4) cm. long; calyx
5-7 mm. long, tomentose, 10-nerved, 5-angled, with deltoid acute
lobes, petals about as long or somewhat longer; carpels 5, brown,
coriaceous, 2.5-3.5 mm. long, antrorsely puberulent toward apex
(where dehiscing) including the 2 short spines, dorsally rugose-
reticulate, glabrous; seeds trigonous, about 2 mm. long. — The
Peruvian collections if correctly determined may have been casual
introductions, but the range of the species is wide. The var. angus-
tijolia (Lam.) Griseb. is said to be a form with subcuneate based
leaves. Probably should be drawn to include as variants S. alba and
S. salviaefolia which compare. Illustrated, Amer. Journ. Bot. 33:
468, figs. 2, k-
Lima: Callao, Wilkes' Exped. — Apurimac: Edge of cultivation,
Goodspeed Exped. 1057 U (det. Standley). To eastern North America.
Sida urens L. Syst. Nat. ed. 10. 1145. 1759. S. pseudo-urens
Baker, Journ. Bot. 30: 294. 1892.
Perennial, erect or the slender branches often sprawling, notably
yellowish-brown hirsute, even to the 5 calyx angles, with long simple
trichomes and with some shorter stellate indument, this mostly on
the membranous leaves; stipules linear, 3-5 (6) mm. long; petioles
1-2.5 cm. long; leaves ovate or oblong-ovate, cordate, acute or
acuminate, serrate, the 5-7 nerves, especially beneath, pubescent,
usually about 5 or 6 cm. long; flowers sessile or shortly peduncled.
Flora of Peru 587
often several in dense clusters, axillary or later sometimes disposed
in terminal heads or spikes with 1 or 2 foliose bracts; calyx 6-8 mm.
long, the acuminate lobes usually slightly exceeding the yellow,
usually red-blotched petals; carpels five, 2 mm. long, glabrous,
bicuspidate. — Ex char., the plant of Baker seems to have no dis-
tinguishing marks unless it lacks the hirsute indument mentioned
only for the calyx, the pubescence elsewhere apparently "rufo-
stellate," thus suggesting S. aggregata Presl of more northern range.
Your reduction is quite right (B.P.G.H.). Illustrated, Mart. Fl.
Bras. 12, pt. 3: pi. 60.
Hudnuco: Pozuzo, U669. Chanchamayo Valley, Schunke 537;
588. — Junfn: La Merced, 5889. — Loreto: Aguaitia, Woytkowski
3jU84. — Cuzco: Marcapata, Herrera 1158. Tropical America;
Africa; Java.
Sida veronicaefolia Lam. Encycl. 1:5. 1783. S. repens Dombey
ex Cav. Diss. 1: 7, pi. 8, fig. 2. 1785. S. Dombeyana DC. Prodr. 1:
463. 1824. S. veronicaefolia Lam. var. Dombeyana (DC.) Baker,
Joum. Bot. 30: 293. 1892.
FVostrate-ascending, sometimes becoming lignescent at base,
more or less spreading, pilose-hirsute or glabrescent; stipules sub-
ulate-filiform; petioles about as long as the leaves, these cordate-
subrotund, obtuse or acute, serrate, mostly 2-3 cm. long and wide;
peduncles almost filiform, solitary or 2-3 in the axils, commonly
longer than the reduced upper leaves; calyx angulately pentagonous,
4-5 mm. long, the basally ovate lobes acute, ciliate, the slightly
longer petals yellowish with purple lines; carpels 5, reticulate, about
3.5 mm. long, muticous (?) or more or less birostrate. — Schumann
in Mart. Fl. Bras. 12, pt. 3: 319. 1891, employs the name of Lamarck
from the East Indies for this apparently variable or not understood
plant and quite possibly there is only one species concerned. How-
ever, ex char., the plant of Lamarck being erect with geniculate
peduncles and lanceolate calyx lobes seems possibly distinct. Ap-
parently the Peruvian names apply to the same species, in spite of
the fact that the carpels seemingly may be little muticous or aristate
(Cavanilles). Apropos, Hochreutiner has added: it is certain that
you are right and the proof that Lamarck's name is prior is that
Cavanilles has cited it along side S. repens in his book, 1: 7. 1785;
a cosmopolitan weed as far as I know and therefore very variable.
Illustrated, Hook. Bot. Misc. 2: 209, pi. 89 (S. Dombeyana). F.M.
Negs. 29787; 29795 (as S. veronicaefolia).
588 Field Museum of Natural History— Botany, Vol. XIII
Cajamarca: Huancabamba, Weherbauer 7111. — Lima: River bed,
Obrajillo, Wilkes' Exped. Near Lima, Jos. de Jussieu, type; Dombey
(carpels rostrate). South America? .
Sida Weberbaueri Ulbr. Bot. Jahrb. 54: Beibl. 117: 68. 1916.
Erect grayish annual (or persisting and more or less lignescent?),
the branchlet tips and leaves more or less tomentose becoming
glabrescent; stipules persisting, linear, fimbriate, to 1 cm. long;
petioles 1-2 cm. long; leaves lanceolate-ovate, cordate, obtuse or
subcuneate at base, acute, 4-7 cm. long, about 1-2 cm. wide, serrate,
the pinnate nerves prominent beneath; flowers 2-2.5 cm. across,
mostly solitary in the axils; pedicels 2-6 mm. long, articulate 1-2
mm. below the 5-angled calyx, the acute slightly fimbriate lobes
glabrous only within; petals oblong-spathulate, white to pink,
purplish or dark red, rarely yellowish within toward base, minutely
ciliolate, 7-8 mm. long; stamen tube 2.5 mm. long, stellulate; fruit
included in calyx, subglobose, 4 mm. high, nearly glabrous, brown,
the mostly 7 carpels 3.5 mm. high, compressed, trigonous, laterally
transversely rugulose and somewhat reticulate, tardily separating,
shortly beaked, the solitary seed glabrous. — Allied to S. acuta with
shorter peduncles and petioles, longer stamen tube, different fruit
characters (Ulbrich). Very likely a form of S. acuta, that cosmo-
politan weed with innumerable variations (B.P.G.H.). Flowers of
Horton 1157 U noted as "cream yellow." Illustrated, Svenson, Amer.
Journ. Bot. 33: 468. ^gr. 5.
Piura: Rocky slopes, Cerro Prieto, (Haught & Svenson 11582).
Hacienda Nomala, Chauro and Serrdn, Weberhauer 5959, type; 598^;
5997. La Brea, Horton 1157J^; 11578 (both det. Johnston). Parinas
Valley, Haught 136; i56.— Ayacucho: Huanta, Killip & Smith 233 ItS
(det. Killip).
21. ANODA Cav.
Reference: Hochreutiner, Ann. Cons. Jard. Bot. Geneve 20:
29-68. 1916.
Low growing somewhat hispid or glabrate herbs with entire to
serrate often hastately 3-lobed or rarely dissected leaves and pe-
dunculate yellow or purple flowers axillary (Peru) or in terminal
racemes. Petals spreading, exceeding the rather stout stamen-tube;
style branches abruptly capitate- or discoid-stigmatose. Carpels
opening laterally by an evanescent septum, spurred dorsally, the
thin endocarp falling with the seed, or merely angled or with a dorsal
Flora of Peru 689
umbo, the endocarp thin, reticulate, becoming sac-like or sometimes
joined with seed; yet again not separating from the pericarp. —
Probably there is only one variable entity in Peru since A. extrema
Hochr., I.e. 64, judging from its relationship, is more likely from
Mexico as is often the case for specimens labelled simply "Pav6n."
Anoda cristata (L.) Schlecht. Linnaea 11: 210. 1837; 44. Sida
eristata L. Sp. PI. 685. 1753. A. hastata Cav. Diss. 1: 38. 1785.
Simple or little branched, early erect, soon decumbent-ascending,
a dm. to several dm. high, more or less hispid-setose especially above
including the deeply parted calyces; leaves variable but commonly
triangular-hastate, sometimes with obscure lateral lobes, often
rather coarsely serrate, usually 4-5 cm. long, 3-4 cm. wide, truncate
or sometimes cordate at base, acute to acuminate; peduncles soon
exceeding leaves; calyx lobes about 7 mm. long, acuminate; flowers
infundibuliform, white, roseate or lavender, usually 1.5- nearly
2 cm. long; carpels 10-20, hispid and rather long-spurred dorsally,
the thin endocarp separating from the pericarp to envelop the seed
firmly. — For full synonymy and varieties of this vegetatively
variable herb see Hochreutiner. Often along trails and in neglected
gardens. Several of the following distributed as A. hastata.
Cajamarca: Socota, Stork & Horton 10100. Libertad: Trujillo,
KiUip & Smith 21515. Hudnuco: Cani, 3U6. Hudnuco, 2067; 3225.
— Lima: Cavanilles' herb., type, A. hastata. — Huancavelica: Man-
taro Valley, WeberbaiLer 7607. — Apurlmac: Abancay and Chincheros,
Goodspeed Exped. 1055S; 10779.— Cuzco: Valle de Santa Ana,
Herrera 979. Prov. Anta, Vargas 219. San Miguel, Cook & Gilbert
1031. Chile to Arizona. "Ruppu" (Cook & Gilbert).
22. BASTARDIA HBK.
Glandular pubescent herbs or small shrubs with small, usually
yellow flowers, mostly solitary in the axils, the petals as calyx lobes
spreading in anthesis. Style branches same number as the uniseriate
carpels, the stigmas apical. Ovules solitary, pendulous. — Fruit
technically a capsule, the 5-8 muticous to aristate thin walled
carpels remaining attached to the axis, not separating septicidally,
opening loculicidally; may be called a schizocarp (B.P.G.H.).
Flowers 10-12 mm. long; carpels biaristate; seeds partly glabrous
or puberulous.
Stamen tube setose; pedicels short or obsolete B. limensis.
590 Field Museum of Natural History— Botany, Vol. XIII
Stamen tube glabrous (typically); pedicels 5-10 mm. long in
flower B. hivalvis.
Flowers about 6 mm. long; seeds described as pilose.
Petals glabrous; nectary 5-lobed; carpels biaristate. .B. parvifolia.
Petals ciliate at base; nectary 5-angled; carpels not arista te.
B. viscosa.
Bastardia bivalvis (Cav.) HBK. Nov. Gen. & Sp. 5: 255. 1822.
Sida bivalvis Cav. Diss. 1: 13, pi. 11. 1785.
Shrub often a meter or two high, the slender branches, petioles,
peduncles and calyces more or less glandular-villous and finely tomen-
tulose or the former indument mostly lacking except on the younger
parts; petioles mostly 1.5-4 cm. long or the lower 8-15 cm. long;
leaves cordate-ovate, acutely acuminate, fine to coarsely crenate-
serrate, lightly to sometimes densely stellate-tomentose both sides
or green and minutely stellulate in age, commonly 4-7 cm. long, 3-5
cm. wide, sometimes 12 cm. wide; pedicels 5-10 mm. long to twice
as long or longer in fruit; calyces deeply parted, the ovate-lanceolate
lobes awned or apiculate, about 5 mm. long; petals 10-12 mm. long,
marginally ciliate at base; stamen column typically glabrous (or
sometimes sparsely hirsute?); fruit 10-12 mm. wide, the carpels
shortly villous, 5-6 mm. high, aristate, the solitary seeds glabrous or
puberulent. — The var. aristata (Turcz.) Hochr,, Ann. Cons. Jard.
Bot. Geneve 20: 144. 1917, is merely tomentulose except the pedicels
and calyces or younger parts, and I suggest the B. limensis is a
synonym (B.P.G.H.).
Piura: Cabo Blanco, Haught 257. Amotape Hills, Haught 172.
— Lima: Matucana, 385. — Huanuco: Pozuzo, 1^623 (var. aristata,
det. Hochreutiner). — Arequipa: Chala, Worth & Morrison 15607
(det. Johnston). — Loreto: Salinas de Pilluana, Huallaga, (Ule 670 Jt,
stamen tube with 4 or 5 trichomes, intermediate to B. limensis, fide
Hochreutiner). To southern North America.
Bastardia limensis R. E. Fr. Sv. Vet. Akad. Handl. ser. 3.
24, no. 2: 23, pi- ^- 1947.
Much branched shrub, the slender younger branchlets early
glutinose-tomentose, some hirsute trichomes more persisting; petioles
mostly shorter than 1 (1.5) cm.; leaves membranous, at first whitish
tomentose, finally green and sparsely stellate puberulent both sides,
triangular-ovate, very openly cordate at base, rather long-acuminate,
obtusely crenate, to 4.5 cm. long, 2.3 cm. wide, usually smaller,
Flora of Peru . 591
densely reticulate veined; flowers subsessile or shortly pedicelled,
most numerous at the tip of the branchlets; calyx 3-8 mm. long, the
oblong acute lobes apiculate, tomentose; corolla yellowish, about
2 cm. wide (in herb.), the petals glabrous except densely ciliate at
base; stamen tube 3 mm. long, setose; carpels 5, tomentulose and
glandular hirsute, 4 mm. long, the slender rigid beaks about 1 mm.
long; seeds glabrous or hirsutulous, subglobose, 2.5 mm. long. — So
glutinous that the paper on which the specimens lay is colored
yellow (Fries). Nevertheless I doubt if the plant is more than
a variant of B. bivalvis (see note under that species). B. sptnifex
Tr. & PI., recently found as near as Ecuador (cf. Fries, I.e. 23, for
description), has spreading indument, narrowly cordate leaves to
7 cm. long, secund flowers, pilose calyx and carpels, these with beaks
3 mm. long.
Lima: San Bartolom^, 1,500 meters, (Asplund 11202, type);
11207. Ambar, shelter of boulders. Stork 11^60 (det. Johnston,
B. bivalvis).
Bastardia parvifolia HBK. Nov. Gen. «fe Sp. 5: 255. pi. ^72.
1822. B. viscosa (L.) HBK. var. parvifolia (HBK.) Griseb. Fl. Brit.
W. Ind. 80. 1864.
Similar to B. bivalvis but leaves often merely acute or if more or
less acuminate not abruptly, petals entirely glabrous, about 6 mm.
long, the seed pilose except the funicle; the nectary is 5-lobed. —
After Hochreutiner, I.e. 143. Fries, Sv. Vet. Akad. Handl. ser. 3. 24,
no. 4: 19-23. 1947, follows Grisebach's disposition which may
indicate best the taxonomic value but account of the awned carpels
it seems allied to B. bivalvis, its character largely defined by its
smaller (1-2 cm. long) canescent-tomentose leaves, pedicels 7-15
mm. long. However, the type is Peruvian; Fries questions the
identity of Cuban specimens, so it is expedient to leave it within
Peru at present as distinct.
Cuzco: Pachachaca, Abancay, 2,200 meters, Vargas US2 (det.
Standley, B. bivalvis). — Cajamarca: Pongo de Rentama to Tome-
penda, Ja^n de Bracamores, (Bonpland, type).
Bastardia viscosa (L.) HBK. Nov. Gen. & Sp. 5: 256. 1822.
Sida viscosa L. Syst. Nat. ed. 10. 1145. 1759.
Velvety pubescent and viscid (except in one variant) half-shrubs
sometimes a meter high, often also with many simple trichomes;
leaves cordate-ovate, acuminate, rather repandly dentate, varying
592 Field Museum of Natural History— Botany, Vol. XIII
in size, often 2-5 cm. long, sometimes much larger; pedicels usually
1.5-3 (4) cm. long; calyx 3.5-5 mm. long, tomentose, the lobes
acuminate, shorter than the mature 5-8-celled fruit, this 5-7 mm.
wide, the 3 mm. long carpels muticous; seeds white-pilosulous, 1.7
mm. long. — Occurs in Peru according to Britton & Millspaugh, and
certainly as near as Guayaquil; also in Argentina (Fries).
Peru (probably). To Mexico and the West Indies.
23. TETRASIDA Ulbr.
Becoming a small tree, the leaves entire, the flowers many,
yellow, fascicled in terminal panicles. Involucre wanting. Calyx
cupulate, with 4 broad unequal lobes. Carpels 5, the solitary ovule
pendulous. Fruit depressed, breaking the persisting patelliform
calyx, the 1-seeded carpels parting from central axis but cohering at
base, longitudinally dehiscent laterally (Ulbrich, Notizbl. Bot. Gart.
Berlin 11. 533. 1932). — Absence of involucel and character of ovule
suggest Sida (Ulbrich) but the calyx and almost completely loculi-
cidal carpels make this a very distinct genus (Kearney) .
I have seen only a specimen labelled Tetrasida polyantha, Wil-
liams 6663, rather poor, without flowers, but with several good
fruits. All of them have at least 8 carpels (loculicidal and septicidal) .
In Notizbl. Bot. Gart. Berlin 11: 533. 1932 the author states that
there are 10 carpels. If the number of 5 styles given by the author
is right it means a most extraordinary fact (we know in the Ureneae
that there are 10 styles for carpels), but if it is true we would have
here 5 styles for 10 carpels. Description of S. tulla being incomplete
it is impossible to tell the truth (B.P.G.H.).
Tetrasida polyantha Ulbr. Bot. Jahrb. 54: Beibl. 117: 66. 1916.
Sida tulla Ulbr. Notizbl. Bot. Gart. Berlin 11: 534. 1932?
Branches, leaves, calyces and petioles, these to 3 cm. long, gla-
brate in age; stipules promptly caducous, linear, 4 mm. long;
leaves ovate or oblong-ovate, cordate, acuminate, to 12 cm. long,
5.5-6 cm. wide, early more or less tomentose, entire, prominently
7-nerved and reticulate beneath; pedicels 1-1.5 cm. long; calyx 4
mm. long, the broadly ovate obtuse to subacuminate lobes unequal;
corolla sub-spreading, nearly 1 cm. long, connate 1 mm. with gla-
brous stamen tube, the obovate petals only ciliate at base, emar-
ginate; fruits 6-7 mm. across, 4 mm. high; carpels subligneous,
obliquely oblong, 2.5-3 mm. broad, more or less appressed stellate
Flora of Peru 598
pubescent as the triangular ovoid seeds. — Tree to 8 meters high
(Raimondi). I am indebted to Dr. Kearney for the probable
disposition of Sida tuUa but in Leafl. West. Bot. 7: 121. 1954 he has
commented on it as follows: the leaves are broader, with a more
open basal sinus, their stellate hairs are somewhat longer and the
flowers are larger in all their parts; the globose flower buds, deeply
reniform anthers and large capitate stigmas are especially note-
worthy; until fruit is available the relationship remains uncertain.
Cajamarca: Ja^n to Bella vista, 600 meters, Weberbauer 6208,
type. Toward Cutervo, 1,400 meters, Raimondi. — Amazonas: Jagu-
anga to Bagua, Raimondi. — San Martin: Tarapoto, Williavis 6663.
Cuzco: Santa Ana, Rio Urubamba, {Cook & Gilbert 1505, type,
S. tuUa).
BOMBACACEAE. Balsa Family
Trees or shrubs characteristically with notably enlarged trunk
with alternate now digitate now simple leaves and often large
flowers solitary or few in the leaf axils. Indument stellate or lepidote.
Calyx coriaceous, generally closed in bud and at anthesis, usually
irregularly lobed or truncate. Petals 5, asymmetrical, contorted in
bud, distinct but commonly connate at base with the staminal tube,
this 5-branched. Stamens 5- many, connate into 1-several bundles,
the anthers 1-2 (or more) -celled. Ovary 2-5-celled, each cell with
2-many ovules ordinarily imbedded in soft or lanate tissue. Fruit
ligneous, usually splitting loculicidally into 5 valves, the smooth
seeds rarely arillate.
Kapok, which has become the common (market) name for the
wooly fibers that fill the mature fruits of a number of species,
principally those of Ceiba pentandra, pantropic but cultivated,
mostly in Java, and Balsa (Ochroma), one of the lightest woods
(or the lightest in markets), are the useful and most outstanding
products, well-known, too, because their exceptional characteristics
caught the public fancy; for an entertaining and informative account
of these trees see Standley and Steyermark, Fl. Guatemala (Fieldi-
ana: Bot. 24, pt. 6: 387-402. 1949, after the various genera); also
Record, Trop. Woods 59: 1-20. 1939, and Williams, Field Mus.
Bot. 15: 309-317. 1936, the two last for technical and economic
information.
Besides the following, the highly interesting Patinoa Cuatr., Rev.
Inst. Bot. App. Agric. Trop. 369-370: 306-313. 1953, could occur
594 Field Museum of Natural History— Botany, Vol. XIII
in Amazonian Peru. Two species are known: P. almirajo Cuatr., I.e.
309, the pulp of the chocolate shaped fruits agreeable, and P.
sphaerocarpa Cuatr., I.e. 312; the former is cultivated as "Almirajo"
in Brazil. The genus resembles Matisia, a part of Quararibea, as
to staminal structure but is characterized by its entire leaves,
tubular calyx, 5-celled ovary and large baccate fruit with coriaceous
epicarp and large lanate seeds; the genus is named for the collector,
Victor Manuel Patino, distinguished agronomist.
Leaves digitately divided.
Stamens numerous, the long filaments from a much-divided tube;
trees usually aculeate 1. Bombax.
Stamens (fertile) few or the tube 5-10-parted with few apical
anthers.
Stamens 5, with partly free filaments, the tube 5-parted.
2. Ceiba.
Stamens with quite sessile anthers, tubes obsoletely 5-dentate.
3. Chorisiai
Leaves simple, rarely unifoliate and articulate, som times lobed.
Flowers as fruits 1 dm. long or longer, the latter filled with silk]
fibers 4. Ochromal
Flowers as fruits much smaller.
Leaves pseudo-simple, articulate; stamen tube cleft.
5. Huberodendronl
Leaves not articulate; stamen tube 5-dentate or divided oi
obsolete.
Stamens united into 5 fascicles; fruit samaroid.
7. Cavanillesiai
Stamens united into a single tube; fruits not samaroid.
Drupes (or similar fruits) with 1-5 seeds; anthers short J
flowers solitary or fascicled 8. Quararibea^
Capsules with many winged seeds; anthers vermiform^
septate; flowers subumbellate 6. Septotheca\
1. BOMBAX L.
Pachira Aublet, PI. Guian. 2: 725. 1775.
Trees, often large, the digitate leaves with 2-9 subentire or re
pand, sometimes crenate-serrulate leaflets, the usually white or yel-j
lowish flowers rather short to long and slender, solitary on single oi
Flora of Peru 696
fascicled axillary or terminal peduncles or rarely paniculate. Calyx
firm or coriaceous, truncate, obsoletely or irregularly dentate
or lobed, always much exceeded by the conspicuously pubescent
and narrow petals. Stamens many, connate toward or at base.
Ovary early 5-celled. Capsules at maturity lanate within or less
often the dissepiments fleshy, the many seeds usually surrounded
by the wool-or cotton-like fiber that is derived from the endocarp.
— Pachira, which, at least practically, ought to be included, but
with apparently the basic character, fruits not lanate-filled, the
seeds larger, has been shown by Ducke, Arch. Jard. Bot. Rio Jan.
5: 160-161. 1930, to be in fact properly as well as expediently a part
of Bombax, since B. obtusum without wool-filled fruits has the
typically small seeds of Bombax, B. paraense lanate-producing
fruits but large seeds of Pachira, and calls to attention the presence
of seeds equipped for water or wind dispersal within the same genus
in several other families as Vochysiaceae, Leguminosae and Och-
naceae. Anyway, the classification is more useful, being sensible,
and equally scientific, or erudite shall we say, with a species of
Bombay which is always that in flower and is still a Bombax even
when in fruit.
Besides the following, WeberbaiLer ^250 from near Celendin,
Cajamarca, may be a new species comparable, at least in Peru, to
B. balanoides Ulbr, but with petiolulate leaflets; flowers about 8 cm.
long; it was given a name by Weberbauer (seemingly never pub-
lished) after the locality (F.M. Neg. 9530).
Leaflets not articulate with the apical petiole disk; petals 5-12 cm.
long; trees unarmed.
Flowers more or less precocious, to 6.5 cm. long; leaflets glabrous.
B. munguba.
Flowers borne with the somewhat pubescent leaves and to 12 cm.
long B. marginatum.
Leaflets articulate; trees usually somewhat aculeate.
Flowers few, showy, 4-many cm. long.
Leaflets retuse or rarely rounded apically; flowers 12-many
cm. long.
Calyx glabrous; petals 12-15 cm. long B. paraense.
Calyx tomentose; petals 3 dm. long or longer.
B. Spruceanum.
Leaflets usually acute or flowers smaller.
596 Field Museum of Natural History— Botany, Vol. XIII
Flowers 2-4 dm. long; leaflets oblong-lanceolate.
B. aquaticum.
Flowers 5-8 cm. long; leaflets oblong-obovate . .B. balanoides.
Flowers at most 2 cm. long, paniculate.
Leaflets obovate, abruptly apiculate B. Ruizii, B. discolor.
Leaflets oblong-lanceolate, acuminate.
Leaves as calyces puberulent B. Weherhaueri.
Leaves as calyces glabrous B. Vargasii.
Bombax aquaticum (Aublet) Schum. in Mart. Fl. Bras. 12,
pt. 3: 233. 1886 (as Pachira aquatica) or Pflanzenfam. 3, Abt. 6:
62. 1890. Pachira aquatica Aublet, PI. Guian. 2: 726. pis, 291, 292.
1775.
Tree, glabrous except the flowers, flowering as a shrub but
becoming 20 meters tall or taller, the narrow high-buttressed trunk
with smooth pale bark; petioles to 2 dm. long or about as long as
the narrowly oblong or oblong-lanceolate obscurely acute or shortly
acuminate leaflets, these 5-7, firm-membranous or subcoriaceous,
yellowish-green; flowers solitary, the truncate ferrugineous calyx
1.5 cm. long, the stellate-pubescent petals 2-3 dm. long; style with
shortly 5-lobed stigma; stamens connate into 15 repeatedly parted
fascicles; fruit to about 2.5 dm. long, ferrugineous, tomentulose
without, the angular seeds 4 cm. long. — Petals reddish-brown and
greenish, stamens white (Ducke); white or greenish-yellow, the =
filaments purple or reddish (various authors). Illustrated, Mart.
Fl. Bras. 12, pt. 3: pi. A6, fig. 2.
San Martin: Juanjui, Klug ^57 (det. Standley). — Loreto: Yuri-
maguas, Killip & Smith 28121; Williams 3937 (both det. Ulbrich).
Caballo-Cocha, Williams 2^1 (det. Ulbrich). "Bellaco caspi,
"wimba" (Williams). To the Guianas.
Bombax balanoides Ulbrich, Notizbl. Bot. Gart. Berlin 6:
156. 1914.
Smooth and glabrous (except the petals), the brown barkec
branches of the small crown terete; stipules promptly caducous^
ovate, 6 or 7 mm. long, 4-5 mm. wide; petioles to 12 cm. long,!
sub terete, with apical disk 8-12 mm. across; leaflets sessile, 5-7,J
obovate or oblong-obovate, subobtusely acuminate to rounded-
obtuse, entire, 14-18 cm. long, 5-9 cm. wide, subcoriaceous, nearly
concolorous but somewhat lighter beneath, the 9-12 moderately
Flora of Peru 597
remote lateral nerves equally obvious both sides; flowers solitary
or few in the upper axils on stout peduncles 2.5-4 cm. long, these
squamulose and with 3 rudimentary bractlets; calyx cupulate,
ligneous, truncate, glabrous also within; petals ovate-lanceolate,
subobtuse, about 4 cm. long, 1.5-2 cm. wide, coriaceous, white (dry-
ing brown-olive), reflexed, finely tomentulose without, sericeous
tomentose within; stamen tube nearly 1 cm. long and about 8 mm.
thick, brownish tomentose except the 10 (more or less) apical
phalanges of many glabrous filaments, these about 3 cm. long; ovary
squamulose, style 5-7 cm. long; fruit (immature) 4 cm. long, 1 cm.
thick, glabrous toward apex. — Flower buds simulate those of oak
(Qiiercus); allied to B. septenahim Jacq. (B. cumanense HBK.) of
Venezuela, but petioles and filaments longer, leaves, stamen tube
and style shorter (Ulbrich). Tjrpe with columnar trunk 6-12 meters
high. F.M. Neg. 9527.
Rio Acre: Seringal Auristella, Ule 9596.
1^ Bombax discolor HBK. Nov. Gen. & Sp. 5: 299. 1822.
Unarmed, the terete younger branchlets puberulent; petioles
6 cm. long or longer, finely canescent tomentose; leaflets 5, sessile,
oblong, cuneate at base, acuminate, the larger about 1 dm. long,
3 cm. wide, crenulate, reticulate veined, membranous, green but
hirsute above, softly stellate-tomentose beneath; flowers paniculate,
pedicels 3-4 mm. long; calyx urceolate, truncate, obsoletely denticu-
late, canescent tomentulose; petals 5, sericeous tomentose without,
pubescent within, barbate-tomentose toward obtuse apex; stamens
60-70, longer than corolla, filaments connate below; ovary hirsute,
5-celled; stigma simple. — Backhuizen referred this, with query, to
the Brazilian B. pvhescens Mart. Nov. Gen. & Sp. 1: 91. pi. 58,
a possibility I have not been able to study; however, ex char, that
species has retuse leaflets. F.M. Neg. 35357.
Cajamarca: San Felipe, Prov. Ja^n de Bracamoros, Bonpland,
type. Chala, Martinet. Cascas, Raimondi.
Bombax marginatum (St. Hil.) Schum. in Mart. Fl. Bras.
12, pt. 3: 223. 1886. Pachira marginala St. Hil. Fl. Bras. Merid.
1:260. pi. 51. 1827.
Branchlets, petioles (1.5-2.5 dm. long) and solitary angled
peduncles (3-6.5 cm. long) glabrous but the 7-9 foliolate leaves
conspicuously ferrugineous tomentose or glabrate above in age;
leaflets sessile and inarticulate on the dilated petiole apex, lanceolate-
598 Field Museum of Natural History — Botany, Vol. XIII
oblong or obovate-oblong, entire, 6-17 cm. long, about 3-8 cm.
wide, attenuate to base, obtuse or obtusely acuminate; calyx cupu-
late, entire, glabrous or tomentulose, 12-18 mm. long, somewhat
wider; petals acute, greenish tomentose or toward base villous and
whitish villous within, 8-12 cm. long, about 1 cm. wide; stamen
tube about as long as calyx, villous at base, finally glabrous; pistil
exceeding filaments; ovary glabrous, stigma obsoletely 5-lobed;
capsules 7.5-9 cm. long, 3-3.5 cm. thick, ovoid, glabrous as the
ovate seeds, these 3-3.5 mm. long. — The leaves of the Peruvian
collection are notably reticulate-veined beneath. Illustrated, Mart.
Fl. Bras. I.e. pi U-
Junin: Chanchamayo Valley, Schunke 1557 (det. Ulbrich).
Brazil.
Bombax munguba Mart. Nov. Gen. «& Sp. 1: 93. pi. 99. 1826.
Stout branches subverticillate; petioles 2 or more cm. long,
dilated at apex; leaves terminally crowded, with 7-8 inarticulate
oblong-lanceolate leaflets, these attenuate at base, the petiolules
to 2 cm. long, acute or acuminate, subcoriaceous, about 1.5-3 dm.
long, 5-11 cm. wide; peduncles 6.5-9 cm. long; calyx 1 cm. long,
tuberculate, entire; petals 5-6.5 cm. long, tomentulose, finely white
puberulent within as the 1 cm. long stamen tube, this apically
parted into 10 phalanges of many filaments, the anthers straight;
ovary as style at base glabrous. — Sometimes 50 meters tall, the
inner bark used for cordage (Williams).
Hudnuco: Tocache, Poeppig. — Loreto: Rio Nanay, Williams 260;
610 (det. Ulbrich). Caballo-Cocha, Williams 2506. La Victoria,
Williams 2866 (det. Standley). Amazonian Brazil. "Huina caspi,"
"punga blanca" (Williams), "monguba" (Brazil).
Bombax paraense Ducke, Archiv. Jard. Bot. Rio. Jan. 4:
124. 1925.
Nearly glabrous tree (except the leaves beneath and flowers),
the branchlets obscurely or not verruculose; petioles to 1 dm. long,
strongly dilated apically; leaflets 5-7, articulate, oblong-obovate,
broadly retuse or rarely rounded at tip, cuneately attenuate to very
short petiolule, sometimes 18 cm. long, 7 cm. wide, papyraceous
(not in Peru), lustrous above, minutely lepidote and opaque beneath,
the fine nerves and veins prominent both sides; peduncles often
1.5-2.5 dm. long, rather stout, striate; calyx at anthesis 1.5-2 cm.
long, finely rugose, glabrous without except often tomentulose at
Flora of Peru 599
apex, sericeous within, eglandular at base, truncate; petals 12-15
cm. long, 6-9 mm. wide, brunnescent stellate-tomentulose without,
rubescent and finely tomentulose within; stamen tube 3-5 cm. long,
a little tomentulose below the apex, the purple stamens greenish
white at tip. — Ducke, I.e. 5: 161. 1930, described the fruit as ovoid,
1.5 dm. long, about 9 cm. thick (dried), thinly ferrugineous tomen-
tose, filled with brown wool, the oblong-elliptic seeds not angled,
2.5 cm. long, 1.5 cm. wide, and remarked: this species is exactly
between typical Bombax and the section Pachira in several of its
characters: the wool filling the mature fruit is quite that of the first
but the seeds have the appearance of those of the last. Surely
doubtful is the determination for the Peruvian specimen, said to
have large white flowers, the heavy coriaceous (2-3) leaflets to 1.5
dm. long, 6 cm. wide, immature obovoid fruit 6 cm. high, 5 cm.
across depressed top, brown-lanate within; common in dense forests
according to the collector, who gives wood description. Field Mus.
Bot. 15: 312. 1936. F.M. Neg. 9540.
Lore to: Manfinfa, Upper Rio Nanay, WiUiams 1097 (det. Ul-
brich, with query). Amazonian Brazil. "Punga blanca de chamisal."
Bombax Ruizii Schum. Bot. Jahrb. 25: Beibl. 60: 17. 1898.
MiUea ecuadorensis Standley, Field Mus. Bot. 17: 199. 1937, fide
Cuatrecasas.
Stout branches quite glabrous; petioles early puberulent, 11-18
cm. long; leaves herbaceous, the elliptic or obovate leaflets shortly
and acutely acuminate, 7-15 cm. long, at or above the middle
3.5-7.5 cm. wide, serrulate, minutely and sparsely pubescent on the
castaneous upper surface, softly and rather canescently pubescent
beneath and the often more than 20 lateral nerves conspicuous;
panicle many flowered, in the type 12 cm. long, nearly as broad, the
branches divaricate; pedicels scarcely 5 mm. long; calyx 2 mm.,
petals 13-14 mm., androecium 11 mm. long. — No known species
has flowers so small, inflorescence so composite (Schumann). The
roots bear many tubers, some attaining 1.5 X 1 dm. (Haught).
Svenson, Amer. Joum. Bot. 33: 415. 1946, suggested this might be
the same as B. discolor while Bakhuizen considered it a possible
synonym of B. globosa Aublet of the Amazon, which, from range
alone, seems less likely. The type by Tafalla from Guayaquil.
F.M. Neg. 9543.
Piura: Cerro Viento, Haughl 105 (det. Cuatrecasas; det. Weber-
bauer, B. discolor). "Pisaiya" (Haught), "quinihuc" (Tafalla).
Southern Ecuador.
600 Field Museum of Natural History— Botany, Vol. XIII
Bombax Spruceanum (Dcsne.) Ducke, Archiv. Jard. Bot.
Rio Jan. 4: 126. 1925. Pachira Spruceana Dcsne. Fl. Serres 23:
46. 1880; Misc. Bot. 7. 1880. P. insignis of Schumann not (Sw.)
Sav. which is B. spectabilis Ulbrich, fide Ducke, I.e.
Glabrous tree, except the flowers, the branches stout; petioles
elongate, much enlarged at base; leaves coriaceous, lustrous above,
the leaflets usually 7 (rarely 8 or 9), oblong-obovate, always rounded
or retuse; peduncles short, stout; flowers 3-4 dm. long; calyx cupu-
late, 2.5 cm. long and broad, tomentulose without as the petals,
especially within, these as stamens deep purple but terminally
white, petals without brownish-red, calyx velvety brown (Ducke).
— The fruit is said to be depressed as in the species with which it has
been confused. Magnificent tree, sometimes 20 meters tall; leaflets
usually 7 (-9); petals 3-4 dm. long, purple within as stamens,
reddish brown without; fruit ovoid, apparently longer than that of
B. aquatica (Ducke). F.M. Neg. 9544 (fide Ducke); 35353.
Loreto: Clearing, Mishuyacu near Iquitos, Klug lJf.79 (det.
Ulbrich). Rio Acre and Amazonian Brazil. "Mamorana grande.
Bombax Vargasii Cuatr. Phytologia 4: 468. 1954.
Small essentially glabrous tree, the nodes of the terminal branch-
lets little enlarged, the few 5-foliolate leaves borne at their tips on
slender rigid petiolules 5-6.5 cm. long, these enlarged at base and
ampliated into a short pentagonous disk at apex; leaflets articulate,
lanceolate, attenuate to the sessile subobtuse base, acutely long-
acuminate, serrulate, 6-11 cm. long, 18-28 mm. wide, subherbaceous,
pale green, sparsely and minutely granulose squamulate both sides,
sublustrous and obscurely venose above, prominently so beneath,
the secondary filiform nerves 1.5-2 mm. distant; flowers as many as
10 in short cymes at tip of exfoliate branchlets; pedicels 3-5 mm.
long, often minutely 3-bracteolate; calyx cupulate, obsoletely 5-
denticulate, 3 mm. long, long-barbate within at base; petals linear-
oblong, about 27 mm. long, 3.5^ mm. wide, whitish pubescent |
within, about one fourth connivent below, reflexing above; stamen
tube glabrous, 6 or 7 mm. long, filaments about 90, shorter than;
petals; ovary sericeous, style glabrous, filiform, about 16 mm. long,
the stigma smooth. — Allied to B. discolor HBK. and to B. Ruizii]
Schum., both with pubescent narrower leaflets, those of the former]
larger, the inflorescence of the latter pubescent, the flowers smaller.
A specimen by Gay, from Cuzco, referred with query by Schumann
to B. discolor probably belongs here or to B. Weberbaueri.
Cuzco: Sisal, Prov. Anta, 2,100 meters, Vargas 1087, type.
Flora of Peru 601
Bombax Weberbaueri Cuatr. Phytologia 4: 469. 1954.
Stout branchlets scarred by the enlarged nodes and petiole bases;
petioles about 5 cm. long, enlarged at base, the apical amplification
small; leaflets 5, obovate-oblong, cuneately narrowed to base, obtuse
and shortly mucronate, serrulate above the middle, 4-6.5 cm. long,
to about 2.5 cm. wide, densely puberulent both sides, the trichomes
above simple or fasciculate, those beneath stellate, the secondary
nerves 2.5-3 mm. distant; terminal panicles leafless, 2-7 cm. long,
stellate-pubescent; pedicels to 4 mm. long; calyx cupulate, 3 mm.
high, truncate, tomentose without, sericeous within; petals linear-
oblong, 20-22 mm. long, 3.5-4 mm. wide, villous-hirsute without,
pubescent and dull salmon color (in herb.) within; stamen tube
6 mm. long, glabrous, ventricose above the base, the 40-60 filaments
little shorter than the petals; ovary hirsute toward the apex, the
filiform style more or less 3-5-parted; capsules ovoid, lustrous
brown, 3.5 cm. high, 2.5 cm. fn diameter (medial); seeds oval,
glabrous, nearly black, 6 mm. long, about 5 mm. thick. — Allied to
B. Ruizii and B. discolor, both with much larger leaflets, different
pubescence. To 10 meters high, large, spreading tree, the bark
brown, smooth; flowers pale yellow, wool brown.
Apurimac: At 2,500 meters, Weberhauer 58U9, type. At 1,800
meters, Herrera 1968, Pacachacos Valley, West 3792. — Cuzco:
Puente Cutuctay, valley of Apurimac, (Bues 1967). "Pati."
2. CEIBA Gaertner
Eriodendron DC. Prodr. 1: 479. 1824. Spirotheca Ulbrich,
Notizbl. Bot. Gart. Berlin 6: 159. 1914, fide Bakhuizen.
Like Bombax — capsules lanate within — but the stamens 5 con-
nate into a tube. — The calyx may be truncate and the spiralled
anthers sometimes much elongate, in one species 4-celled. In my
original preparation of this account I argued for the maintenance
of Ulbrich's genus but actually it seems preferably classified as
a new section, since elsewhere in the family the calyx may be either
truncate or lobed in the same group and the anther character, while
extreme (apart from the number of cells, also variable in the family)
is alone scarcely a generic indication. The branches of young trees
may be somewhat aculeate, as the trunk.
Petals 3-6 cm. long.
Flowers 1-5, calyx to 18 mm. long; ovary glabrous. .C. pentandra.
Flowers 1-2, calyx 3-4 mm. long; ovary sericeous. C trichistandra.
m
602 Field Museum of Natural History— Botany, Vol. XIII
Petals 8-12 cm. long.
Anthers elongate, more or less coiled at least in age; leaves entire.
Petals salmon-rose, merely puberulent; calyx cupulate.
C salmonea.
Petals sericeous lanuginose.
Calyx much shorter than the stamen tube . . . C. trichistandra.
Calyx and stamen tube subequal C. samauma.
Anthers short, to 7 mm. long, scarcely coiled; leaves sharply
serrate C. pubiflora.
Ceiba pentandra (L.) Gaertn. Fruct. 2: 244, pi. 133. 1791.
Bombax pentandrum L. Sp. PI. 511. 1753.
Becoming a beautiful tree, the columnar trunk flanked in age
toward base with thick broad buttresses, the cupola-shaped crown
with very thick branches (these spiny only in young trees) ; petioles
elongate or about as long as the 5-7 lanceolate acuminate often
finely serrate leaflets; flowers precocious, fasciculate on the stems;
calyx about 15 mm. long, campanulate, irregularly 4-6-lobed, the
petals two or three times longer, densely white velvety tomentose
especially without; stamen tube short, the 5 filaments with spirally
twisted anthers; fruits (1) 2-3 dm. long, the many lanate imbedded
seeds about 5 mm. long. — The silky hairs that originate from the
inner wall of the fruit are cylindrical, full of air, impermeable to
moisture and therefore extremely buoyant. Trees sometimes 65
meters high.
Kapok of commerce is the floss of the pods widely used for
buoys, etc. but probably being replaced by plastics and treated
rubber.
Hudnuco: Tingo Maria, Burgos 4.Jf. (det. Standley). — Puno:
Sandia, Weberbauer 13^8. — Rio Acre: Mouth Rio Macauhdn, Kru-
koff 56j^8. Tropics of both hemispheres. "Ceiba," "huimba"
(Burgos).
Ceiba pubiflora (St. Hil.) Schum. in Mart. Fl. Bras. 12, pt. 3:
213. 1886. Eriodendron pubiflorum St. Hil. Fl. Bras. Merid. 1:
266. 1827. C. Mandoni Britten & Bak. f. Journ. Bot. 34: 175. 1896,
fide Bakhuizen.
Branchlets smooth; petioles 1-1.5 dm. long; leaflets 6-7 (petio-
lules to 12 mm. long), ovate or ovate-lanceolate, acuminate, about
5-10 cm. long, 2.5- about 4 cm. wide at or a little above the middle,
Flora of Peru 603
coarsely serrate, membranous; peduncles to 12 mm. long; calyx
campanulate, glabrous without, about 2.5 mm. long, the lobes white
tomentose within; petals 8 (more or less) cm. long, oblong-ovate,
white sericeous especially without; stamen tube cylindrical, sub-
equaling calyx, the stamens about as long as the petals. — The
staminodial annulus is toward the apex of the joined portion of the
filaments; description that of Britten and Baker f. The anthers are
only 6-7 mm. long. The leaflets may be only 3 or 4, entire or lightly
serrulate (C. boliviana Britten & Bak. f. I.e. 174, fide Bakh.); it is
not entirely clear that the Andean tree with more or less coarsely
serrate leaflets is not at least varietally distinct. F.M. Negs. 35366;
23784 (C. Mandoni).
Apurimac: At 2,400 meters, Weberbauer 587 U (det. Ulbrich,
C. Mandoni). — Cuzco: Sta. Rosa, Prov. Convenci6n, Vargas 7195
(det. Cuatrecasas, C. Mandoni). Bolivia; Brazil. "Pati" (Weber-
bauer).
Ceiba salmonea (Ulbrich) Bakh. Bull. Jard. Bot. Buitenz. ser.
3. 6: 198. 1924. Spirotheca salmonea Ulbrich, Notizbl. Bot. Gart.
Berlin 6: 160. 1914.
Branches terete, glabrous but sparsely aculeate with tiny conoid
prickles; leaves lacking in flowering type; peduncles solitary, mi-
nutely squamose, 5-8 mm. long; calyx campanulate-cupulate,
truncate, 10-12 mm. high, 13-14 mm. across or in fruit to 20 mm.,
glabrous without, suberose-ligneous, yellowish sericeous within;
petals linear, obtuse, 7-10 cm. long, 13-15 mm. wide, coriaceous,
tomentose both sides, salmon or salmon-rose; stamen tube cylindric,
puberulent tomentose, 1-1.5 cm. above the articulation glabrous
as the 2-5 cm. long filaments, the linear anthers about 3 cm. long,
2.5-3 cm. wide, the 4 spiralled cells convolute in bud about the
filaments and style; ovary cylindric, 12 mm. long, ferrugineous
tomentose, style 5-8 cm. long, stigma 5-lobed. — A 3 meter shrub
allied to C. Rivieri Dene. (5. Rivieri Ulbrich) of eastern Brazil with
much smaller flowers, conoid stamen tube and ovary (Ulbrich).
F.M. Neg. 9547.
Puno: Sandia to Chunchusmayo, Weberbatier 1116, type. —
Hudnuco: Rio Pozuzo, Weberbauer 6770.
Ceiba samauma (Mart.) Schum. in Mart. FI. Bras. 12, pt. 3:
210. 1886. Eriodendron samauma Mart. Nov. Gen. & Sp. 1: 89.
pi. 98. 1826.
604 Field Museum of Natural History— Botany, Vol. XIII
Fine tree, 20-30 meters tall or taller, the bulging or somewhat
barrel-shaped trunk aculeate, the slender flowering branchlets
smooth, glabrous except for the strikingly pubescent flowers, the
petals orange sericeous- villous without, within, as the calyx, finely
puberulent; petioles 5-10 cm. long, to 2 mm. thick, apical disk
5 mm. across; leaflets shortly petiolulate (2.5-5 mm.), oblong or
oblong-lanceolate, acute at base, obtusely acuminate, 6-15 cm.
long, medially 2.5-6 cm. wide, drying purplish or reddish and
lustrous above, chartaceous; peduncles 1-1.5 cm. long; calyx cam-
panulate, to 4 cm. long, the lobes about 1 cm. long; petals oblong-
spathulate, 10-11 cm. long, 3 cm. wide toward the obtuse tips,
subequaled by the stamen tube, free filaments 6 mm. long, anthers
orange; style glabrous toward tip, villous at 5-angled base; capsule
oblong-ovoid, densely lanate within, the small seeds rusty tubercu-
late. — Martius noted the beauty and possible usefulness of the
lustrous white wool of the fruits. F.M. Negs. 19670; 23783.
San Martin: Juanjui, Klug U2JtIf (det. Standley). — Loreto:
Yurimaguas, Poeppig 123 It. Mishuyacu, Klug 94^9. Mouth Rio
Santiago, Tessmann 1^653 (det. Ulbrich). Bolivia; Amazonian
Brazil. "Samahuma" or "samauma" (Martius; Guarani name),
"huimba" (Klug).
Ceiba trichistandra (Gray) Bakh. Bull. Jard. Bot. Buitenz.
ser. 3. 6: 196. 1924. Eriodendron trichistandrum Gray, Bot. U. S.
Expl. Exped. 1: 182. 1854.
Trunk ventricose about medially to 1.5 meters in diameter, the
stout conical spines to 1.5 cm. long; branches densely aculeate,
branchlets smooth, glabrous; petioles 1-1.5 dm. long; leaflets articu-
late, 5-9, oblong-subobovate, narrowed to base, abruptly caudate-
acuminate, subentire, glaucous beneath, 0.5-1.5 dm. long; inflo-
rescences from axils of defoliate branchlets, the flowers solitary or
several; peduncles stout, somewhat hirsutulous, 2-2.5 cm. long, in
fruit to 1 cm. thick; calyx fleshy, 3^ cm. long, 2.5-3.5 cm. wide,
irregularly 5-6-lobed, the rounded lobes reddish velutinous without,
sericeous within; petals reflexing, obliquely obovate, glabrous within,
sericeous-languinose without except apex and margins (in each case),
4-6 cm. long, 2.5-4 cm. wide, stamens about as long, tube conical,
3-5 times shorter than calyx, glabrous within and without, pha-
langes 3-parted below the middle, the anfractuose anthers versatile;
stigma simple, globose; capsules pendulous, obovoid, glabrous
without, 15-17 cm. long, 7-8 cm. in diameter, seeds to 1 cm. thick.
Flora of Peru 606
glabrous, sparsely scrobiculate. — After Bakhuizen, who illustrates
the flowers and fruits, I.e. pis. SO, SI.
Peru: Type from a garden at Lima (Wilkes Exped.); also in
Java, country of origin unknown.
3. CHORISIA HBK.
Aculeate, branched trees, the digitate leaves with 5-7 entire or
serrate leaflets, the rather large rubescent flowers borne on axillary
or subracemose peduncles that are 2-3-bracteolate before anthesis.
Calyx cupulate, irregularly 2-5-lobed. Petals linear or oblong.
Stamen column annulately 5-10-lobed below the middle, the teeth
2-antheriferous. Ovary 5-celled, the cells many ovuled; style
obscurely 5-lobed. Capsules 3-valved. — Trunk developing as
a beautiful smooth light-colored ventricose column. Named for
L. I. Choris, who accompanied Kotzbue around the world.
Leaflets somewhat serrulate C. insignis.
Leaflets quite entire C. integrifolia.
Chorisia insignis HBK. Nov. Gen. & Sp. 5: 297. pi. j^5,
fig. 1. 1822.
Aculeate bottle-shaped tree, the branches slender; petioles 3-9
cm. long, only about 1 mm. thick, the apical disk 1-2 mm. across;
leaflets 5-7 (petiolules 2-5 mm. long), obovate-oblong, abruptly and
acutely short-acuminate, 5-12 cm. long, 3-4.5 cm. wide medially or
a little above the middle, somewhat serrulate toward the apex,
chartaceous, lustrous above, opaque beneath; peduncles to 2 cm.
long; calyx glabrous, irregularly 2^-lobed, 15-17 mm. long; petals
oblong-obovate, obtuse, plane or more or less undulate-crisped
above, roseate or crimson, canescent puberulent both sides, most
densely without, 5-7 cm. long, about 1.5 (-2.5) cm. wide; stamen
tube 4.5-5 cm. long, staminodia rufous puberulent; ovary glabrous,
attenuate into the basally villous style, this somewhat enlarged un-
der the stigma; capsules attenuate at base, oblong-ovoid, 10-14 cm.
long. — Apparently variable in size of petals and degree of undulation.
C. crispiflora HBK., without description, I.e. pi. 4S5, fig. 2 (type,
Langsdorf, Brazil), is scarcely to be included in C. insignis with
smaller flowers; except for the strongly undulate petals it appears
to be nearer C. speciosa St. Hil. to which species extra Peruvian
material determined as C. insignis probably belongs.
606 Field Museum of Natural History— Botany, Vol. XIII '
Piura: Rio Guiros, Weberhauer 63Jf9. — Cajamarca: Tomependa
and Chamaya, Bonpland, type. — San Martin: Juanjui, Klug IfSOU
(det. Standley, C. crispiflora). Near Tarapoto, Spruce 3928 (det.
Schumann). — Hudnuco: Shapajilla, Woytkowski 8. Pampayacu,
Sawada 6.— Cuzco: Hacienda Santa Rosa, Soukup 513. — Loreto:
Yurimaguas, Poeppig (with ined. name). — Rio Acre: Seringal Auri-
stella, Ule 9575 (det. Dahlem, C. crispiflora). "Lupuna," "guito
algodon," "huimba" (Woytkowski).
Chorisia integrifolia Ulbr. Bot. Jahrb. 54: Beibl. 117: 77. 1916.
Completely glabrous except the flowers, the young terete dark
colored branches rugulose, the broadly ovate bractlets slightly
fimbriate; petioles 6-8 cm. long; leaflets obovate or oblong-oval,
acuminate, 6-10 cm. long, (3) 5-6 (4) cm. wide, entire, the petiolules
about 5 mm. long; peduncles to 2 cm. long; flowers white, about
7 cm. long, the thick calyx to 1.5 cm. long, 3-4-lobed, white serice-
ous within; petals firm, oblong-oblanceolate, 15-18 mm. wide,
marginally entire, undulate, sericeous tomentose without and within
toward the subacute tip, otherwise glabrous; stamen tube glabrous,
6 cm. long, curved, enlarged apically, staminodia fuscous villous,
anthers 4-5 mm. long; ovary sessile, glabrous as filiform style, this
exceeding stamens by 6 or 7 mm. — Much smaller flowered than
C. insignis HBK. and C. speciosa St. Hil. both too, as C. crispiflora
HBK., with more or less serrate leaflets, the latter however with
similar flowers (Ulbrich) ; seems doubtfully distinct from C. insignis
or the latter as interpreted less variable. F.M. Neg. 9551 (narrow
leaved var.).
Cajamarca: Prov. Ja^n, 900 meters, Weberbauer 6195, type.
Junin: Chanchamayo, Raimondi (det. Ulbrich).
4. OCHROMA Sw.
References: Rowlee, Journ. Wash. Acad. Sci. 9: 157-167. 1919;
John H. Pierce, Trop. Woods 69: 1-2. 1942; 70: 20-23. 1942.
Trees with ample simple palmately nerved and angulately lobed
leaves, and large peduncled flowers terminal on the branchlets.
Calyx lobes unequal, 2 acute, 3 obtuse. Stamen tube lobes short,
covered from middle to apex with adnate 1-celled cohering spirally
twisted anthers. Ovary 5-celled; stigmas exserted from the stamen
tube, spiralled. Capsule subligneous, about cigar-shaped but
elongate, the many seeds imbedded in an abundance of cottony
fibers. — See Pierce, I.e. for synonymy and discussion. Record,
Flora of Peru 607
Trop. Woods 59: 15-18. 1939, gave a good summary of the com-
mercial uses of this important timber tree, cultivations, technical
description of the wood (after Williams) and pertinent bibliography,
partly repeated by Standley and Steyermark in their entertaining
account, Flora of Guatemala (Fieldiana: Bot. 24, pt. 6: 396-399.
1949). Williams, Field Mus. Bot. 15: 314-315. 1936, besides techni-
cal wood description quotes at length P. A. Means' description
(Trans. Conn. Acad. Arts & Sci. 27. 1925) of the balsa rafts (carrying
as many as fifty persons) constructed by the Incas (in use in 1530, on
arrival of the Spanish). This author wrote, with the appreciation
of a traveler from eastern North America, that the tree, very showy
in flower, suggests Liriodendron (Tulip tree).
Ochroma Lagopus Swartz, Prodr. 98. 1788. Bombax pyra-
midale Cav. ex Lam. Encycl. 2: 552. 1788. 0. pyramidcde (Cav.)
Urban, Repert. Sp. Nov. Beih. 5: 123. 1920. 0. peruviana Johnst.
Contr. Gray Herb. 81 : 95. 1928. 0. boliviana Rowlee, Joum. Wash.
Acad. Sci. 9: 166. 1919.
Attaining only 20 or 30 meters but the often basally buttressed
trunk a meter in diameter, the pale or reddish brown bark smooth,
the crown small or depressed ; petioles stout, shorter than the ample
more or less angulately lobed and cordate broadly ovate or orbicular
membranous leaves, these in Peru green and glabrous above, fer-
rugineous stellate beneath; flowers 1-1.5 dm. long, the fleshy white
petals much longer than the rounded unequal lobes of the tubular-
campanulate calyx. — Probably the Peruvian tree is var. bicolor
(Rowlee) Standi. & Steyer., Field Mus. Bot. 23: 62. 1944, since
the Peruvian form or race (0. boliviana) has leaves dark green above,
tawny white beneath. Pierce, I.e. 69: 2, observed with reason, since
priority is uncertain, Urban's combination based on uncertain data
and 0. Lagopus, so widely used it would seem wise to maintain it.
Wood important, especially for insulation, weighing 6-8 pounds
per cubic foot, dry; not the lightest wood known but the only one
with equal strength. Some botanists claim to recognize ten or more
species, but for all practical purposes there is only one, 0. Lagopus
Swartz (0. pyramidale (Cav.) Urban), of which the others are
varieties or forms (Record).
San Martin: Juanjul, Klu^ Jk398. Tarapoto, WilHams 5965.
— Amazonas: Chachapoyas, 1,600 meters, Weberbau£r JtSlO. —
Hudnuco: Valley of Rio Chinchao, 1,200 meters. Stork & Horton
9882. Pampayacu, Kanehira 226. Utcubamba, Maraiion, Weber-
608 Field Museum of Natural History— Botany, Vol. XIII
hauer, 191. Cuchero, Poeppig. — Junin: La Merced, 5250; Killip &
Smith 2Jf088; Isern 2069. Puerto Yessup, Killip & Smith 26895.—
Ayacucho: Rio Pieni, Weberhauer 5639. — Cuzco: Potrero, Vargas
8267. — Loreto: Rio Itaya, Williams 166; 3Jt39. Iquitos, Williams
36 U; Klug 1J^66. Yurimaguas, Poeppig 2^70; Killip & Smith 28170.
Puerto Limon, Tessmann 3856 (det. Ulbrich). — Rio Acre: Mouth
Rfo Macauhdn, Krukoff 5308. Bolivia to Mexico; West Indies.
"Topa," "palo de balsa," "balsa," "huampo."
5. HUBERODENDRON Ducke
Tall high-buttressed unarmed trees with pseudo-simple 1-
foliolate leaves (petioles rather distinctly articulate at apex) and
small (for the family) slightly zygomorphic white flowers borne
(in type species) in ample terminal panicles of racemiform cincinni.
Calyx campanulate, valvate, irregularly 5 (3-4) -dentate. Petals
5, elongate-oblong, at anthesis somewhat revolute. Stamen tube
unilaterally cleft below apex, above parted into 5 lobes, the more or
less contorted anthers biseriate, 5 per lobe. Pistils shorter than
stamens. Ovary sessile, 5-celled, with 5 (6) ovules in each series in
each cell. Capsules fusiform or ellipsoid, 5-celled, pericarp ligneous,
5-valved, endocarp forming 5 papyraceous sacs enclosing the 2
longitudinally seriate seeds, these terminally long-alate. — A photo-
graph of a typically buttressed tree and illustrations of the fruits
(H. Patinoi Cuatr.) of this genus of magnificent trees has been given
by Cuatrecasas in Fieldiana: Bot. 27, no. 1: 89-91. 1950. Filiform
styles with 5-lobed stigmas usually exserted from cleft of stamen
tube. Allied by the author to both Septotheca and Bernoullia
Oliver, the former without stamen tube cleft, the latter with scarlet
flowers, leaves of 5 or 6 leaflets, long-exserted style but with similar
inflorescence which recalls that of Raputia (Rutaceae) while the
flowers suggest those of Styrax. Meritoriously honors the active
and able Swiss botanist, Jacques Huber. Perhaps the wood specimen
(Yale 2187; George Barrel) from Amazonian Peru, with native
name "Lupuna," apparently identical to that of Bernoullia (Record)
belongs here, but the name is also recorded for Chorisia.
Huberodendron swietenioides (Gleason) Ducke, Archiv. Inst.
Biol. Veg. Rio Jan. 2: 72. 1935. H. styraciflorum Ducke, I.e. 59.
Bernoullia swietenioides Gleason, Phytologia 1: 109. 1934.
Branchlets robust as the alternate petioles, these to 7 cm. long;
leaves subovate- or ovate-elliptic, obtuse or narrowly rounded at
Flora of Peru 609
base, obtuse or shortly acuminate, often 1-1.5 dm. long, 5-10 cm.
wide, herbaceous-coriaceous, somewhat lustrous and equally finely
reticulate veined both sides, with 7 or 8 lateral nerves; panicles
laxly ferrugineous stellate-tomentose, the cincinni with 6-15 secund
flowers; pedicels to 1 cm. long, stout, about equaled by the char-
taceous calyx, this to 9 mm. wide at apex, appressed whitish-
sericeous within; petals to 2 cm. long, 5-6 mm, wide, finely canescent
sericeous both sides as the stamen tube below, this about 6 mm. long
the obtuse lobes 5 mm. long, not extended beyond the anthers; ovary
densely fulvous tomentose, the style glabrous; capsule fusiform,
glabrous, narrowed to obtuse apex, 2 dm. long, 7 cm. in diameter,
seeds about 14, dark chestnut, the suberose wing 6 cm. long, 1.5 cm.
wide. — To 60 meters high or higher, not rare in inundated forests
of the Purus and Acre rivers (Ducke), thus in adjacent Peru in all
probability. The other known species, H. ingens Ducke, I.e., 60,
has the long subulate stamen-tube lobes extended beyond the
anthers, H. Patinoi Cuatr., I.e., 87, petals only 1.5 cm. long, fruit
ellipsoid, 10-12 cm. thick, seeds larger, the wings 2.5-3.5 cm. wide.
Rfo Acre: Seringal Arisema, {Ducke). Mouth Rio Macauhdn,
Krukoff 5609, tjrpe. Adjacent Brazil.
6. SEPTOTHECA Ulbrich
Stately trees with very large alternate coriaceous leaves, the
younger parts lepidotely stellate. Flowers slightly irregular, sub-
umbellate on elongate peduncles in the upper axils. Calyic tubular,
the upper lobe obtuse, the 3 lower subacute, rigid-coriaceous. Petals
5, cuneate or spathulate. Stamen tube exserted shortly, 5-lobed
at apex, the sessile anthers vermiform, obliquely septate. Ovary
sessile, 5-celled, the many super}K)sed ascending ovules anatropous;
style branches 5, contorted or reflexed. — Differs mostly from Quar-
aribea (Matisia) in the subumbellate inflorescence and vermiform
anthers, according to Ulbrich, who gives a detailed description and
analysis, Notizbl. Bot. Gart. Beriin 9: 128-135. 1924.
Fibers of twigs, petioles and peduncles are of remarkable strength
(Ducke).
Septotheca Tessmannii Ulbr. Notizbl. Bot. Gart. Berlin 9:
129. 1924.
Younger branchlets and leaves both sides or the latter beneath
densely squamate-stellate; stipules lanceolate, 4 cm. long; petioles
6 cm. long; leaves oblong-ovate, deeply cordate, subacute, 3 dm.
610 Field Museum of Natural History — Botany, Vol. XIII
long, 1.5 dm. wide, (more or less), prominently nerved; peduncles
about 1.5 dm. long, 3-5-flowered, the pedicels 2-2.5 cm. long; calyx
about 4 cm. long, the lobes 4-5 mm. long, granulate with large
stellate scales; petals greenish, thick-coriaceous, 3.5-4 cm. long,
stellate without; stamen tube 5-5.5 cm. long, glabrous; anthers
3-4 mm. long, irregularly vermiform, longitudinally dehiscent; fruit
a ligneous loculicidal oblong-cylindrical capsule 8-10 cm. long,
about 4 cm. thick, obtusely beaked, with 5 shallow grooves, slightly
rough, the many seeds extended into a thin subtruncate wing 2.5
cm. long, 8 mm. wide. — Type trunk at 2 meters 6 dm. in diameter;
fruit from Ducke 1657 from Tabatinga at boundary. Ducke, Bol.
T^cn. Inst. Agron. Norte, Bel^m 4: 21-22. 1945, has described in
detail the fruit, observing that it splits into 5 triquetrous parts,
long pendulous from peduncles, the inner portion and septum sub-
papyraceous; fruiting calyx funnelform, about 2.5 cm. long, 2 cm.
wide. Illustrated, Schultes, Bot. Mus. Leaf!. Harvard, 13: pi. 29.
Loreto: Yarina Cocha, Tessmann 3214-, type. Colombia; adja-
cent Brazil. "Utucuru," "sapote rana" (Ducke).
7. CAVANILLESIA R. & P.
Pourretia Willd. Sp. PI. 3: 844. 1800.
Large trees, the palmately nerved more or less lobed leaves
appearing in the crown after anthesis of the roseate flowers, these
disposed in umbelliform cymes. Calyx 5-parted. Petals 5, glandular
at base within. Stamen column contracted above the base, finally
separating into many pentadelphous filaments each with 1 reniform
anther. Ovary 3-5-celled, the ovules geminate and erect from the
cell base. Fruit indehiscent with 5 membranous wings much en-
larged laterally, often with only 1 narrow seed. — Commemorates,
of course, the talented contemporary of the authors.
Leaves palmately lobed, subpeltate, with odor of cumarin.
C. phtanifolia.
Leaves entire, cordate, faintly if at all with aroma.
Flowers red; trunk enlarged medially C. umhellata.
Flowers ochraceous; trunk straight or little enlarged.
C. hylogeiton.
Cavanillesia hylogeiton Ulbrich, Notizbl. Bot. Gart. Berlin
6: 163. 1914.
Flora of Peru 611
Trunk smooth, not or little enlarged, the slender branches
forming an open crown; stipules ovate, 5-6 mm. long, 3-4 mm. wide,
caducous; petioles 5-10 cm. long or longer, subtomentose; leaves
oval, entire, truncate or cordate at base, obtuse or obtusely apiculate,
to 2.5 dm. long, 2 dm. wide, glabrescent both sides but the pinnate
nerves floccose-tomentulose, these very prominent beneath; flowers
precocious, fasciculate at ends of branchlets and subumbellate on
divided 2-4 cm. long peduncles, the pedicels 1 cm. long, tomentulose;
calyx campanulate, 15-17 mm. long, 10-12 mm. broad, subtomentose
without, sericeous within except lobes; petals 2.5-3 cm. long, 7-8
mm. wide, subspathulate, obtuse, ochraceous, glabrous within,
sericeous without; stamen tube cylindric glabrous, 1.5 cm. long,
filaments to 2 cm. long in fascicles or some solitary, anthers reni-
form, 1.5 mm. long; ovary tomentose, ovoid; style to 3 cm. long,
glabrous above as truncate stigma; samara to 18 cm. high, nearly
20 cm. wide, the 5 oval wings about 9 cm. broad; seed oblong-fusi-
form, about 4 cm. long, 7-8 mm. broad. — Distinguished by the
author from C arhorea (Willd.) Schum. "of Brazil and Peru" that
is said to be a shorter tree with thicker rough-barked trunk, the
branches more or less pendent, the leaves and blossoms much
smaller, the latter red, and suggests that C. arhorea may be the same
as C. umbellata. To about 40 meters (Ule). F.M. Neg. 9548.
Rio Acre: Seringal Auristella, Ule 95H; 9595, types. Mouth
Rio Macauhdn, Krukoff 5621. Bolivia; Brazil. "Embirana,"
"pucca lupuna" (Ule).
Cavanillesia platanifolia (Humb. & Bonpl.) HBK. Nov. Gen.
& Sp. 5: 306. 1823. Pourettia platanifolia Humb. & Bonpl. PI.
Aequin. 2: 162. pi. 133. 1817.
Tall beautiful tree, the trunk with smooth pale porous bark, the
crown rather open, subglobose; petioles elongate; leaves sinuate-
cordate at base, the 5-7 nerves and reticulate veins prominent
beneath, somewhat stellate-puberulent both sides, 5-7-(authors) or
often 3-lobed, or younger leaves cordate-oval, entire or 1-2-lobed;
flowers precocious; calyx finely ferrugineous tomentose, campanu-
late, coriaceous, the ovate spreading lobes unequal; petals ligulate,
oblong-obtuse, membranous, to 2 cm. long, flesh-colored without,
reddish puberulent within; stamens red, the tube about half as long
as the petals; ovary pubescent as calyx, 5-celled, stigma 5-parted;
fruit wings semicircular, reticulate veined, to 7.5 cm. wide, mem-
branous; cells 1-seeded, the oblong seed acute both ends. — Some-
612 Field Museum of Natural History— Botany, Vol. XIII
times more than 30 meters tall, the huge trunk considerably smaller
at base than the flask-shaped upper portion, the crown small, open;
the Peruvian specimen has four leaves from one tree, the largest
with petiole 13 cm. long, blade 27 cm. wide, 23 cm. long, the 3 broad
acutish lobes about a fourth as long; other leaves about half as
large; it seems probable that later leaves may have the 5-7 lobes
described for the original tree.
Dr. E. L. Little, Jr., of the Forest Service, U. S. Dept. of Agri-
culture, thoughtfully shared with me (in 1949) his correspondence
with Dr. Petersen, who apparently first discovered the tree in Peru
and called attention to the few leaf -lobes; he found the tree limited
to two small areas.
In part of the range of the species the sweet oily seeds are eaten
and the mild oil used in food and medicinally.
Tumbez: El Caucho and in ravine Trapazola, G. Petersen (det.
Elbert L. Little, Jr.). To Panama. "Pretino."
Cavanillesia umbellata R. & P. Prodr. 97. pi. 20. 1794.
Pourettia arhorea Willd. Sp. PL 3: 844. 1800. C. cordata [R. & P.]
Spreng. Syst. 3: 125. 1826. C. arhorea (Willd.) Schum. in Mart.
Fl. Bras. 12, pt. 3: 237. 1886, probably only as to name.
Trunk stout, enlarged medially, the wood spongy; leaves un-
known in type but said to be cordate; flowers umbellate; pedicels
about 6 mm. long; petals 2 cm. long, elliptic, obtuse, canescent
without, red, stamens about half as long. — Willdenow merely
renamed the Ruiz and Pavon tree but Schumann took up his name
for Brazilian collections, which possibly are referable to C. hylogeiton.
Flowers precocious, short lived, the rapidly forming fruits with
4-5 big wings appearing like a multitude of small lanterns on the
branches; to 30 meters tall or taller, the wood so soft that two men
felled a tree with a few strokes, though they could not embrace the
swollen trunk; crown nearly spherical; the native name refers to the
use of the bark for drum hoops (Ruiz & Pavon). F.M. Neg. 23786.
Hudnuco: Near Pozuzo, Ruiz & Pav6n, type. Brazil? "Huan-
carssacha," i.e., "arbol del tambor."
8. OUARARIBEA Aublet
Matisia Humb. & Bonpl. PI. Aequin. 1: 9. 1805.
Trees or shrubs with entire pinnately nerved leaves but 3-10-
nerved from the base and solitary or fasciculate flowers often sub-
Flora of Peru 613
sessile in the axils or on lateral 2-3-bracteolate peduncles. Calyic
irregularly 2-5-lobed. Petals 5, linear to oblong-obovate. Stamen
column with 5-6 episepalous teeth at apex or more or less parted,
the 6-15 stamens with 2-celled anthers. Ovary 2-5-celled, each cell
with 2 (-3) ovules. Stigma sub-bilobed or capitate, sometimes
5-sulcate, rarely the style lobed, the stigmas capitate. Fruit sub-
drupaceous, coriaceous-fibrous, indehiscent, 1-5-seeded. — The dif-
ferences between Matisia and Quararibea are not constant enough
to justify retaining both of them as Baillon, Adansonia 10: 146. 1871
(Hist. PI. 4: 155. 1873), and Vischer, Bull. Soc. Bot. Geneve s^r. 2.
11: 199. 1919, demonstrated (Cuatrecasas) ; actually the diagnostic
characters are variously developed, all species considered; never-
theless, to avoid the problem, I retained both genera in my original
preparation of this account but later discovered the statement of
Cuatrecasas; at time of press, he has published under both names.
In this group of families number of ovary-cells is often variable, and
therefore not of prime importance in classification. It seems to me
obvious that this is another instance where practical taxonomy may
employ a single group name without disturbing the classification,
this equally satisfied by subgeneric indications. Matisia, as a section,
still commemorates, for the historically interested, M. Matis, "one
of the most distinguished artists of the Royal Botanical Expedition"
(to Colombia).
The foliage of most species has been described as having an
aroma reminiscent of curry, of fenugreek (Trigonella) or of the inner
bark of slippery elm, in accordance, probably, with the earlier
experiences of the writer.
Leaves little or scarcely narrowed to the broadly rounded and more
or less cordate 5-10-nerved base; fruits (known) 4-5-celled
and -seeded; stamen tube more or less parted.
Leaves suborbicular, strongly cordate, about as broad as long.
Indument of leaves obscure or lacking Q. cordata.
Indument obvious, especially on leaves beneath . . Q. stenopetala.
Leaves rather oblong-elliptic, longer than wide, inequilateral.
Q. rhombifolia, Q. inaequilatera.
Leaves more or less narrowed from about the middle to the 3-nerved
base, this rarely openly or minutely cordate at petiole.
Ovary 5-celled; stamen tube deeply parted unless in Q. lomensis.
Calyx not alate, 2-3 cm. long, one side often parted to base.
614 Field Museum of Natural History— Botany, Vol. XIII
Leaves scarcely narrowed to the openly cordate base.
Q. huallagensis.
Leaves narrowed to the acute or rarely rounded base.
Calyx indument red or orange tomentose (in Peru) .
Calyx base rounded; indument soft Q. lasiocalyx.
Calyx base narrowed; indument scabrous. .Q. ochrocalyx.
Calyx indument yellowish-green.
Calyx unequally parted; fruits 2-3-celled.
Q. ohlongifolia.
Caljrx 4-5-dentate; fruits often 5-celled.
Q. putumayensis.
Caljrx alate carinate, scarcely 1.5 (-2) cm. long.
Q. lomensis, Q. bracteolosa.
Ovary 2-celled; stamen tube obsoletely to shortly 5-dentate; calyx
1.5- scarcely if ever quite 2 cm. long except Q. velutina,
Q. machin.
Flowers 3.5-4.5 cm. long.
Stigma peltate; calyx nerves fine or hidden in indument.
Calyx 4 mm. wide medially; petals suboblong, 3 mm. wide.
Q. machin.
Calyx 7 mm. wide medially; petals obovate Q. velutina.
Stigma bilobed; calyx strongly nerved Q, amazonica.
Flowers about 2 cm. long.
Calyx tubular, to 1 cm. long; leaves glabrous Q. Duckei.
Calyx campanulate, about 1.5 cm. long; leaves at least
younger often lepidote Q. Wittii.
Ouararibea atnazonica Ulbrich, Verb. Bot. Ver. Brandent.
50: 91. 1908.
Similar to Q. machin but said to lack the strong odor of Trigon-
ella and the broader calyx conspicuously nerved and angled, the
obtuse subspathulate petals to 11 mm. wide and barely exceeding
the stamen tube, this about 33 mm. long, and, especially, the stellate
style with 2 strongly dilated tomentose lobes about 5 mm. long, the
peltate stigmas 2.5-3 mm. across. — Type 10 meters high. The
herbarium specimen of Klug is strongly scented, but it seems to have
the bilobed style, apparently unique. F.M. Neg. 9552.
Loreto: Florida, Rio Putumayo, Klug 2192 (det. Cuatrecasas).
Amazonian Brazil. "Mutseguinia" (Huitoto).
Flora of Peru 615
Quararibea bracteolosa (Ducke) Cuatr. Lloydia 11: 191. 1948.
Matisia bracteolosa Ducke, Bot. T^n. Inst. Agron. Norte, Bel^m
4: 17. 1945.
Younger branchlets, i)etioles, these 3-13 mm. long, peduncles,
12-18 mm. long, and calyces without rufous tomentose, the last
remarkable by the lateral extension of 5 longitudinal ribs into wing-
like appendages to the 5 elongate- triangular lobes; stipules subulate,
canescent tomentose; leaves oblong-obovate, cuneate attenuate to
base from the upper third, shortly and abruptly acuminate, 1.5-3
dm. long, 5.5-13 cm. wide, flexible-papyraceous, more lustrous
beneath than above, pubescent both sides with some fasciculate
rigid trichomes, more frequently asperous on the nerves, these
prominent beneath, the veins reticulate; peduncles with 3 apical
bractlets 1-1.5 cm. long, 5-8 mm. wide at base, long-acuminate,
persisting; calyx 1.5-2 cm. long, tubular, white sericeous within,
subtruncate at base; petals apparently white, 2.5-3 cm. long, to
8 mm. wide at broadened apex, whitish tomentose without, re-
flexing, the glabrous stamen tube about as long, the lobes 8 mm.
long; fruits conical ovoid enclosed by the cupulate calyx, this with
5 well-developed longitudinal crests. — Moist places of upland forests,
Tabatinga, (Ducke 1782).
Peru : No doubt, as now collected at the boundary as noted above.
Amazonian Brazil.
Quararibea cordata (Humb. & Bonpl.) Vischer, Bull. Soc. Bot.
Geneve s^r. 2. 11: 206. figs. 1,2,3. 1919. Matisia cordata Humb. &
Bonpl. PI. Aequin. 1: 10. pis. 2a, 2b. 1805.
Much branched above a straight cylindrical rugose trunk, the
branchlets green; petioles hardly as long as the cordate suborbicular
leaves, these to 3 dm. long and nearly as wide, membranous, gla-
brous, 7-nerved at base; fascicles of 3-several pale rose flowers
on the branches, the pedicels with 2 or 3 bracts near base; calyx
2-5-lobed, tomentose within and without; corolla sub-bilabiate,
about half again as long as calyx, 2 petals a little smaller than the
other 3, all obovate; stamen tube with 5 linear lobes with about
12 anthers on each lobe; style shorter than stamens, puberulent as
the 5-angled ovary, the stigma 5-sulcate capitate; fruit oval, 1-1.5
cm. long, tomentose, 5-celled, 5-seeded. — Type of Matisia. Tree,
15 meters, light brown bark, thick gummy yellow sap (Mexia);
attains 40 meters, said to be cultivated for its edible fruits.
San Martin: Cultivated, Tarapoto, Spruce Jt21U- Moyobamba,
Weberbauer -4^71.— Hudnuco: Above Cayumba, Mexia 8305 (det.
616 Field Museum of Natural History — Botany, Vol. XIII
Standley). Pozuzo, Ruiz & Pav6n. — Junin: Chanchamayo, Weber-
bauer 1952. — Loreto: Rio Itaya, Williams 170; 3515. Near Iquitos,
Klug 1U70; Killip & Smith 27273. Yurimaguas, Killip & Smith
27652. Caballo-cocha, Williams 2182. Mouth of the Ucayali,
Tessmann 3087 (det. Ulbrich). Adjacent Brazil; Ecuador, "Zapote,"
"sapote de monte," "sapotillo."
Ouararibea Duckei Huber, Bull. Soc. Bot. Geneve s^r. 2. 6:
186. fig. 3. 1915.
Branches and branchlets slender, the younger as the petioles
(1 cm. long) and tubular-campanulate calyces (7-10 mm. long)
furfuraceous tomentulose; stipules 3 mm. long; leaves elliptic,
slightly oblique at obtuse base, obtusely short-acuminate, to 2 dm.
long, nearly half as wide, glabrous, membranous, indistinctly 5-7-
pli-nerved, the venation prominent beneath including the reticulate
veins; flowers on terminal or pseudo-lateral branches; pedicels
5 mm. long, minutely bracteolate at base; calyx 7-10 mm. long, the
lobes unequal, 3 or 4, rounded; petals spathulate, less than 2 cm.
long, 4-5 mm. wide near the rotund or obliquely lanuginate tip,
subequaling the furfuraceous-lepidote stamen tube, this shortly
(2 mm. long) 5-dentate, 4-6 anthers per lobe; ovary 2-celled, the
stigma dilated; fruit immature but with 2 seeds. — Transitional to
Matisia (Huber). Tree 15 meters on terra firma, flowers white
(Krukoff). F.M. Neg. 9553.
Rio Acre : Near mouth Rio Macauhdn, Krukoff 5721 . Amazonian
Brazil; Guiana?
Ouararibea huallagensis (Cuatr.) Macbr., comb. nov. Matisia
huallagensis Cuatr. Phytologia 4: 476. 1954.
Robust sulcate branchlets sparsely stellate-pilose; petioles rigid,
about 2 cm. long, tomentulose; leaves oblong-elHptic, rounded-
cordate at base (apex unknown), apparently about 2.5 dm. long and
half as wide, glabrous, rather conspicuously reticulate veined both
sides, the 6 or so lateral nerves prominent beneath; flowers opposite
the leaves, solitary, the peduncles 5 cm. long, gradually enlarged
toward apex, densely ochraceous tomentose with stellate and fas-
ciculate trichomes; buds elUptic-oblong, 3 cm. long, with 4 apical
auricles 2-3 mm. high; calyx densely tomentose, little striolate, 3
cm. long, coriaceous, attenuate at base, irregularly 4-lobed, the lobes
5-6 mm. long, somewhat plicate-alate; petals to 5 cm. long, 8-10
mm. wide at the obtuse tip, glabrous within, stellate-tomentose
without, subequaled by the puberulent stamen column, this with
Flora of Peru 617
5 linear lobes 8-10 mm. long, the few anthers 4-5 mm. long; fruit
rotund-depressed, seemingly 5-celled, rounded at base, flat at top,
appressed tomentose, nearly 5 cm. broad, 2-8 cm. high, the sub-
elliptic seeds about 2.5 cm. long, 1 cm. broad, the subenclosing
calyx coriaceous, the peduncle robust. — Distinguished from the
other species of the section Matisia by the cordate leaves.
Loreto: San Ram6n, Yurimaguas, Williams J^72 (det. Ulbrich,
Q. guianensis) . "Huayhuash sapote."
Quararibea inaequilatera Cuatr. Lloydia 11: 187. 1948.
Known to attain 20 meters; petioles rigid, 7.5 cm. long; leaves
strongly oblique, ovate-elliptic, rather deeply emarginate at base,
one side narrowly and truncately lobate, the other lobe much longer,
rounded, apically obtuse or apiculate, rounded, to 3 dm. long or
longer, about 2 dm. wide, green and glabrous above except lightly
puberulent nerves, densely stellate- tomentose beneath; nerves
palmately radiating, 10, elevated, with 2 lateral ones most prominent,
secondary lateral nerves 7, ascending, the transverse veins reticulate;
peduncles on trunk, 2 cm. long, sparsely pilose; fruits pyriform in
cupulate cal3rx, stellate-tomentose, 2.5-3 cm. long, 5-celled, the
5 seeds 2 mm. long. — Unique among the 3 other species with asym-
metrical leaves in the pyriform fruit, tomentose leaves (author).
Peru (undoubtedly). Adjacent Colombia.
Quararibea lasiocalyx (Schum.) Vischer, Bull. Soc. Bot.
Geneve s^r. 2. 11: 206. 1919. Matisia lasiocalyx Schum. in Mart.
F\. Bras. 12, pt. 3: 239. 1886.
Glabrous except petioles, these 1-2 cm. long, enlarged both ends,
stipules (5-6 mm. long), peduncles and flowers, the indument
tomentulose, this ferrugineous on the peduncles (2 cm. long) and
calyces (20-23 mm. long); leaves oblong-elliptic or -lanceolate,
rounded or subcuneate at base, acuminate, often 3-4 dm. long,
7-10 cm. wide at the middle, membranous, 3- or obscurely 5-nerved;
calyx campanulate, 8-9 mm. in diameter; petals oblong-spathulate,
rounded at apex, 2.5 cm. long, 7-8 mm. wide, subtomentose above,
glabrescent toward narrowed base, finely puberulent within; stamen
tube finally twice as long, tomentose only without above the base,
5-lobed; stigma simple. — Fruit depressed obconical, 6 cm. high,
7 cm. wide, in the horizontally dilated calyx appearing like a slightly
concave plate, 5.5 cm. across; the finely areolate and rugulose fruit
618 Field Museum of Natural History — Botany, Vol. XIII
is covered with a tenuous brown tomentum (Ducke). Illustrated,
Schumann, I.e. pi. 48.
Loreto: Balsapuerto, Klug 3025 (det. Cuatrecasas). Amazonian
Brazil.
Ouararibea lomensis Cuatr. Lloydia 11: 187. 1948.
Like Q. putumayensis at least as to leaves, but the nervation
more elevated; fruiting peduncles 1-1.5 cm. long with 1-3 persisting
apical bracts, the calyx 4-5-lobed, 14-18 mm. high, fruit ovoid-
subpyriform, 25-28 mm. long, 12-15 mm. broad. — The Peruvian
collection has elliptic-obovate leaves to about 1.5 dm. long, half as
wide, attenuate to rounded or acute base, acutish, chartaceous,
minutely and sparsely scurfy stellulate below on the basal prominent
nerves, the conspicuous reticulate venation apparent on the upper
surface; peduncles and calyces, these 1.5 cm. long, brownish stellulate
puberulent, the lobes seemingly (from dried calyx) carinate-alate;
petals glabrous within, lightly sericeous without, oblong-spathulate,
about 2 cm. long; stamen tube 3 cm. long, shortly dentate, the
anthers few. — Perhaps not distinct from Q. bracteolosa (Ducke)
Cuatr. Fruit edible; flowers white; 8 meters tall (Klug).
Loreto: Florida, Rio Putumayo, Klug 2039 (det. Cuatrecasas).
Adjacent Colombia. "Zapote."
Ouararibea machin Macbr., spec. nov.
Ut videtur Q. guianensis perafRnis, foliis oblongo-ellipticis,
obtuse acutis, circa 2 dm. longis, 8-10 cm. latis; floribus lateralibus,
4-5 cm. longis; pedicellis 5-10 mm. longis; calycibus irregulariter
4-5-lobatis. — The species of Aublet is apparently unknown within
Peru but this collection simulates it except for the smaller flowers
borne laterally and the more lobed calyx, this with a tendency even
to split on one side; as in the related species it is very slender, finely
striate and minutely tomentulose as the linear petals both sides and
the tardily exserted stamen-tube. The fruit is unknown, but in all
probability is also 2-celled as indicated by the similar stigma. Type
8 meters high, trunk 4 dm. in circumference, flowers white, leaves
slightly aromatic.
Loreto: River bank, Gamitanacocha, Rio Mazdn, Schunke 290,
type (det. Standley, Q. guianensis). "Machin."
Ouararibea oblongifolia (Poepp. & Endl.) Vischer, Bull. Soc.
Bot. Geneve s^r. 2. 11 : 206. 1919, as ohlongiflora. Matisia oblongifolia
Poepp. & Endl. Nov. Gen. & Sp. 2: 35. pi. 150. 1838.
Flora of Peru 619
Small glabrous tree, the erect trunk about 5 cm. in diameter;
branches few, slender, flexible; petioles 3.5-6 cm. long, .angled;
leaves elliptic-oblong, rarely obovate, acute at base, shortly and
obtusely acuminate, 2-3 dm. long, 8-11 cm. wide at the middle;
peduncles terete, 3-4 cm. long; calyx obconic or campanulate,
unequally 3-5-parted, one side always slit to base, 2.2-2.5 cm. long,
10-12 mm. wide at apex, finely rugulose; petals oblong-obovate,
very obtuse, 28 mm. long; stamen tube medially 5-parted, puberulent
within; drupes ovoid, obtuse, 2-3-celled. — Strange that this has not
been collected since the type; possibly there has been a misinter-
pretation in character and it will be found to include one or more
of the more recently described forms.
Loreto: Yurimaguas, Poeppig, type.
Quararibea ochrocalyx (Schum.) Vischer, Bull. Soc. Bot.
Gendve s^r. 2. 11 : 206. 1919. Matisia ochrocalyx Schumann in Mart.
Fl. Bras. 12, pt. 3: 238. 1886.
Younger slender terete branchlets, petioles (3^ cm. long),
peduncles (2-3 cm. long) and especially calyces orange-ochraceo-
tomentulose with multiradiate trichomes; leaves broadly oblong or
obovate-lanceolate, acute at base, shortly and abruptly acute or
cuspidate, 3-pli-nerved, 1-2 (3) dm. long, 5-7 cm. wide; calyx
campanulate, attenuate at base, (1.5) 2-3 cm. long, irregularly
3-4-dentate; petals oblong-spathulate, rounded at tip, 2.5-4.5 cm.
long, 7-9 mm. wide, subtomentose without, finally glabrous within,
the stamen tube twice as long, tomentose only without; fruits ovoid,
nearly enclosed in the rough puberulent calyx, at least 3 cm. long,
2 cm. in diameter. — Tree to 15 meters tall or taller of upland forests.
Wood of this species like that of Quararibea (Record). Deter-
minations by Standley except as noted. M. lasiocalyx has similar
fruits but brown sericeous tomentose, calyx thinner (Ducke).
Loreto: Florida, Klrig 2205. Yurimaguas, WiUiams WO.
Iquitos, Killiv & Smith 27020; 27025 (det. Ulbrich); WiUiams 8791;
Klug 1110; 1137. Rio Itaya, Williams 3389. Pebas, Williams 1576.
Balsapuerto, Killip & Smith 28622. Rio Nanay, Williams 6j^8
(det. Ulbrich). To French Guiana. "Macacha-ey" (Klug), "ma-
chinnaccha" (Schunke), "zapotillo" (Williams).
Quararibea putumayensis Cuatr. Lloydia 11: 186. 1948.
Slight tree with ochraceous-green branches and calyces, these
densely and minutely sublepidote-stellate tomentulose, glabrous
620 Field Museum of Natural History— Botany, Vol. XIII
subcoriaceous leaves; stipules linear-lanceolate, acute, 1 cm. long;
petioles stout, 6-16 mm. long; leaves elliptic-lanceolate, attenuate
both ends, apiculate at apex, often 1.5-3 dm. long, 4.5-11 cm. wide,
basally triplinerved and with 4-5 secondary nerves arcuate-ascend-
ing; peduncles 3 (-4.5 in fruit) cm. long; calyx coriaceous, tubular-
campanulate, 2 cm. long, the 4 or 5 triangular teeth obtuse or acutish,
sericeous villous within; petals to 3 cm. long, about a third as wide,
white, villous without, sparsely pubescent above; stamen tube 5
cm. long, with 5 linear obtuse divisions 6.5-8 mm. long, anthers 6;
fruit indurate in herb., densely brownish-green tomentulose, often
5-celled, 5-seeded, 2.5 cm. long, 3 cm. broad. — Similar and perhaps
also in Peru is Q. lomensis Cuatr. I.e., 187, the leaf nerves more
elevated, peduncles 1-1.5 cm., with 1-3 persisting apical bracts,
fruiting calyx cupulate, more or less 4-lobed, 14-18 mm. high, fruit
ovoid-subpyriform, 25-28 mm. long, 12-15 mm. broad. Here, by
my key, largely expedient, would be sought Q. lecythicarpa (Ducke)
Cuatr. (Matisia, Ducke, Bol. T^cn. Inst. Agron. Norte, Bel^m 4:
18. 1945), the calyx indument tomentulose apparently as in Q.
lasiocalyx but the fruits depressed pentagonal instead of pyriform.
Loreto: Soledad, Rio Itaya, Killip & Smith 29727 (fide Cuatre-
casas). Adjacent Colombia.
Ouararibea rhombifolia (Cuatr.) Macbr., comb. nov. Matisia
rhombifolia [Standi.] Cuatr. Phytologia 4: 478. 1954. Q. rhombifolia
Standi. Bol. Mus. Hist. Nat. Lima 4: 476. 1940, name.
Branchlet tips, petioles, these about 6 cm. long, and the very
prominent 9 basal leaf nerves beneath more or less ferrugineous with
a dense stellate-tomentose indument; leaves subobovate, somewhat
pandurate, openly cordate at base, rounded at apex, the few known
about 2-2.5 dm. long, to 6 cm. wide, dark green but minutely and
sparsely stellulate above, the conspicuous secondary nerves and
reticulate veins puberulent beneath with fascicled or simple tri-
chomes; peduncles 2.5 cm. long; fruit subglobose, mucronate at
apex, about 3.5 cm. thick, 5-celled, the 5 seeds about 16 mm. long,
7 mm. broad. — To 15 meters tall, the round-tapering fruit appearing
velvety golden-brown at a distance, coarse-textured in hand (col-
lector).
Hudnuco: In jungle at 630 meters, Shapajilla, Woytkowski, type.
Ouararibea stenopetala (Standi. & Cuatr.) Macbr., comb. nov.
Matisia stenopetala Standi. & Cuatr. Phytologia 4: 568. 1954.
Flora of Peru 821
fir Small pubescent tree, striking by reason of its relatively large
orbicular deeply cordate leaves — several dm. across — and its red-
violet flowers with petals to 5 cm. long but only 6 or 7 mm. wide;
indument early ferrugineous tomentose, the leaves becoming gla-
brate above, the nerves and reticulate veins beneath finely and
rather densely stellate puberulent; pedicels fasciculate, 2.5-3 cm.
long, stellate- tomentose; calyx tubular, 1.5 cm. long, 5 mm. broad,
pale ochraceous stellate-tomentose, sericeous within, unevenly
3-4-lobate; petals membranous, pubescent only without, the tri-
chomes simple; stamen column tomentose, 4 cm. long, the 5 linear
antheriferous lobes 8 or 9 mm. long, the 6 cells contiguous; style
stellate-setose, the stigma capitately lobed; fruit globose, tomentose,
5-celled, 5-seeded, the persisting calyx coriaceous. — AfRne, accord-
ing to authors, Q. bicolor (Ducke) Cuatr. (Matisia bicolor Ducke),
Archiv. Jard. Bot. Rio Jan. 3: 210. 1922, with different leaves and
indument, calyx and stamen-column. To 20 meters tall, the flowers
borne on the main stems. F.M. Neg. 9561.
Loreto: Balsapuerto, Klug 2972, type. Mouth of the Santiago,
Teasmann S98S (n. sp., Ulbrich, in herb.).
Quararibea velutina Cuatr. Phytologia 4: 475. 1954.
Eugulose branchlets at tips, petioles, these 5-7 mm. long, and
linear-lanceolate acute stipules (about 11 mm. long) tomentulose;
leaves subelliptic, oblong-elliptic or slightly obovate, oblique at
obtuse or rounded base, obtuse or obtusely apiculate at apex, 9-14
cm. long, 4-6 cm. wide, minutely and sparsely stellulate above, the
secondary nerves there obvious, sparsely stellate-pilose beneath and
the 5 secondary nerves there prominent; flowers solitary; peduncles
about 1 cm. long, 2-3-bracteolate above the base, enlarged toward
the apex, velvety tomentose as the tubular-conical 16 mm. long calyx
without with trichomes stellate or fasciculate, the branches fine and
coarse, villous sericeous within, sub-trilobed ; petals white, linear,
membranous, about 4 cm. long, 8-10 mm. wide near the rounded
tip, narrowed to base, stellate pilose both sides; stamen column 3.5
cm. long, stellate tomentose, the 5 obtuse lobules 2 mm. long, the
anthers 5-seriate, about 6 cells in each series; style pubescent,
strongly enlarged at apex; ovary 2-celled, the cells biovulate. —
Allied to Q. loretoyacensis Cuatr. but leaves and flowers different in
size and, especially, pubescence of calyx similar to that of Q. ochro-
calyx.
Loreto: Rio Mazdn, JosS Schunke 235, type (det. Standley,
Matisia ochrocalyz). "Machinfiaccha."
622 Field Museum of Natural History — Botany, Vol. XIII
Ouararibea Wittii Schum. & Ulbr. Verhandl. Bot. Ver. Brand-
enb. 50:90. 1908.
Appearance of Q. ochrocalyx but more allied to Q. machin; stipules
lanceolate, 5 mm. long; petioles 1-1.5 mm. long, much enlarged
beneath the oval leaf-blade, this obtuse or narrowed at base, obtuse
or subacuminate at apex, about 1.5 dm. long, 8-11 cm. wide, early
almost microscopically lepidote-stellate, soon glabrescent, the
midnerve and pinnate nerves very prominent beneath; peduncles
to 2 cm. long or longer; calyx infundibuliform-campanulate, yellow-
ish with minute stellate-lepidote indument, about 1.5 cm. long, un-
equally (3 and 6 mm.) bilobed, the 4 oblong segments rather obtusely
acuminate, sericeous tomentose within, in fruit to 2 cm. long,
lignescent; petals 2.5 cm. long, canescent stellate, the stamen tube
nearly as long, shortly 5-lobed, the anthers sessile; style subpeltately
5-lobed, ovary 2-celled; fruit oblong-ovoid, densely lepidote-stellate;
seeds about 1 cm. long, 3 mm. across. — Type 8-12 meters, (Tess-
mann) tree trunk 1.5 dm. in diameter. F.M. Neg. 9556.
San Martin: Chazuta, Klug Jf-lJ,.! (vel afRne). — Loreto: Mouth
Rio Santiago, Tessmann Jf233 (det. Ulbrich). Yurimaguas, Williams
U869 (det. Ulbrich).— Rio Acre: Ule 9592. Amazonian Brazil.
"Sapotillo," "hujTiash-sapote."
STERCULIACEAE. Cocoa or Cola Family
Usually stellate-pubescent soft- wooded trees or shrubs (rarely >
climbing) with alternate stipulate simple serrate or lobed leaves and
panicled or cymose often showy flowers, the petals if present
and contorted in bud, the 5 (or 3) sepals connate, valvate. Stamens
many and connate or few and nearly distinct; anther cells always
2 or more, globular or linear. Ovary free with 2-5 (rarely 10-12)
more or less connate styles or in Waltheria only 1, sessile or stiped,
each with 1 or more ovules. Fruit dry or fleshy, usually 5-celled,
loculicidally dehiscent or separating into folicles or cocci.
Cocoa (and therefore chocolate) is supplied by the bean-like
seeds of the fruits of Theobroma (T. Cacao L.) of South America,
while Cola or Kola nuts (Cola acuminata (Beauv.) Schott & Endl.)
are native to West Tropical Africa; the family is important eco-
nomically also as the source of several fibers.
Flowers hermaphrodite; petals present.
Petals soon caducous.
Flora of Peru 623
Ovary as stamens on long stipe finally longer than spiralled
fruit; petals red or white 8. Helicteres.
Ovary sessile or shortly stiped, fruit not spiralled; petals brown-
purple or in Gttazuvia yellowish.
Ovary (as fruit) smooth; anthers 2 or more in each stamen-
tube sinus; leaves ample.
Petals produced into an elongate narrow appendage; leaves
digitate 5. Herrania.
Petals with a short spatulate appendage; leaves entire.
6. Theobroma.
Ovary (as fruit) tuberculate or spinescent.
Anthers 3 in each stamen-tube sinus 7. Guazuma.
Anthers 1 in each stamen-tube sinus.
Petals smooth or with stiped gland; fruit tubercled,
3. Ayenia.
Petals appendaged; fruit spinescent 4. Buettneria.
Petals withering, persisting.
Ovary 5-celled 1. Melochia.
Ovary 1-celled 2. WaUheria.
Flowers unisexual or polygamous, apetalous; leaves ample, divided
or lobate 9. Sterculia.
1. MELOCHIA L.
Shrubs, herbs or trees, stellate pubescent or sometimes with
simple trichomes but these not or rarely intermixed, stipulate
serrate leaves and usually small flowers variously disposed in the
axils or terminal. Calyx campanulate, rarely inflated, 5-dentate or
parted. Petals 5, marcescent, plane, opposite the 5 more or less
connate stamens, staminodia none or minute. Ovary 5-celled, the
cells 2-ovuled; styles 5, free or united medially. Capsules 5-valved,
the cells sometimes separating completely as 1 (2) -seeded cocci.
M. hirsuta Turcz., Bull. Soc. Nat. Mosc. 36, pt. 1: 570. 1863,
based on Mathews 1552 without data, is essentially a nomen nudum
the few lines of description without diagnostic characters, but
possibly it is M. tomentosa; it is a pity that there is not a general
revision of the group as will be apparent from the following synopsis
of the Peruvian species.
Leaves subrotund or in any case rounded apically, at most about
3 cm. long.
624 Field Museum of Natural History— Botany, Vol. XIII
i
Stems slender, trailing; capsules pyramidal M. crenata.
Stems stout, erect; capsules apparently globose. . . .M. peruviana. ,
Leaves longer than broad or in any case mostly or all much larger. *
Flowers all in dense usually sessile axillary clusters; capsules
depressed-globose M. nodiflora.
Flowers variously disposed other than above.
Inflorescence (at least principal) an interrupted terminal spike
of dense sessile purplish flowers; capsules 5-lobed globes,
the cocci bivalved M. villosa.
Inflorescences various, often axillary, sessile or peduncled cymes
or corymbs or if terminal more or less paniculate or in-
frequently spiciform.
Flowers purplish, mostly pedicelled, cymose-umbellate; cap-
sules pyramidal.
Calyces to 4 mm. long, cymes rather opposite the often
glabrate leaves M. pyramidata.
Calyces 5-6 mm. long; cymes in the axils of the canescent
(beneath) leaves M. tomentosa.
Flowers yellow or white (unknown, M. pseudonodiflora),
usually many, often subsessile; capsules like 5-lobed
globes.
Fruiting calyx papyraceous; petioles slender, 1-2 cm. long;
flowers slender-pedicelled in axillary clusters or on
axillary branchlets M. lupulina.
Fruiting calyx firm; petioles stout, the upper rarely 1 cm.
long except M. pseudonodiflora; flowers in panicles or
often peduncled corymbs.
Leaves more or less pilose above with mostly simple
trichomes; flowers about 6 mm. long in rather
irregular or open inflorescences M. pilosa.
Leaves stellate-pilose both sides; flowers about 5 mm.
long, at least in part in dense corymbs or ample
panicles.
Leaves serrulate, shortly petioled; inflorescences sub-
corymbose M. leucantha.
Leaves coarsely serrate, slender-petioled; flowers,
except lower, in an ample panicle.
M. pseudonodiflora.
Flora of Peru 626
Melochia crenata Vahl, Symb. Bot. 3: 86. pi. 68. 1794. M.
Chamaedrys St. Hil. Fl. Bras. Merid. 1 : 161. 1825, at least as to Peru?
Trailing suflfrutescent, the elongating pilose stems almost filiform
and with 1-3 (5)-flowered cymules opposite the roundish crenate-
serrate leaves, these 1.5-3 cm. long, glabrate (Peru) above, pilose
beneath; stipules ovate-subulate; calyx 4 mm. long, the lobes
subulate; petals 5-6 mm. long; stamens about medially connate, the
styles basally; capsules pyramidal, the 5 angles rounded, about 1
cm. long, with 2 seeds in each cell. — Petals rose colored with brown
veins (Weberbauer). F.M. Negs. 21602; 9600 (M. Chamaedrys).
There may be two species but the negatives do not suggest it.
M. hermannioides St. Hil. is similar but the more numerous flowers
are subcapitate.
Cajamarca: I*rov. Ja^n, Weberbauer 6220 (det. Dahlem, M.
Chamaedrys). Colombia; Jamaica.
Melochia leucantha Macbr., sp. nov.
Fruticosa 4-8 dm. alta; ramulis, petiolis, foliisque (imprimis
subtus) stellulato-puberulentis; stipulis ovato-lanceolatis, ad 3 cm.
longis; petiolis 5 (-7) mm. longis; foliis late ovatis interdum fere
subrotundatis, basi truncatis vel obscure cordulatis, ad apicem late
acuminatis vel rotundatis apice ipso obtusis, plerumque 3.5-4 cm.
longis, 3-3.5 cm. latis, serratis, nervis lateralibus subtus conspicuis
supra paullo notatis; inflorescentiis subterminalibus mediocriter
contractis circa 5-8 cm. latis et longis, pedunculis 2-5 cm. longis,
ramulis 5-10 mm. longis, pedicellis 1-3 mm. longis; sepalis sub-
sericeo-pilosis, ovatis, longe acuminatis, firmo-chartaceis, 5-7 mm.
longis, petalis paullo brevioribus; fructibus ut videtur depresso-
globosis. — As noted by Cuatrecasas in herb, this is not M. globifera
Tr. & Planch, of Colombia (as determined by Standley) which is
well-named ; the entire inflorescence in the Peruvian shrub is a broad
pseudo-corymb. Noted by the Goodspeed collectors as conspicuous
in full flower, the corollas white or white with yellow markings at
base; the other specimens cited are placed here on the basis of their
reference to the Colombian species and may belong to some other
species.
Huancavelica: Gravelly shrub-lands, 2,400 meters, east of Mejo-
rada, Goodspeed Exped. 10908, type. — Apurimac: Shrub-land, 2,000
meters, Rio Pachuchaca, Goodspeed Exped. 10546.
Melochia lupulina Swartz, Prodr. 97. 1788. Mougeotia in-
fiata HBK. Nov. Gen. & Sp. 5: 330. pi. 484. 1823.
626 Field Museum of Natural History— Botany, Vol. XIII
Often a meter or more high and suffnitescent below, the slender
virgate branches early stellate puberulent; stipules and petioles
slender; leaves ovate, sometimes broadly, serrate, subcordate to
rounded-truncate at base, acute or short-acuminate, soon glabrate
or persistently a little pubescent beneath, membranous, commonly
5-8 cm. long and more than half as wide; flowers slender pedicelled
but densely clustered in the leaf -axils or on axillary branchlets or in
corymbs, these sometimes pedunculate; petals white, yellow at
center, little longer than the calyx, this about 4.5 mm. long, in
flower soon inflated-accrescent, membranous and pale in fruit;
filaments medially united; capsules about 3 mm. high, puberulent,
the cocci opening slightly at separation along the inner angle. —
Illustrated, Fawcett & Rendle, Fl. Jam. 5: 163 (flowers and fruit).
F.M. Neg. 35380 (M. inflata).
Cajamarca: Cascas, Raimondi (det. Ulbrich). Tambillo, Jelski
282 (det. Szyszyl., M. inflata). — Piura: Raimondi (det. Ulbrich). —
Libertad: Trujillo, Killip & Smith 21501 (det. Standley). — Junin:
Colonia Peren^, Killip & Smith 2506 Jf. (det. Standley). — Huanuco:
Near Huanuco, 2060. — Lima: Chorillos and Miraflores, Martinet;
Raimondi. San Lorenzo near Callao, Andersson. Chosica, ^96.
Near Lima, Ball. — Loreto: Yurimaguas, Poeppig 2^25; Spruce 3898.
To Central America; West Indies.
Melochia nodifiora Swartz, Prodr. 97. 1788.
Resembles M. lupulina but the dense usually sessile clusters with ,
conspicuous bracts, the calyces not at all or little accrescent in '
fruit; flowers pink or rose-striped; filaments completely connate; '
capsules depressed-globose.
Peru (undoubtedly). Bolivia; Colombia to Mexico; West Indies.
Melochia peruviana Desr. in Lam. Encycl. 4: 83. 1797(?);
cf. Journ. Bot. 4, 4: 318. 1906.
Tomentose in all parts, the lignescent branches about 1 dm.
long; leaves oval, obtuse or acutish, serrate, very shortly petioled;
stipules lanceolate, caducous; flowers axillary, solitary (-5), citron
yellow, the peduncle about as long as petioles; calyx segments
lanceolate, acute, slightly shorter than the oblong-ovate clawed _^
petals; ovary globose, tomentose, the 5 reddish styles shorter than
the basally united stamens. — After Lamarck, who gives the type X;
locahty as "Peru." Perhaps a part of M. depressa Mill, of Cuba, as ,
illustrated by Cav., Diss. 6: pi. 173, fig. 1, but the Cuban plant
Flora of Peru 627
noted by Lamarck as having larger leaves, roseate flowers, probably
different fruit (depressed, pentagonous), by Grisebach as similar to
M. venosa. The petioles are 3-7 mm. long, leaves 1-3 cm. long and
about as wide, densely white stellate-tomentose especially beneath;
flowers about 5, axillary-fasciculate, white with a yellow center, the
clusters shortly peduncled. F.M. Neg. 29771.
Hudnuco: Near Hudnuco, 2S2Jt^; Ruiz & Pavdn; Dombey, type.
Melochia pilosa (Mill.) Fawcett & Rendle, Fl. Jam. 5: 164.
1926. Sida pilosa Mill. Gard. Diet. ed. 8. 1768. M. venosa Swartz,
Prodr. 97. 1788. Mougeotia polystachya HBK. Nov. Gen. & Sp. 5:
328. pis. Jt83, Wa. 1823, fide Fawcett & Rendle. Melochia poly-
stachya (HBK.) Tr. & PI. Ann. Sci. Nat. s^r. 4, 17: 341. 1862. M.
venosa Swartz var. betonicaefolia [R. & P.l Schum. in Mart. Fl. Bras.
12, pt. 3:37. 1886.
Pilose-sericeous shrub sometimes 1 meter tall with notably
nervose leaves especially beneath and narrow or sometimes branched
terminal and axillary panicles; stipules linear-lanceolate, 7-8 mm.
long; petioles rather stout, usually about 1 cm. long; leaves typically
about 4-8 cm. long, about half as wide, apparently sometimes much
larger; calyx 5 mm. long, the lobes acuminate; petals 6.5-8 mm.
long, yellow, apparently sometimes white; filaments completely
connate; capsule depressed-globose, villous, the finally separating
cocci splitting apically by 2 short valves. F.M. Negs. 9609 (var.);
23866 (var.).
Cajamarca: Valley of the Tabaconas, Weberhaiier 62H- Ja^n,
Weberbaim 6200.— San Martin: Tarapoto, Williams 5802; 6685.—
Hudnuco: Cuchero, Poeppig 1269. Below Rio Santo Domingo, I^212.
Tingo Maria, Soukup 22^6. Near Pozuzo, Jt761?—C\xzco: Prov.
Convencion, Soukup 911; Vargas 793. Lares Valley, Prov. Galea,
Weberbauer 7932 (det. Ulbrich). Echarate, Goodspeed Ezped. 10506
(det Standley). West Indies; Tropical South America.
Melochia pseudonodiflora Hochr. Ann. Gons. Jard. Bot.
Geneve 21: 430. 1920.
Ferrugineous tomentose or pilose, especially toward the top;
petioles to 14 mm. long; leaves ovate, mostly a little cordate,
coarsely serrate, to 3.5 cm. long, about 1.5-2.5 cm. wide, stellate-
pilose both sides, basal nerves 3-5, villous, prominent beneath;
flowers 1-3 in axils of reduced leaves, the uppermost forming an
ample panicle, the pedicels 2-3 mm. long; calyx stellate-pilose,
628 Field Museum of Natural History— Botany, Vol. XIII
glabrescent within at base, about 5 mm. long in fruit, the lobes half
as long, the obovate petals little longer, inserted slightly above the
short subcylindrical stamen column this bearing apically the
scarcely applanate filaments; styles free; capsules globose, sulcate,
pubescent, the 5 carpels separating and dehiscing loculicidally.
— Much resembles M. nodiflora but leaves a little thicker, serrations
better marked and, especially, lacking the inflated stamen column
with broad short filaments, the petals not decurrent on its base
(author).
Peru(?) : Without locality, Ruiz [& Pavdn], type.
Melochia pyramidata L. Sp. PI. 674. 1753. Sida Mathewsii
Turcz. Bull. Soc. Nat. Mosc. 36, pt. 1: 565. 1863, fide Baker, Journ.
Bot. 30. 1892.
Slender-stemmed, suffrutescent at least below, rarely 1 meter
tall, usually glabrate or the ovate or oblong-lanceolate leaves not
infrequently (at least in Peru) more or less cinerous stellate pu-
bescent; cymes sessile or pedunculate, opposite the slender petioles,
usually few (-10) -flowered; calyx 3.5-4 mm. long, the lobes lanceo-
late-subulate, the purplish-pink petals twice as long; capsules 5-6
mm. high, somewhat broader, acutely angled. — S. Mathewsii not seen.
Cajamarca: Cascas, Raimondi (det. Ulbrich). — San Martin:
Tarapoto, Williams 58^0 (det. Standley). — Junin: Colonia Peren^,
Esposto (det. Ulbrich). Chanchamayo Valley, Schunke 1537. La
Merced, 5312. Uspachaca, 1310. — Lima: Near Lima, Savatier ^.05.
— Loreto: Rio Paranapura, Klug 3927 (det Standley). Widely
distributed in the warmer regions of the New World and naturalized
elsewhere.
Melochia tomentosa L. Syst. Nat. ed. 10. 1140. 1759. M.
Turpiniana HBK. Nov. Gen. & Sp. 5: 323. pi. U82. 1823.
Shrubby, more or less canescent with a close stellate indument
the broadly ovate to oblongish leaves most densely so beneath;
flowers in lax or dense axillary or terminal cymes mostly obviously
pedicelled; calyx 5-6 mm. long, the narrow lobes acuminate the
usually purplish petals about twice as long; capsules broadly pyra-
midal, rostrate, the angles more or less rounded. — F.M. Negs.
23865 (var.); 35386 {M. Turpiniana).
Peru (possibly). To southern North America; West Indies.
Melochia villosa (Mill.) Fawcett «fe Rendle, Fl. Jam. 5: 165.
1926. Sida villosa Mill. Card. Diet. ed. 8. 1768. M. hirsuta Cav.
Flora of Peru 629
Diss. 6: 323. pi. 175. 1788. M. vestita Benth. in Hook. Journ. Bot.
4: 130. 1842, fide Uittien.
Well-marked by the dense sericeous villous pubescence usually
spreading on the long younger branches that terminate in spiciform
inflorescences composed of dense sessile or subsessile clusters, these
bracteolate and sometimes bracted by reduced leaves; calyx 4 mm.
long, the violet petals nearly twice as long; capsule globose, the 5
cocci finally completely separating and dehiscing. — In the specimen
of Cavanilles by Jussieu the flowers are in a globose head and
axillary in sessile fascicles, probably a young state. F.M. Negs.
35388 (M. vestita Benth.); 35389 {M. hirsuta).
Cajamarca: Sucse River Valley, Socota, Stork & Horton 10102?
Fence rows, Chota, Stork & Horton 100^6 (young; det. Standley,
M. glohifera). — San Martin: Tarapoto, Williams 5798; 61 U9. San
Roque, WiUiams 7102. Zepelacio, Klug 3705. To Mexico and the
West Indies.
2. WALTHERIA L.
Resembles Melochia but indument always entirely stellate, ovary
1-celled, the capsule bivalved. — The flowers are densely glomerate,
the leaves velvety tomentose in the Peruvian species, at least
beneath.
Leaves usually ovate-rotund, subequally yellowish tomentose both
sides; filaments free above W. ovata.
Leaves usually ovate-oblong or ovate-elliptic, less whitish tomentose
above; filaments completely united W. americana.
Waltherla americana L. Sp. PI. 673. 1753. W. indica L., I.e.
W. erioclada DC. Prodr. 1: 493. 1824.
More or less suffrutescent and canescent with a tomentose
indument that extends to the sessile or pedunculate bracted axillary
and terminal globose inflorescence, least dense on the upper surface
of the leaves; petioles often only 5-10 mm. long; leaves generally
ovate-oblong or -lanceolate or subelliptic, rounded or subcordate
at base, obtuse or slightly acute, usually 3-6 cm. long, serrate;
calyx 4-5 mm. long, the lobes lanceolate-subulate, little exceeded
by the bright yellow petals; style not longer than the connate fila-
ments; capsules 2 mm. long.— F.M. Negs. 8002; 23849 {W. erioclada).
Arequipa: Mejia, (Guenther & Buchiien 189, det. Bruns). —
Apurlmac: (Weberhaner 5877, det. Ulbrich). — Cuzco: Savanna,
630 Field Museum op Natural History— Botany, Vol. XIII
Prov. Convencion, Weherbauer 79 J/^ (det. Ulbrich); Raimondi.
New World tropics and naturalized in the Old.
Waltheria ovata Cav. Diss. 6: 317. pi. 171. 1788. W. reticulata
Hook. f. Trans. Linn. Soc. 20: 231. 1847. W. sericea Turcz. Bull.
Soc. Nat. Mosc. 31, pt. 1: 214. 1858, fide Svenson.
Prostrate or bushy and erect (Svenson), sometimes 2 meters
tall, equally covered by a felt-like yellowish-white indument; leaves
commonly cordate-ovate, rarely linear-oblong, rounded at tip, more
or less obviously reticulate veined especially beneath, often 3-6
cm. long, 2.5-4.5 cm. wide; flowers yellow, mostly in several sessile
clusters often borne along together on short axillary branchlets;
petals 5 mm. long, exceeding the densely pubescent calyx; style
longer than the partly free filaments. — W. incana R. & P. ined. is
the same. The sulphur yellow color of the leaves is characteristic
(Svenson). Illustrated, Svenson, Amer. Joum. Bot. 22: 244. pi. 3.
Piura: Payta, Gaudichaud; D'Urville. Negritos, H aught 3.
Lower Parifias Valley, Haught 118. — ^Ancash: Ocros, Weherbauer
2653. — Huanuco: Chulki, Sawada 48. Huacho, Stork & Horton
9U12. Near Hudnuco, 20^3; Kanehira 267; River o; Ruiz & Pav6n.
— Lima: Amancaes, Andri. Chosica, 55^. Near Lima, Soukup;
Raimondi. Callao, Domhey; Jussieu, type; Gaudichaud. Surco,
Martinet. San Bartolom^, Savatier Ii.92. — Ayacucho: Rio de Lomas,
Weherbauer 57^5 (det. Standley). Ecuador; Galapagos. "Mem-
brillejo" (Raimondi).
3. AYENIA L.
Much like Guazuma but subherbaceous or more or less suffrute-
scent, the calyx 5-parted, the slender-clawed petals cucullate and
more or less cleft at apex and, especially, the anthers (sometimes
3-celled) solitary in the sinuses of the very short staminal tube, the
ovary cells only 2-ovuled. — Besides the following, A. magna L. may
occur; it is a shrub with ovate-cordate acuminate leaves soft-
pubescent beneath, 5-9 cm. long, 4-6 cm. wide and perhaps (Svenson,
Amer. Joum. Bot. 33: 469. 1946) has been found in adjacent Ecuador.
Named for the Due d'Ayen.
Leaves acutely acuminate, the coarse serrations strongly unequal.
A. serrata.
Leaves obtuse or acute, the small serrations irregular but subequal.
Stems erect, the ascending branches villous A. erecta.
Flora of Peru 681
Stems as branches diffuse, glabrate, retrorsely puberulent or
tomentulose A. piisilla.
Ayenia erecta Mart. Fl. Bras. 12, pt. 3: 103, pi. 23, fig. 1. 1886.
Erect from fibrous root, the stem simple or with many spreading-
ascending densely leafy branches villous with spreading trichomes,
to 2 dm. high; petioles 5-10 mm. long; stipules filiform, 1 mm. long;
leaves oblong-elliptic, acutish or obtuse, narrowly cordate at base,
2-3 cm. long, 14-17 mm. wide, the lower and uppermost much
smaller, irregularly serrate, sparsely appressed pubescent above,
finely villous beneath; pedicels usually in threes, filiform, equaling
the calyx, this 2 mm. long, the lanceolate 1-nerved acuminate
segments hirsute without, subglandular within; petals 3 mm. long,
red, the capillaceous claw purple; gynophore 3 times longer than
stamen tube, this 0.7 mm. long; capsules depressed globose, finely
echinate with fragile spines, the seeds deeply and irregularly foveo-
late. — Apparently ex char, and illustration, I.e., distinctive, and
presumably determination correct but originally from Bahia, Rio
San Francisco near Joazeiro.
Cajamarca: Callacate, (Jelski 321, det. Szyszylowski). Brazil.
Ayenia pusilla L. Syst. Nat. ed. 10: 1247. 1759.
Subdecumbent-ascending or the many stems or branches sub-
erect, sometimes several dm. tall, the sparse to numerous trichomes
short, recurved; leaves usually basally ovate- or elliptic-oblong,
ordinarily 1-1.5 (4) cm. long, rarely more than 1.5 cm. wide, closely
serrulate from subtruncate or openly cordate base to the apiculate
truncate or acutish tip; flowers reddish-purple, solitary or 2-3-
fasciculate, calyx deeply and acutely lobed, 2-3 mm. long, the
petals with slightly longer capillaceous claw and roundish bilobed
limb; ovary long-stipitate; capsules depressed-globose, 3 mm. long,
4-5 mm. broad, pubescent or glabrate, more or less short-echinate.
—Illustrated, Mart. Fl. Bras. 12, pt. 3: pi. 2U.
Lima: Ambar to Huacho, Stork 1H68 (det. Johnston). — Hudn-
uco: Near Hudnuco, Ruiz & Pavdn; Kanehira 225; Soukup 2227.
Ambo, 3169. — Huancavelica: Colcabamba, Weberbauer SHI- —
Cuzco: Cunyac, Prov. Anta, 2,050 meters, Vargas 9^30. Widely
distributed.
Ayenia serrata Ruiz, in herb.
Fruticosa ad 5 dm. alta ramulis junioribus subherbaceis dense
retrorseque puberulis; petiolis gracilibus circa 1.5 cm. longis; foliis
632 Field Museum of Natural History— Botany, Vol. XIII
ovatis, plerumque circa 5 cm. longis, 2.5 cm. latis, basi rotundatis,
acute acuminatis, irregulariter serratis, membranaceis, glabris vel
subtus sparse hispidulis; pedicellis filiformibus 3-5 mm. longis,
3-5-fasciculatis; sepalis petalisque ut videtur albidis vel petalis
purpureis, tenue unguiculatis, exsertis; capsulis ignotis. — The
original material probably collected by Tafalla as labeled from
Guayaquil and noted in herb, by Mildbraed as apparently a new
undescribed species. J. R. Lawrence has labeled the Peruvian
collections as new, using a name in reference to the irregular ser-
rations, the teeth being of two sizes, but the publication not found.
Tumbez: Rainy green formation, 150 meters, Hacienda La
Choza, Weberbauer 7701; also Hacienda Chicama, 600 meters,
Weberbauer 7672. Ecuador.
4. BUETTNERIA Loefl.
Reference: Schumann in Mart. Fl. Bras. 12, pt. 3: 83-101. 1886.
Suffrutescent and erect or sometimes scandent or sarmentose,
often aculeate, the leaves various, the small purplish or greenish-
white or yellowish flowers more or less pedicellate in umbels or now
and then in axillary or subterminal sessile or peduncled cymes.
Calyx deeply 5-parted, the 5 concave inflexed petals with long
linear terminal and 2 small lateral appendages. Stamen tube and
petals connate, with 5 sessile 2- or 3-celled anthers. Ovary 5-celled,
the cells 2-ovuled; style 5-parted. Capsule globose, echinate or
spiny, with partly dehiscing cocci. — There are usually one or more
glands on the medial and lateral leaf -nerves beneath; the name has
been spelled variously but as in Luehea the original may be followed
except to indicate the umlaut by the diphthong. Basic characters
of the many species are probably in the flowers and fruits but these
are unknown in detail for many species; my key therefore is doubtless
burdened, for expediency, with characters of doubtful value.
Peduncles and pedicels subfiliform or slender, soon either or both
longer than petioles or usually about 1 cm. long; fruits except
as noted, unknown; leaves often more or less pubescent beneath.
Leaves about oblong to ovate, sometimes broadly ovate or obovate
but obviously longer than wide or small; stems usually more
or less aculeate.
Indument if present more or less villous-hirsute; sepals not
subfiliform-acuminate.
Flora of Peru 633
Plants unless inflorescences glabrous or glabrate; leaves often
entire; fruits unknown except B. glabrescens.
Leaves about oblong-elliptic, entire or nearly, shortly
acuminate or acute.
Leaves ample, 4-9 cm. wide, acuminate.
B. myriantha, B. coriacea.
Leaves small, about 2 cm. wide, acute. . .B. rhamnifolia.
Leaves ovate, long-acuminate, serrate; fruits long-aculeate.
B. glabrescens.
Plants densely pubescent, at least the serrate leaves beneath;
fruits echinate, spines a few mm. long.
Flowers white or yellowish; ovate-lanceolate leaves soon
glabrate or glabrous above.
Branches aculeate; trichomes not or little crisped.
B. parviflora, B. cordata.
Branchlets not aculeate (type); trichomes crisped.
B. eriogona.
Flowers red; cordate-ovate leaves tardily glabrate above.
B. hirsuta.
Indument minutely stellate on upper stems, scurf y-tomentose;
sepals subfiliform-acuminate B. urosepala.
Leaves ample, subrotund to broadly ovate, but little longer than
wide; stems unarmed B. pescapraeifolia, B. catalpaefolia.
Peduncles relatively stout, usually shorter than petioles or scarcely
1 cm. long unless in fruit, the pedicels often shorter; fruits
unknown, B. ancistrodonta, B. aurantiaca, B. benensis, B.
peruviana, B. eriogona; species glabrescent except B. divaricata,
B. discolor.
Petioles subtending flowers about 5 mm. long; leaves never
lepidote-stellate, stems (t3T)es) armed, not densely pubescent.
Leaves obtuse or barely acute; fruits long-aculeate B. ovata.
Leaves obviously acute or acuminate.
Leaves rugosely nerved beneath, acute or very shortly
acuminate; fruits short-echinate B. Weberbaueri.
Leaves finely nerved, acuminate; fruits long-aculeate (known).
Leaves ovate-lanceolate, about 2 cm. wide, serrate toward
tips; stems terete or subquadrate only in age.
B. tereticaulis.
634 Field Museum of Natural History— Botany, Vol. XIII
Leaves if ovate broader, usually about elliptic; stems more
or less angled or leaves entire or subentire.
Leaves often distinctly 2-3-serrate below the caudate
acumen, notably reticulate; petal appendage in-
flated B. ancistrodonta.
Leaves entire or subentire, not very reticulate; petal
appendage compressed or little dilated.
Leaves oblong-elliptic; petal appendage subfleshy.
B. aurantiaca.
Leaves ovate; petal appendage little dilated.
B. peruviana.
Petioles subtending flowers about 1 cm. long or longer or if
shorter leaves pilose or lepidote-stellate beneath and stems
unarmed.
Leaves glabrous or glabrate, mature at base acute, rounded or
subcordate; fruits long-aculeate (known).
Stems not aculeate (types).
Leaves long-acuminate, abruptly coarsely serrate.
B. benensis.
Leaves short-caudate, serrulate B. boliviana.
Stems aculeate; leaves short-acuminate.
Leaves somewhat serrate toward tips; peduncles and
pedicels subequal B. aculeata.
Leaves quite entire; peduncles often a little longer than
short pedicels B. Tessmannii, B. acuminata.
Leaves softly pubescent beneath; fruits short-echinate.
B. divaricata.
Leaves rufescent beneath with a compact minute indument.
B. discolor.
Buettneria aculeata Jacq. Sel. Stirp. Amer. Hist. 76. 1763.
B. carthaginensis Jacq. Sel. Stirp. Amer. Pict. 41. 1780. B. lateralis
Presl, Rel. Haenk. 2: 144. 1835.
Shrubby, the hollow angled stems or branches armed with]
recurved prickles and usually tangled or scandent; leaves rounded-|
ovate to oblong-lanceolate, very variable in size, ordinarily mem-
branous and crenate or serrate toward the tip, glabrous to rarel]
densely pilose- tomentose beneath; flowers almost minute, brown<j
purple or green, the sessile or often pedunculate and umbellifoi
Flora of Peru 686
cymes few-flowered; fruit body 7-10 mm. broad, glabrous or densely
pubescent, armed with many long slender or stout hard spines. —
Synonymy and description after Standley and Steyermark, Fieldi-
ana: Bot. 24, pt. 6: 406. 1949, who remark that "no good basis for
separating any of the Central American forms is apparent." This
situation seems to exist in the case of one or more Peruvian species,
which perhaps ought to be included in this variable (?) entity, but
the problem is one for a student who can acquaint himself with the
plant of Jacquin.
Peru (possibly). To Mexico.
Buettneria acuminata Bredem. ex Roem. & Schult. Syst 5:
470. 1819.
Stems terete or nearly, glabrous, the prickles 2-3 mm. long;
petioles to 1 cm. long; leaves ovate or oblong, entire, rounded or
crenate-rounded at base, long-acuminate, to 5 cm. long, half as wide,
puberulent only beneath at base and in nerve axils; inflorescence
consisting of 1-3 2-3-flowered umbels, the puberulent peduncles
and glabrous pedicels to 7 mm. long; calyx 3 mm. long, the lanceolate
acuminate segments 2 mm. long with some longer trichomes at the
angles; petals 4 mm. long, claw 1 mm. long, obovate-lanceolate, the
lateral processes triangular, ligule linear, subclavate apically, puberu-
lent only basally; staminodia 3-dentate, puberulent at tip; capsules
8 mm. in diameter, spines 1.5 cm. long, the black seeds with purple
chalaza and raphe, 5 mm. long, 2 mm. thick. — After Schumann.
Collectors of Peruvian material have noted the flowers as rose and
cream colored, greenish-yellow or -white. Tamayo 275 from the
type region, Venezuela, a well-prepared specimen seen at the Uni-
versity of California, seems to match the Peruvian material, but the
relationship of the ubiquitous (presumably) B. aculeata is open to
question. It pleases me greatly to record here the cooperation given
me by Herbert Mason and the staff of the University Herbarium —
a great collection effectively ordered and maintained, constantly
important in the preparation of this work. F.M. Neg. 9805 (Brede-
meyer), which was not seen until after preparation of this account.
San Martin: Chazuta, Klug 2987; m7 (both det. Standley).—
Junln: Puerto Yessup, Killip & Smith 26293 (det. Mildbraed). La
Merced, 55P^ .^—Loreto: Yurimaguas, Poeppig 2360; Spruce 3900
(both det. Schumann). Balsapuerto, Klug 2987 (det. Standley).
Venezuela.
636 Field Museum of Natural History — Botany, Vol. XIII
Buettneria ancistrodonta Mildbr. Notizbl. Bot. Gart. Berlin
11:142.1931.
Glabrous, the branchlets and leaf -nerves aculeate; petioles
scarcely 5 mm. long; leaves elliptic-lanceolate, obtuse at base,
subcaudate-acuminate, 6-11 cm. long, 2-4.5 cm. wide (or to 18 cm.
long, 8 cm. wide), toward apex with 1-3 crenate serrations, the 6-8
lateral nerves and densely reticulate veins prominent; pedicels very
short; sepals ovate-lanceolate, acute, 4-4.5 mm. long; petal claw
broadly linear, cucullate, little dilated, the lateral processes fleshy,
clavate, divaricate, 1 mm. long, the narrowly ovoid fleshy appendix
1.4 mm. long, obtuse; staminodia 2 mm. long, the hamate middle
tooth much longer than the wing-like lateral. — Extraordinary in the
inflated petal appendage and the form of the elongate staminodia
teeth (author); flowers dark brown to carmine (Tessmann). The
Puerto Lim6n specimen was at one time referred to B. coriacea.
F.M. Neg. 17935.
Loreto: Soledad on the lower Itaya, Tessmann 529^, type.
Puerto Limon on the Marafion, Tessmann 3861. Rio Mazdn,
Schunke 8j^. Mishuyacu near Iquitos, Klug 1593 (det. Standley,
B. acuminata).
Buettneria aurantiaca Mildbr. Notizbl. Bot. Gart. Berlin
11:141.1931.
Branchlets shortly aculeate, the younger as the petioles, these to
1 cm. long, sparsely puberulent with crisped trichomes; leaves
oblong-elliptic to lanceolate, rounded and more or less cordate at
base, obtusely acuminate, entire or irregularly and distantly crenate-
dentate above, only the younger both sides on the midnerve with
some scattered trichomes, basally 3- or obsoletely 5-nerved, the 6
or so lateral nerves with transverse veins prominent beneath;
peduncles about 2 mm. long, pedicels little longer; sepals ovate-lan-
ceolate, acutely acuminate, scarcely 3 mm. long, very sparsely pilosu-
lous; petal claw scarcely 1 mm. long, lateral processes rather semi-
lunate, semifleshy, stout appendix compressed, obtusish, 1-8 mm.
long, glabrous, orange; medial staminodia tooth very broad, obtuse.
— The species is recognizable by the orange lanceolate somewhat
fleshy petal appendix (Mildbraed). F.M. Neg. 17936.
Loreto: Flood-free wood, Soledad, Tessmann 5292, type.
Buettneria benensis Britton, Bull. Torrey Club 16: 155. 1889.
Scandent, unarmed, glabrous except for a few scattered stellate
trichomes on the branches and petioles, these 3-4 cm. long, and
Flora of Peru 687
a minute indument on the leaves beneath, these strongly cordate,
ovate, sagittate or ovate-lanceolate, long-acuminate, serrate, dis-
tinctly nerved and reticulate veined, 8-13 cm. long, 5-7 cm. wide;
peduncles short; calyx hirsute, the lanceolate sepals 5 mm. long. —
Resembles B. cordifolia Sagot of French Guiana but smooth (Brit-
ton), and in view of the variation in this respect of B. glabrescens
and doubtless other species, it may indeed be a part of Sagot's
species; t3rpe from junction of Beni and Madre de Dios rivers,
Bolivia, and thus to be expected within Peru.
Peru (probably). Bolivia.
Buettneria boliviana Britton, Bull. Torrey Club 16: 155. 1889.
Glabrescent, unarmed liana with cordate ovate acuminate
serrulate leaves 6-8 cm. long, 4-5 cm. wide, reticulate veined both
sides, the petioles slender; capsules 1-5 cm. long, 2 cm. in diameter,
the spines 2-3 mm. long; seeds 5 mm. long, 2 mm. broad, glabrous,
black. — With B. benensis, of which, ex char., it seems to be the
fruiting state, but actually, as indicated in the key, it must be
distinct; however, types of neither of these were seen until after
preparation of this compilation.
Peru (probably). Bolivia.
Buettneria catalpaefolia Jacq. Hort. Schoenbr. 1: 21, pi. ^6.
1797; 91.
Unarmed finally nearly glabrous liana, the stems terete, the
petioles slender, several cm. long, the leaves subrotund or basally
ovate, usually a dm. or two long, acutely acuminate, sometimes
especially the younger minutely or floccosely stellulate pubescent
beneath and with 5 prominent nerves from the subtruncate or more
or less cordate base; cymes axillary or terminal, lax, pedunculate,
often ample; flowers white; sepals 5-6 mm. long; petals 10 mm. long,
claw 2 mm. long, lateral processes minute, ligule 8 mm. long, finely
acuminate; fruit body globose, 2.5-3.5 cm. in diameter, the slender
prickles sharp; seeds 8 mm. long, 6 mm. in diameter, reddish with
yellowish raphe, purple chalaza. — Illustrated, Mart. Fl. Bras. 12,
pt. 3: pi. 20.
Cajamarca: Above Ocros, stream banks, Weberbauer, 163. —
San Martin: Zepelacio, Kliig 3386. Juanjui, Klug ^2^2. Tarapoto,
Spruce Jt956. — Hudnuco: Chicoplaya, Ruiz & Pav6n (det. Mild-
braed). — Loreto: Balsapuerto, Klu^ 2857; 291^9. Yurimaguas,
Poeppig 2262; KiUip & Smith 28351. To southern Mexico.
638 Field Museum of Natural History— Botany, Vol. XIII
Buettneria cordata Lam. Encycl. 1: 522. 1785. B. sulcata
R. & P. Fl. Peruv. 3: 10. 1802? B. corchorifolia Turcz. Bull. Soc.
Nat. Mosc. 25, pt. 2: 152. 1852.
Stems diffuse, pilosulous, more or less angled and sulcate, aculeate
(as rarely petioles and leaf midnerve beneath); petioles to about
1 cm. long; leaves rather densely short-pilose beneath especially
on the 3-5 basal nerves, finely serrate base to often obliquely
acuminate apex, glabrous or nearly above, cordate-ovate, usually
3.5-5 cm. long, 1.5-2.5 cm. wide (on vigorous shoots to three times
as large), the upper often bractiform; peduncles and pedicels
filiform, about 1 cm. long, the latter shorter, often somewhat nutant;
calyx segments narrowly lanceolate-acuminate, spreading, about
3 mm, long, the filiform-appendaged petals to nearly twice as long;
fruits rotund, about 1 cm, in diameter, the short spines 2-3 mm.
long; seeds obtusely trigonous, rostrate, scabrous-punctate. — Schu-
mann suggested that B. sulcata could be an earlier name for B.
acuminata Bredem. and I did not find the type at Madrid; however,
it seems ex char, to be a part of B. cordata, the principal differences
noted by the authors being the aculeate petioles and leaf midnerve,
characters which of course at once suggest also B. tereticaulis, but
that is otherwise different; also the locality is in the center of dis-
tribution for B. cordata. F,M, Negs. 9579; 29763; 23838 (B. cor-
chorifolia).
Lima: Quive, Prov. Canta, (Ruiz & Pav6n, type, B. sulcata).
Near Lima, Dombey, type; Wawra 259 U. Santa Rosa, Martinet.
Chancay and Huarai, Ruiz & Pavdn. On Rio Santa Eulalia, 1,700
meters, Horton 10992 (det. Johnston, B. parviflora); at 1,400 meters,
Goodspeed & Stork 114^83 (det, Johnston, B. hirsuta). River bottom,
Chosica, 2863. Ecuador. "Yerba de la Arana" (Ruiz & Pavon).
Buettneria coriacea Britton, Bull. Torrey Club 16: 156. 1889.
Glabrous except for many-flowered inflorescences; branches
terete, sparsely aculeate; petioles stout, 1 cm. long; leaves oblong-
lanceolate, 12-14 cm. long, 4-5 cm. wide, coriaceous, prominently
nerved beneath; pedicels and bracts pubescent; sepals tomentose,
ovate, acute, 2 mm. long. — Mildbraed, Notizbl, Bot. Gart. Berlin
11: 141. 1931, has added the following from the Tessmann col-
lections: petioles 5 mm. long, rugulose; leaves papyraceous, nar-
rowly ovate or ovate-lanceolate, rounded or obtuse at base, gradu-
ally long-acuminate, to 7 cm. wide, lateral nerves 8 or 9, the densely
reticulate veins more conspicuous above than below; peduncles
Flora of Peru 639
very slender, 1 cm. long, the pedicels scarcely half as long; sepals
lanceolate, acutely acuminate, 4 mm. long; petals 3 mm. long,
claw 1.5 mm. long, slender lateral processes little deflexed, fleshy
appendage dorsally compressed, especially marginally, ciliate,
staminodia hardly 1 mm. high; ovary tuberculate. — Well-marked
by the black-red (dried) petals with fleshy lanceolate brown ciliate
appendage (Mildbraed); liana, to 10 meters, 5 cm. in diameter,
much branched (Tessmann). Type from junction of the rivers Beni
and Madre de Dios, Bolivia, peduncles 6-8, pedicels 3-4 mm. long;
veins lax, same both sides.
Loreto: Soledad, Tessmann 5291. Pongo de Manseriche, Tess-
mann JtlOJt. Leticia, Williams 3171 (det. Mildbraed). Florida,
Klug 2301; 2310; 2330 (both distr. as B. ancistrodonta?) . Bolivia.
Buettneria discolor Benth. in Hook. Joum. Bot. & Kew Misc.
3:165. 1851; 92.
High-climbing, unarmed, the branchlet tips ferrugineous tomen-
tulose as the younger petioles, these finally 1-2 cm. long; leaves
oval or some oblong-lanceolate, rounded to subcordate at base, very
shortly acuminate, acute or obtusish, mostly about 1 dm. long (-1.5),
7 cm. wide, coriaceous, glabrous and drying brownish red above,
minutely lepidote-stellulate beneath; inflorescence of 3-5 umbels,
3-6-flowered, the puberulent peduncles and pedicels to 5 mm.
long; calyx tomentulose only without, 4 mm. long, the segments
ovate, acute; petals purple, 4 mm. long, claw obcuneate, the lateral
processes little incurved, fleshy, as the lanceolate acuminate ligule;
staminodia emarginate, carinate; anthers barely free; capsules 1.5
cm. in diameter or larger, densely aculeate with stout spines about
2 mm. long.— Illustrated, Mart. Fl. Bras., 12, pt. 3: pi. 21.
Loreto: Rio Mazdn, JosS Schunke 166 (det. Cuatrecasas). Ama-
zonian Brazil.
Buettneria divaricata Benth. in Hook. Joum. Bot. 4: 124.
1842; 100.
Younger scandent divaricate branches minutely aculeate,
tomentulose; petioles 6-15 mm. long; leaves orbicular-ovate to
broadly oblong-lanceolate, cordate to truncate at base, serrulate,
5-10 cm. long, half as wide, scabriusculous above, tomentulose and
8-5-glandular on nerves near base beneath; peduncles stout, short,
bearing 4-5 umbels of 5-7 flowers; pedicels puberulent; calyx 5 mm.
long, the lanceolate segments acuminate, hirsute without and within
640 Field Museum of Natural History — Botany, Vol. XIII
at tip; petals 5 mm. long, cucullate, the lateral processes horizontal,
triangular, rounded at apex, the thick ligule puberulent at acuminate
tip, dilated at base; staminodia acute, serrate, stamens sessile, the
anthers approximate; pistil tubercled; capsules 1.5 cm. in diameter,
tomentose, the approximate conical prickles 3 mm. long. — After
Schumann. Has been collected as near as Solimoes. F.M. Negs.
19637; 35435.
Peru (probably). To British Guiana.
Buettneria eriogona Mildbr. Notizbl. Bot. Gart. Berlin 11:
144. 1931.
Puberulent-tomentulose with crisped simple trichomes especially
the younger branchlets, petioles, these to 12 mm. long, and leaves
beneath the branches soon glabrate except the obvious angles,
unarmed; leaves cordate-ovate, acutely acuminate, 4-6 cm. long,
3^.5 cm. wide, minutely crenulate, the crenations apiculate, 5
(-7) -nerved from base with 3-4 additional lateral nerves; inflo-
rescences or peduncles about 1 cm. long; sepals free nearly to base,
ovate-lanceolate, 3 mm. long; claw of petals strongly incurved, the
thin linear lateral processes erect-divaricate, appendix filiform,
about 5 mm. long; staminodia short, broadly retuse. — The species
is characterized by the indument and by the erect-spreading ap-
pendages of the petals (Mildbraed); it is, I think, a part of B.
cordata Lam. F.M. Neg. 17938.
Lambayeque: Hacienda de Chinama, Raimondi ^11, type.
Buettneria glabrescens Benth. Bot. Voy. Sulphur 71. 1844.
Stems (these rarely aculeate, Svenson) and younger leaves
puberulent, the older glabrate, ovate-lanceolate, rounded or rareljr
subcordate at base, acuminate, serrate, mostly 5-10 cm. long, at
least half as wide or upper narrower; peduncles 12-25 mm. long,
6-10-flowered, the buds long-acuminate; sepals 3 (-5) mm. long,
narrowly lanceolate, sometimes long-acuminate; petal appendages
filiform; fruit long-aculeate. — AfRne B. cordata but glabrate and
seems distinct by form of leaves (Bentham). Tjrpe from Guayaquil;
the stems are marked by 5 light colored ridges, semi-woody, in
clumps to moi*e than 2 meters high, rarely show any trace of spinel
(Svenson; see illustration, .Amer. Joum. Bot. 33: 476, fig. 1). The
southern Peru specimens have larger flowers, shorter peduncles and
may be at least varietally distinct; both referred by Standley to
B. cordata but glabrous and the fruits long-aculeate. See also
B. Dielsii mentioned under B. parviflora.
Flora of Peru 641
Piura: Amotape Hills, (HaugfU & Svenson 115S8). Farinas
Valley, Haughl 71; 168 (distr. as B. parviflora), — Apurlmac: Oropeza
Valley, 2,700 meters, Vargas 9771— Cuzco: Collcha-Pururo, 2,820
meters, Vargas 9736. Ecuador.
Buettneria hirsuta R. & P. Fl. Peniv. 3: 10. 1802; 99.
Stems densely hirsute, much branched, the 5 angles strongly
aculeate, the branches less so as the petioles, these 2.5 cm. long;
leaves cordate, acute, crenate, 7-nerved, the nerves somewhat
aculeate, the indument more softly hirsute beneath than above;
peduncles to 3 times as long as petioles, filiform, many-flowered;
pedicels umbellate, capillary, unequal; petals lutescent the lower
half, purple the upper; otherwise like B. sulcata (Ruiz & Pav6n);
and to this original description Schumann has added: leaves ovate,
10-13 cm. long, 6-9 cm. wide, scabriusculous above or early orange
sericeous, the medial and lateral nerves glandular; peduncles to
1.5 cm. long; calyx 5.5 mm. long, hirsute as the lanceolate acuminate
segments within; petals 7 mm. long, cucullus 1 mm. long, linear, the
lateral processes rotund, glabrous as the filiform ligule; staminodia
emarginate, stamens stipitate, anthers free, capsules 16 mm. in
diameter with short conical spines. — The leaves of the type are
velvety in the manner of those of the more northern B. mollis
HBK. which may not be distinct. P.M. Neg. 23841.
Hudnuco: Pillao, Ruiz & Pavdn, type. Yanano, 3799 (det.
Mildbraed). Mufla, 3926. — Arequipa: Challascapa near Mejia,
{Guenther & Buchtien -4, det. Bruns). To southeastern Brazil.
Buettneria myriantha Schum. & Ulbr. Verb. Bot. Ver.
Brandenb. 50: 91. 1908.
High climbing glabrous liana, the terete branches with a few
spines often 3-4 mm. long and 5 mm. wide at base; petioles 6-8
mm. long; leaves oblong-oval, obtuse at base, acuminate (perhaps
not always), entire, 1.5-2 dm. long, 7-9 cm. wide, coriaceous, the
nerves prominent both sides; flowers about 10, the peduncles 1-1.5
cm. long, the slender pedicels 5 mm. long, both pubescent as the
lanceolate sepals, these 2-3 mm. long, basally carinate; ovary
densely verruculose. — Perhaps allied to the puberulent B. discolor
Benth. (authors), but ex char, seems to be a part of B. coriacea.
F.M. Neg. 23842.
Peru (probably). Brazil.
642 Field Museum of Natural History — Botany, Vol. XIII
Buettneria ovata Lam. Encycl. 1: 522. 1785.
Shrub, a meter or two (sometimes 4 meters) high, the 5-angled
glabrous branches aculeate on the angles; petioles smooth; leaves
ovate, glabrous, serrate, about 2.5 cm. long; flowers axillary, in
corymbs or umbels of 3-6, the peduncles and pedicels short; sepals
acute; petals trifid at purple apex; capsules spheroid, echinate with
five 1-seeded cells. — Lamarck; type grown from seeds collected by
Joseph de Jussieu in "Peru," probably rather from the vicinity of
Quito (Mildbraed), who cited a number of collections from central
Ecuador, Bibl. Bot. 116: 106. 1937. The Peruvian specimen is
abundantly aculeate; the leaves of type oval, barely acutish, 2-3
cm. long, 12-15 mm. wide, pinnately nerved above 3 basal nerves
these barbate in the axils, sometimes minutely scabrous on nerves,
the capsule 1 cm. in diameter, the rather few spines to 7 mm. long.
The branches are acutely angled, stout. F.M. Negs. 8003; 29764;
35439.
Libertad: Potaz, Raimondi 512 (det. Diels). Ecuador. "China-
cacha" (Jussieu).
Buettneria parviflora Benth. PI. Hartw. 114. 1843.
Branchlets aculeate, puberulent, soon glabrate; leaves ovate
(or the smaller floral lanceolate), rounded or scarcely cordate at
base, acuminate, 5-7.5 cm. long, 3-5 cm. wide, acutely serrate,
tomentose beneath and the midnerve glandular; peduncles fasci-
culate toward the tip of the branchlets in the axils of bracteiform
leaves, 8-10 mm. long, bearing umbels of 6-10 flowers; calyx and
petal segments ovate-lanceolate, the latter 2 mm. long, acute,
puberulent, with divaricate auriculate lateral processes and filiform
appendages twice as long as the calyx, scarcely or not enlarged above
the middle; spines of fruit short-conical (Bentham). — Included as
some Peruvian specimens referred by me to B. cor data or B. gla-
brescens have been so determined; however the species is not clearly
distinguishable from the former. The apparently well-marked
B. obtusata Benth. ex Hochr. Ann. Cons. Jard. Bot. Geneve 11-12:
5. 1908, of Ecuador, has subrotund leaves, rounded even at apex,
canescent pilose beneath. B. Dielsii Mildbr. Bibl. Bot. 116: 106.
1937, is very near according to the author but branchlets glabrous
or merely puberulent, flowers 5-6 mm. long.
Peru (cf. note above). Ecuador.
Buettneria peruviana Turcz. Bull. Soc. Nat. Mosc. 36, pt. 1^
569. 1863.
Flora of Peru 648
Branches definitely scandent, aculeate, finely puberulent; leaves
ovate, subcordate, obtusely acuminate, entire, glabrous and puncti-
culate above, rarely pubescent beneath at base of the 5 nerves, and
with a depressed oblong gland on midnerve; pedicels of the few-
flowered umbels scarcely longer than the short peduncles; petal
appendages little dilated. — Seems to be near B. hrevipes Benth. of
southern Ecuador but the petal appendages not enlarged into an
ovate-oblong claw and inflorescences not geminate (author); the
leaves of Bentham's species are ovate-lanceolate, subacute at the
rounded base. I am indebted to Annetta Carter of the University
of California Herbarium for finding this description given in Index
Kewensis as "31: 207. 1858."
Amazonas(?): Without data (Mathews 1652, type).
Buettneria pescapraeifolia Britton, Bull. Torrey Club 16:
155. 1889.
Glabrous, the slender petioles not aculeate, subequaling the
leaves, these orbicular or basally ovate, cordate, obtuse or acute,
entire, 8-12 cm. wide, dark green with impressed nerves above,
glaucous and prominently nerved beneath; capsules semiglobose,
2 cm. in diameter, very densely muricate. — After the author, who
allied it to B. discolor.
Junin: Satipo, Seibert 2373 (det. Killip & Idrobo). Bolivia.
Buettneria rhamnifolia Benth. in Hook. Joum. Bot. & Kew
Misc. 3:164. 1851; 96.
Branches erect glabrous except for the puberulent tips and with
a few recurved prickles 5 mm. long; petioles distichous, to 8 mm.
long, stout, canaliculate above; leaves oblong-elliptic or -lanceolate,
rounded at base, acute to subemarginate, 6 cm. long, 22 mm. wide,
coriaceous, entire, glabrous above, puberulent in axils of the nerves
beneath, unarmed; inflorescences sometimes in axillary branchlets
of 1-3 4-6-flowered umbels, the glabrous peduncles to 1.5 cm. long,
the pedicels about as long; calyx glabrous, nearly 4.5 mm. long, the
linear-lanceolate long-acuminate sepals soon reflexing; petals dark
purple, 8 mm. long, cucullus 1 mm. long, linear, dilated at tip
the glabrous wings erect, the ligule long-conical, puberulent; stami-
nodia 3-dentate, anthers free. — The type of B. Weberbaueri was
referred here by Ulbrich. F.M. Neg. 9592.
Peru (probably). Amazonian Brazil.
644 Field Museum of Natural History— Botany, Vol. XIII
Buettneria tereticaulis Lam. Encycl. 1: 523. 1785; 95.
Branches cylindrical, a little pubescent, sparsely aculeate; leaves
ovate-lanceolate, scarcely rounded at base, acuminate, about 5 cm.
long, scarcely 18 mm. wide; capsules of other species but small and
much more densely echinate with divaricate spines 10-12 mm. long.
— Lamarck, the type from Peru by Joseph de Jussieu. Schumann
suggested that this seems to be near if not the same as B. sulcata
(B. acuminata); in the type the leaves are serrate toward the tip
and with a little pilosity at juncture with petiole but not between
the basal nerves and the petioles, these 5 mm. on fruiting branch, as
the young stems pilose only in a line; nervation pinnate. The stems
are quite terete; no specimen from Peru has leaves so narrowly
lanceolate but it could be an extreme form of B. aculeata.
Peru: Without locality, Joseph de Jussieu, type.
Buettneria Tessmannii Mildbr. Bot. Gart. Berlin 11: 140.
1931.
Glabrous unless for some puberulence on the younger branchlets,
irregularly aculeate, the petioles, these 1-2 cm. long, smooth or with
1 or 2 prickles; older branches 5-sulcate, the internodes 3-5 cm. long;
leaves ovate-elliptic or elliptic, rarely broadly lanceolate, obtuse to
shortly cuneate at base, acutely acuminate, 7-12 cm. long, 3.5-6
cm. wide, thin-papyraceous, 3-nerved at base, the lateral nerves
(other than basal) 3-4 curved-ascending to the entire margins;
peduncles scarcely 1 cm. long, pedicels 2-3 mm. long, the few flowers
greenish; sepals 5 mm. long, the triangular acute segments 3 mm.
long; claw of petals 1.5 mm. long, basally 1 mm. wide, lateral
processes spreading, obtuse, appendix filiform-attenuate, 3 mm.
long, little clavate-enlarged; staminodia with acute short hamate
middle tooth; capsules (immature) with many prickles 8 mm. long.
— No observation by author; it seems to be doubtfully distinct from
B. acuminata or B. aculeata or the specimens cited under the former
may belong here. F.M. Neg. 17939.
Loreto: Mouth of the Santiago, Tessmann Jf027, type.
Buettneria urosepala Mildbr. Notizbl. Bot. Gart. Berlin 11:
144. 1931.
Indument minutely stellulate, on the deeply 5-sulcate shortly
aculeate branchlets fulvous-tomentulose; petioles stout, about 5
mm. long;, leaves ovate-lanceolate, rounded at base, rarely slightly
cordate, gradually acuminate, entire (or remotely and minutely
\
Flora of Peru 645
serrulate), 8 cm. long, half as wide, the younger minutely stellate,
punctate above, beneath with some trichomes on the 3-5 prominent
basal and about 6 lateral nerves, the densely reticulate veins more
conspicuous above; pedicels scarcely 1 cm. long; sepals little connate,
lanceolate-ovate, subfiliform-acuminate, 4 mm. long, stellate-pilose;
petal claw dilated at base, the thin lateral lobes deflexed, the filiform
appendix hardly 5 mm. long; staminodia truncate and excised,
barely 1 mm. long. — Without observation by the author. F.M.
Neg. 17940.
Junin: Vitoc, Raimondi, type. Huacapistana, Sandeman W7
(distr. as B. eriogona).
Buettneria Weberbaueri Mildbr. Notizbl. Bot. Gart. Berlin
11:145. 1931.
A shrub 2-3 meters high, the glabrous more or less clearly
5-angulate branchlets armed with strong short recurved prickles,
the leaves especially toward the acute tip very acutely but minutely
serrate; petioles 5-10 mm. long, crisply pilosulous; leaves often
ovate-lanceolate, little cordate, 2.5-5 cm. long, 1.5-3 cm. wide,
basally 5-nerved, barbate in the axils of the 6-7 lateral nerves,
prominent beneath; pedicels nearly as long as petioles; flowers
yellowish-green; sepals glabrous, about 6 mm. long, acuminate;
petal claw obovate-spathulate, cucullate, short lateral processes
deflexed, filiform appendage 6 mm. long; staminodia hardly 1 mm.
high, the minute middle tooth affixed below the retuse margin;
capsules globose-ovoid, 1.5 cm. long, 1 cm. in diameter, rather
densely armed with prickles 2~i mm. long. — Readily known by the
small but very sharp teeth of the leaves (Mildbraed). F.M. Neg.
17941.
Cajamarca: Above Chota, 2,600 meters, Weberbauer U219, type
(det. Ulbrich, B. rhamnifolia) ; Stork & Horton 10038 (det. Standley,
B. cordata).
5. HERRANIA Goudot
Reference: Richard Evans Schultes, Caldasia 2, no. 6: 11-26.
1943; 2, no. 9: 325-336. 1944.
Like Theobroma but notably marked by the digitately divided
leaves and, especially, by the extraordinarily elongate ligule that
exceeds the petal base many times, the aestivation cincinnate-
involute. — Treated as a section of Theobroma by Schumann and
646 Field Museum of Natural History— Botany, Vol. XIII
obviously a part of no other genus; nevertheless it constitutes
a soundly defined separate group, most conveniently bearing its
own name after a General of Colombia (Herran) who became
President and, according to the author, consolidated the peace,
leading his country into prosperity.
Besides the following, H. breviligulata R. E. Schultes, Caldasia
20: 21. 1942; Bot. Mus. Leafl. Harvard 16: 78, 207. pi. 12. 1953,
from Colombia may be found in Peru; it has softly velvety stellate
abruptly acuminate fruits, not at all indented at the peduncle, this
2 cm. long.
Leaves glabrous or essentially, rarely asperous H. nitida.
Leaves more or less pubescent, at least beneath.
Fruits pubescent, with stinging trichomes.
Sepals equal, entire.
Indument stellate-pilose; leaflets moderately attenuate to
base H. Mariae.
Indument soft, dense; leaflets long-attenuate to base.
H. Cuatrecasana.
Sepals unequal, the outer often incised H. kofanorum.
Fruits velvety pilose, with no stinging trichomes; sepals equal,
entire H. nyderodendron.
Herrania Cuatrecasana Garcla-Barriga, Caldasia 1, no. 2:
57, pi. 2. 1941.
Similar to H. Mariae but, as to type, much smaller, to 3 meters
tall with pale maculate bark, indument much softer and denser, and,
especially, the leaflets much longer attenuate at base, the lateral
about 33 cm. long, 9 cm. wide, the central 5 dm. long, 17 cm. wide,
above sparsely stellate, the veins tomentose, beneath stellate-pilose,
the veins more prominent; pedicels stellulate-tomentulose, 2-3 cm.
long, with a basal linear-filiform, pilose bract; staminodes con-
spicuous, purple, rhomboid-lanceolate, attenuate to the obtuse or
acute apex, 2.5 cm. long, 8 mm. wide; fruit ellipsoid, acuminate, the
10 ribs covered with stinging stellate trichomes, rind thin, brittle,
yellow at maturity. — After Schultes, Caldasia 2, no. 9: 329. 1944,
who has described fruits found by him and concludes that while
vegetatively the species appears to be related to H. nycterodendron
"comparison of the fruits reveals that they are rather distinct."
Illustrated, Schultes, I.e. 330 (photograph, fruits).
Peru (no doubt). Adjacent Colombia.
Flora of Peru 647
Herrania kofanorum Schultes, Bot. Mus. Leafl. Harvard 14:
126. 1950.
Habit of other species; trunk to 7 cm. in diameter, the bark
scrobiculate and scabrous; branches and branchlets tomentose,
finally glabrate; stipules caducous, linear, 2.5-3 cm. long, hispidu-
lous without; petioles stout, obscurely sulcate, densely tomentose,
to 3 dm. long; leaflets sessile, notably and regularly undulate-
sinuate, lanceolate-oblong, gradually attenuate to base, acute or
mucronate acumen to 2 cm. long, coriaceous, unequal but mostly
17-30 cm. long, 6-11 cm. wide, glabrous or evanescently and
minutely puberulent above or the veins fusco-hirsute, beneath
densely stellate-pilose the veins ferrugineous tomentose; fascicles
of flowers on lower trunk, the stout densely stellate-puberulent
pedicels about 1 cm. long, the stellate-pilose oblong-globose buds
18 mm. in diameter; sepals strongly unequal, sparsely stellate-
strigose without, the rotund-ovate outer one about 1.5 cm. long and
broad, often incised for 2 mm. the incision extending as a furrow
within to sepal-base, the inner 2 elliptic, acute, 1 cm. long, 6 mm.
wide; petals 4 or 5, red-purple, muricate-papillose, 9 mm. long, 7
mm. wide, the filiform ligule much dilated at granular base, to
1 dm. long; staminodes coarsely papillose both sides, acute, 22 mm.
long, 6 mm. wide; ovary distinctly 10-costate, densely stellate-pilose;
pistil simple; fruit said to be yellow. — After Schultes who compares
it to H. balaensis Preuss of Ecuador with subequal sepals all entire.
Illustrated, Schultes, I.e. pi. 28 (habit photograph of flowers);
pl.SU.
Peru (probably). Adjacent Colombia and Ecuador. "Ko-kee-
ot'-chu" (Kofdn, Schultes).
Herrania Mariae (Mart.) Decaisne ex Gk)udot, Ann. Sci. Nat.
s^r. 3. 2: 233. 1844. Abroma Mariae Mart. Denkschr. Bot. Ges.
Regensb. 3: 297. pis. 8, 9. 1841. Theobroma Mariae (Mart) Schum.
in Mart. Fl. Bras. 12, pt. 3: 71. 1886.
Rather puberulent-tomentulose especially the leaves beneath
including the long (4-5 dm.) basally enlarged petioles and the
2-5-parted roseate calyx (12 mm. long) without, this with basally
elliptic, obtuse, subcymbiform segments sparsely pubescent, within
the simple trichomes purplish; leaflets 7-9, obovate-rhomboid, the
lateral more or less oblique and curved, all attenuate to sessile base,
acuminate, herbaceous, light green; inflorescences fasciculate on
the trunk, the peduncles to 16 mm. long, squamately bracted at
648 Field Museum of Natural History— Botany, Vol. XIII
base, filiform as the pedicels; petal base 6 mm. long, nearly as wide,
yellowish with 5 purple nerves, the ligule 1 dm. long, 1.2 mm. wide,
with 3-5 purple nerves; stamen tube dark red, usually 3-antherifer-
ous, the granulate apically serrate staminodia 10.5 mm. long; styled
tomentose, the stigmas 5. — After Schumann and said to attain:
9 meters with trunk diameter to 3 dm., the beautiful large leaves;
forming a subglobose crown. The var. putumayonis Schultes, Botj
Mus. Leafl. Harvard 14: 129. 1950, differs in leaflets to 5.2 dm. long^
1.8 dm. wide, lanceolate-elliptic, globose flower buds to 17 mm. in'
diameter, ligules often shorter; type several trunks in a clump.
Dedicated to Queen Mary of Saxony for her interest in botany .^
Illustrated, Schultes, I.e. pi. 30 (habit photograph, flowers anc
fruits); Mart. Fl. Bras. I.e. pi. 15.
Loreto: Rio Putumayo between Rios Igaraparana and Yaguas^
(Schultes JfOlO, type, var.). Amazonian Brazil. "Mu-se-naM
(Witoto, Schultes), "cacohuilla" (WilHams).
Herrania nitida (Poeppig) Schultes, Caldasia 2, no. 6: 16. 1943^
Ahroma nitida Poeppig, in Poepp. & Endl., Nov. (Jen. & Sp. 3: 73J
1845. Theohroma nitida (Poeppig) Schumann in Mart. Fl. Bras. 12j
pt. 3: 72. 1886. H. atrorubens Huber, Bull. Soc. Bot. Geneve, s^rJ
2, 6: 187. 1914, fide Schultes, Bot. Mus. Leafl. Harvard, 16: 80j
1953.
Slender tree, the large 7-9-foliolate leaves crowded at the apei
of the strict trunk, this usually only a few cm. in diameter at base^
stipules linear, to 2.5 cm. long, 1 mm. wide, usually stellate-setulosej
petioles enlarged at base, to 4 cm. long, appressed stellulate-tomen-
tulose; leaflets sessile or nearly, lanceolate-oblong, attenuate
base, acuminate, the largest middle ones 2.5-4.5 dm. long, 7-1^
cm. wide, the often asymmetrical much smaller, firm-chartaceous oi
subcoriaceous, entire or undulate-margined, glabrous and lustrous
above, rarely evanescently on the nerves beneath; flowers in fas-
cicles of as many as 30-40 on the lower part of the trunk; pedicels
articulate, densely appressed tomentulose with a few strigose setae,|
about 5 mm. long, outer sepal yellow-red, stellate-setose withoutj
rotund-ovate, about 9 mm. wide, the elliptic acute inner 2 to 6 mm^
wide, twice as long; petal-base broadly obovate, 4-5 mm. wide,j
5-6 mm. long, cucullate, most densely muricate without, yellowish-*
red or roseate, terminating in a pendent coiled pink or scarlet
purple-veined linear ligule to 8 cm. long; stamen-tube alternately
2 and 4 antheriferous, the conspicuous staminodes 9 mm. long.
Flora of Peru 649
wide; stigma simple; fruit ovoid or ellipsoid, acuminate, to
[1 cm. long, 5 cm. thick, the 5 primary and 5 secondary thin cutel-
liform ribs minutely and sparsely hispid with stinging trichomes;
seeds subtriangular, 9 mm. long, 8 mm. wide, 5 mm. thick. — To
4 meters tall, the dark or black thin bark smooth or rough. After
Schultes. Forma sphenophylla Schultes, Bot. Mus. Leafl. Harvard
14: 131. 1950, has narrowly lanceolate-elliptic leaflets, often fewer
flowers; var. aspera (Karsten & Triana) Schultes, I.e. 130, of adjacent
Colombia has asperous leaves, entirely blood-red purple flowers.
Determinations by Schultes. F.M. Negs. 32206; 32204 (var. aspera).
Illustrated, Schultes, I.e. (Caldasia), 17 (photograph of trunk with
flowers).
San Martin: Tocache, Poeppig 1979, tjT^e. — Loreto: Rio Mazdn,
JosS Schunke U5 (t5rpe, forma). Iquitos, Killip & Smith 27Jlt31.
Caballo-Cocha, WiUiaim 2332. On the Amazon, Williams 28It3.
Yurimaguas to Balsapuerto, KiUip & Smith 2823It. Rio Itaya,
WiUiams 33Jt5; SS6U. Peru-Colombia boundary, Williams 281^3;
2816 (var. aspera). Adjacent Colombia. "Cacahuilo" (Killip &
Smith), "cacahuio" (Schunke).
Herrania nycterodendron Schultes, Caldasia 2, no. 6: 21. 1943.
Usually simple-stemmed little tree sometimes 4.5 meters tall
but the trunk less than a dm. in diameter at base, the ample 7-
foliolate leaves crowded at the rarely branched apex; stipules
caducous, linear, to 4 cm. long, 3 mm. wide, tomentose as the stout
terete very obscurely sulcate petioles, these to about 6 dm. long, and
the slender pedicels (2-2.5 cm. long) and linear bracts; leaflets
sessile, lanceolate-oblong, gradually long-attenuate to base, more or
less acutely acuminate (acumen to 2 cm. long), entire or subundulate,
the largest to 6 dm. long, 2 dm. wide, firm-chartaceous or papy-
raceous, somewhat lustrous above, glabrate or sparsely hirsutulous,
the veins minutely tomentulose, these densely ferrugineous tomen-
tose beneath, the interspaces stellate-pilose; flower fascicles on the
lower trunk to 45-flowered; sepals entire, stellulate-pilosulous
without, puberulent within, the outer one rotund-obovate, 19 mm.
long, 15 mm. wide, the inner two elliptic, about 15 mm. long and
half as wide; petals basally rotund, cucullate, muricate-papillose,
pale ochre-purple, 10 mm. long, 8-9 mm. wide, the pendulous
membranous ligule filiform, to 1 dm. long, minutely granular only
at base, deep purple with prominent black nerves; staminodia
reflexed, lanceolate-elliptic, 19-21 nmi. long, 7-9 mm. wide, muricate-
650 Field Museum of Natural History — Botany, Vol. XIII
granular; ovary pilose, the glabrous style deeply 5-parted; fruit
ellipsoid, 10-12 cm. long, 4-5 cm. thick, long-attenuate above and
often constricted below the acute or frequently rounded-obtuse tip,
indented at the ligneous peduncle (this 4-5 cm. long), minutely
but densely velvety stellate-pilose (no stinging trichomes), very
deeply 10-costate, the alternate of the broad rounded ribs smaller,
the rind hard, fibrous, ashy yellow at maturity, the seeds 9x8x8
mm., the white pulp edible. — After Schultes, who remarks "immedi-
ately set apart from the most closely related species H. Mariae and
H. Cuatrecasana by its curious type of fruit." With a happy imagin-
ation he has translated into Greek for the species name the native
one at Remanso, "Palo de murci^lago," i.e. bat-tree, probably in
allusion to the resemblance of the clustered velvety fruits. Illus-
trated, Schultes, I.e. 22 (photographs of fruits) ; 26 (drawing of fruit) ;
Bot. Mus. Leafl. Harvard 14: pi. 35.
Loreto: Remanso, Rio Putumayo, Schultes UOll, type. Florida,
flowers wine-red, Klug 2069. Mishuyacu near Iquitos, Klug 1588.
Colombia. "Palo de murci^lago," "palo de chimbe," "cacao de
murci^lago," "mu-se'-na," "mu-se'-ge-ke" (last two Huitoto, all by
Schultes).
6. THEOBROMA L.
By Josfi CUATRECASAS
Cacao [Toumef.] Aublet.
References: G. Bernoulli, Uebersicht der bis jetzt bekannten
Arten von Theobroma, Neue Denkschrift allg. Schweiz. Gesellsch.
fuer d. g. Naturwiss., 24, 3: 1-15, pis. 1-7. 1871; K. Schum. in
Mart. Fl. Bras. 12, pt. 3: 70-77. 1886; A. Chevalier, Revision du
genre Theobroma, Revue Intern, de Bot. Appl. et d'Agr. Trop., 26:
265-285. 1946.
Tree with usually spreading branches and simple coriaceous,
nodding or pendulous, distichous leaves. Stipules small or medium
sized, geminate, lateral, deciduous. Leaves penninerved, base often
3-5-nerved. Inflorescence cauline or axillary, cymose, few- or
many-flowered. Flowers hermaphrodite, small or medium sized,
pentamerous. Calyx 3-5-parted, lobes ovate or triangular, valvate.
Petals in aestivation valvate, formed by two articulate parts, the
basal one strongly hooded, the superior part laminar, petaloid like
an appendage of variable shape, usually oblong or suborbicular.
Stamens 5 fertile, alternate with 5 staminodia, all connate at the
base, forming a short urceolate tube; the fertile ones opposite the
Flora of Peru 651
petals with a short curved filament, at apex 2-3-parte(i and bearing
two or three bilobate anthers; each extrorse anther hidden in the
petal concavity and its two lobes divaricate; the staminodia petaloid,
broadly or narrowly laminar, acute or obtuse. Ovary sessile,
globose, ovoid, or pentagonal, 5-celled with axile placentation and
many-ovulate cells; style simple or more or less 5-parted; stigmas
simple, acute. Ovule anatropous with dorsal rhaphe and 2 integu-
ments. Fruit usually large, subbaccate, indehiscent, ovoid or
oblong, smooth, rugose or 5-10-costate, tomentose, velvety or gla-
brous; pericarp thick or thin, hard-coriaceous or woody or firmly
fleshy, the meso- and endocarp at maturity a pulp enclosing the
seeds in a single cavity. Seeds medium sized, ovoid but laterally
more or less flatly compressed, integuments filmy, embryo straight,
radicle conic, short, inferior, the cotyledons thick lobulate-corrugate;
endosperm none or reduced to a mucilage within the rugosities of
the cotyledons.
A neotropical genus with 18-20 species distributed in the rain
forests of South and Central America, many of them localized
endemisms, others very much spread, mostly through cultivation.
The seeds of Theobroma are appreciated as food and a stimulating
beverage (cacao, cocoa, chocolate), and one of its species (T. cacao
and varieties) constitutes one of the most important economical
tropical crops. See C. J. J. Van Hall, Cacao (Macmillan & Co. Ltd.,
London, 1932). The name Theobroma, given by Linnaeus, is derived
from the Greek, meaning "food of the Gods" (Theos=God, broma=
food).
Stamens biantheriferous.
Inflorescences on the trunk and old branches.
Leaves flexible-coriaceous, glabrous, penninerved; pericarp
fleshy T. ca^ao.
Leaves rigid-coriaceous, with minutely stellate appressed in-
dument beneath, basal pair of nerves ascendant, distant
from the next pair; pericarp coriaceous T. calodesmis.
Inflorescences axillary; leaves thin-coriaceous, cinereous- tomen-
tose beneath, 5-7-nerved; pericarp ellipsoid, thick, woody,
10-ridged and deeply reticulate-sulcate T. bicolor.
Stamens triantheriferous.
Inflorescences on the trunk and old branches.
Leaves cuneate, trinerved at base; petal ligule subrotundate;
fruit subpentagonal, sulcate, acuminate, imibilicate, about
10 X 5 cm T. calodesmis.
652 Field Museum of Natural History — Botany, Vol. XIII
Leaves at base rotundate, extremely asymmetic, usually 5-
nerved; petal ligule spathulate-cuneate; fruit elliptical,
smooth 10 X 8 cm. scarcely sinuate-sulcate.
T. speciosum.
Inflorescences axillary.
Staminodes petaloid, spathulate; leaves pergaminaceous, pale,
dense indument cinereous, monotrichous beneath; fruit
elliptic-obovate; pericarp coriaceous, tuberculate.
T. obovatum.
Staminodes petaloid, lanceolate; leaves thick-coriaceous with
strong nerves; fruits smooth.
Leaves appressed tomentose, monotrichous beneath, nerves
glabrous; fruits large, about 20 cm. long; flowers large.
T. grandiflorum.
Leaves densely tomentose, heterotrichous beneath, with
a ground of cinereous, appressed minute stellate hairs
and larger reddish stellate more sparse hairs on the
nerves; fruits smaller, about 10 cm. long; flowers medium
sized T. suhincanum.
Theobroma bicolor Humb. et Bonpl. Aeq. 1: 104, pi 30. 1808.
Usually a small tree (3-6 meters) which can attain 18 meters,
with an erect stem and narrow crown; young branchlets, often
horizontal or pendulous, appressed cinereous tomentose by very
minute stellate hairs; leaves thin-coriaceous, usually pendulous;
petioles 1.5-3.5 cm. long, erect or curved, appressed tomentose;
blade ovate-oblong, cordiform at the base, 5-7-palmate-nerved,
acute or acuminate, the edges smooth, 1.5-4 dm. long, 6-23 cm.
broad, glabrous and green on the upper side, densely and compressed
tomentose, lightly cinereous, with minute stellate hairs beneath;
principal nerves little conspicuous above and very prominent be-
neath, the midrib a little thicker with usually 3 secondary curved-
ascending nerves at each side, the transversal nerves separated,
prominent beneath, the small veins rather prominently reticulate;
flowers in small axillary cjnnes equaling or exceeding a little the
petiole; branchlets thin, shortly tomentose and with small linear
bracts; pedicels 3-6 mm. long; sepals 5, ovate-lanceolate, 5-6 mm.
long, 2-2.5 mm. wide, united at the base, reddish appressed-tomen-
tose without; petal-base obovate, 2.2 mm. long, rotundate and
hooded at the apex, the hgule subsessile, ovate-subrotundate, 1-1.2
mm. in diameter, puberulent without, red or purple; staminal tube
Flora op Peru 658
1 mm. high, the staminodes purplish oblong, obtuse-lanceolate,
obtuse or subacute at apex, densely papillose-tomentose, 3.5-4 mm.
long, 0.8-1 mm. broad, the fertile filaments 1 mm. long, bent,
bearing 2 anthers, each bicellular; ovary 5-sulcate, densely hirsute-
tomentose, subobovoid, 1.8 mm. long, the style glabrous; fruit
subglobose-ellipsoid or ovoid, 1.5-2 dm. long, 10-12 cm. in diameter,
green-yellowish with green nerves, the exocarp thin, a little velvety,
the pericarp hard, lignose, 5-8 mm. thick, strongly reticulate with
5 prominent ribs, the pulp yellowish, sweetish; seeds more or less
compressed, ovoid, 20-27 nmi. long, 15-18 mm. broad.
The Williams specimens were erroneously listed by him as T.
quinquenernum Bern. (Woods of Northeastern Peru, Field Mus.
Bot. 15: 324. 1936). Origin uncertain but probably Central Ameri-
can. It is widely but not extensively planted from southern Mexico
through Central America to the Amazonian basin; the seeds are
locally used to make chocolate but are recognized as inferior in
quality to those of T. cacao L. The pulp is also edible and is used
locally to prepare beverages. Its peculiar fruit, elliptic and lignous,
and the leaves, which are whitish, somewhat silvery below, make
this species easily recognizable, and it has been given many different
names in other countries: Central America: "pataiste," "pataste,"
"patashte," "patasht," "cacao pataxte," "cacao bianco," "cacao de
monte." Colombia: "bacao." Ecuador: "patas," "cacao bianco."
Brazil: "cabu-assu," "cupuassu," "cupuacu," "cacao do Peru,"
"cacao baju," "cacao d'anta."
Loreto: Florida, Rio Putumayo at mouth of Rfo Zubineta,
clearing, Klug 2021. Mishuyacu, near Iquitos, clearing, Klug 1528.
Paraiso, Alto Rio Itaya, 145 meters, Williams S3U6. Caballo-cocha,
on the Amazon River, Williams 21^9. — Also Weberhav£r 62j^5 (with-
out locality, at Chicago), "macambo" (Klug, Williams), "majambu"
(Williams SSU6).
Theobroma cacao L. Sp. PI. 782. 1753.
Tree, usually 5-8 meters high, branches in clusters mostly of
5, spreading, gray, glabrous, the very young ones pubescent; leaves
obovate-oblong or oblong, rotundate or obtuse at the base, abruptly
acuminate or cuspidate at the apex, glabrous or slightly pubescent
beneath, the nerves pinnate, the midnerve very strong beneath but
only filiform above, secondary nerves 10-14 at each side, spreading,
curved-ascending, slender and prominent beneath, the other veins
making a lax small but conspicuous reticulum above and more or
654 Field Museum of Natural History— Botany, Vol. XIII
less prominent beneath; petioles 1-2 cm. long, sub terete, shortly-
pubescent or glabrescent; inflorescence on the trunk and on the
larger branches in cymes forming small clusters with short tuber-
cular or nodose woody branchlets, eventually with elongated
branchlets up to 3 cm. long; pedicels slender, villous, to 1.5 cm.
long, with 3 small linear deciduous bracts at the base; calyx pale
reddish, the 5 sepals 5-6 mm. long, lanceolate, almost free, glabrous
within, subglabrous without, stellate-pubescent marginally; petal-
base oblong, hooded, 2.5-3 mm. long, glabrous, appendage obovate-
rhomboid, 1.5 mm. long and broad, cuneate and long pediculate
(pedicel about 2 mm. long, geniculate), yellowish; staminal tubcj
1 mm. long, the fertile filaments curled, biantheriferous; staminodis
subulate, rather fleshy, minutely papillose, scarcely hirsutulous at
the margin toward the base, 4-5 mm. long; ovary ovoid, glabrousj
as the 2 mm. long style; fruit ovoid or oblong-ovoid, obtuse oi
acute, usually pentagonal with more or less pronounced rows
according to the variety; pericarp thick, firmly fleshy, glabrous,^
yellowish or reddish at maturity, about 1.5-3 dm. long, the ovoid'
seeds about 26 mm. long, 18 mm. broad, and 9 mm. thick.
Probably all the specimens found in wild condition in the ter-g
ritory of Peru belong to the subsp. leiocarpum (BernouHi) Cuatr^
comb. nov. (T. leiocarpa Bern., I.e. 6. pi. 2, fig. 11. 1871), whicl
only can be distinguished when with fruits, frequently lacking ii
herbaria. This subspecies is characterized by its ovoid obtus
smooth pods and violet pigmented cotyledons. This is the for
believed to be native in South America, spreading throughout it
tropical regions.
There are few data about the cultivated cacao and its varieties
in Peru, where they never had the economic importance the]
obtained in Ecuador or Colombia. But here and there are plantec
the var. "criollo" with marked ridged acute pods and white coty-
ledons, and chiefly the "forastero" variety with smooth obtuse podsj
and violet cotyledons, and several intermediate forms.
After McLaughHn, Cacao Information Bulletin 2 no. 6: 3 (Turri-|
alba, 1950), according to statistics for the year 1929, there were
11.115 hectares of cacao in Peru. Annual production at the presentj
time is said to be 2,000 to 3,000 tons. The principal producing
regions are near Ja^n (Dept. Cajamarca), Huancabamba (Dept.
Piura) and Valle of Convencion (Dept. Cuzco); there is some pro-j
duction, also, in the Departments of Madre de Dios, Huanuco, Sani
Martin, and Loreto, where there are small, scattered, cacao planta-J
Flora of Peru 655
tions along the lowland rivers, especially near Iquitos. Herrera
recorded the export of a "Cacao chuncho" from the valleys of the
lower Umbamba, known to the inhabitants as "Sargemini."
T. sagittata Pavon ex Chevalier (Rev. Int. Bot. Apl. & Agr.
Col. 26: 274. 1946) and T. sinuosuyn Pavon ex Huber (Bull. Herb.
Boiss. s6r. 2, 6: 274. 1906) are insufficiently known and have not
been seen by the author; presumably they are only forms or varieties
of T. cacao L.
Junin: La Merced, 5278. — Loreto: Caballo-cocha in the Amazon
River forest, Williams 2105, 2349. Alto Rio Itaya, San Antonio,
Williams 3510. Along Rio Itaya, Rfo Mazana, Williams H8.
Lower Rio Huallaga, Recreo, Yurimaguas, WiUiams ^160. San
Antonio on Rio Itaya, dense forest, Killip & Smith 29U3U- Iquitos,
fundo Indiana, Y. Mexia 6399. Balsapuerto, forest, Klug 2938.
Mishuyacu, near Iquitos, Klug 926. "Cacao," "cacao sylvestre"
or "comun."
Theobroma calodesmis Diels, Notizbl. Bot. Gart. Berlin 14:
336. 1939.
Tree to 20 meters with glabra te branchlets; leaves coriaceous;
petioles robust, tomentulose, 8-10 mm. long; blades subelliptic,
acuminate, broader and asymmetric at base, entire, 2-2.5 dm. long,
8-12 cm. wide, the upper surface glabrous, pale beneath, except the
glabrous nerves, covered with a whitish microscopic compressed
stellate indument; midnerve and 5-6 pairs of lateral nerves very
prominent beneath, the inferior subbasal pair stronger, straightly
ascendant, separated 5-7 cm. from the others, which are curved-
ascending; minor nerves transversal and reticulate, prominent be-
neath; flowers in corjmibiform caulinar pedunculate clusters, 10-20
densely fasciculate, borne on pubescent slender 8-10 mm. long
pedicels; sepals narrowly ovate, 8-13 mm. long, 3-4 mm. broad,
reddish, whitish tomentose, pinkish within; base of the petals
obovate-ellipsoid, auriculate at apex, trinervate, 4-5.5 mm. long, 3-^
mm. broad, pink-tinted, the purplish ligule subrotundate, 3.5-5.5
mm. long and broad, abruptly contracted and articulate at base;
staminal tube 2 mm. long, the fertile filaments 2-3 mm. long, flexu-
ose, bearing 2 or 3 bilocular anthers; staminodia lanceolate, purplish,
9-12 mm. long, 1.3-1.8 mm. broad at base; ovary tomentose, 2-2.5
mm. long, the style 2.5-3 mm. long with 5 nearly free branches;
fruit ovoid-oblong, obtusely 5-ridged, rounded and umbilicate at
base, attenuate and acuminate at apex, 10-11 cm. long, 5-5.5 cm.
broad {Frdes 206 Jt5).
656 Field Museum of Natural History— Botany, Vol. XIII j
The type specimen of T. calodesmis (Schultze-Rhonhof 2312) was
collected in Ecuador at Pacayacu River (local name "Chucu"),
affluent of the Bobonanza near the Peru border. It is to be found
undoubtedly in Peru, having been collected recently in Brazil and
Colombia; Brazilian local name "cacao azul" (Froes).
Peru (undoubtedly). Brazil to Ecuador and Colombia.
Theobroma grandiflorum (Spreng.) Schum. in Mart. Fl. Bras.
12, pt. 3 : 76, pi. 8. 1886. Buhroma grandiflorum Spreng. in L. Syst.
Veg. ed. 16. 5, pt. 3: 332. 1826. T. macrantha Bernoulli, I.e. 11. 1871.
Tree frequently 8-10 meters high, but may attain 18 meters;
branchlets, when young, wooly ferrugineous, later glabrate, gray or
brown and more or less cracked or scaled; leaves firm-coriaceous,
petiolate, usually obovate-oblong or oblong, a little narrower toward
the rotundate or sometimes subcordate base, rotundate and abruptly'
acuminate at apex, entire or slightly sinuate, usually 2-3.5 dm. to
6-11 cm. broad, the upper surface lustrous, glabrous and only the
principal filiform depressed nerves conspicuous, the lower with
prominent midrib and about 10 pairs of secondary nerves, these
ascending and curved near the edge, prominent, the transverse
tertiary nerves parallel and others less prominent forming a small
reticulation; the secondary nerves pinnate but the inferior pair
forming a more acute angle and more separated from the next ones;
the venation glabrous, pale and dotted with scattered reddish
minute warts, but the epiderm within the net covered with a dense
whitish tomentum made by very tiny intricated stellate hairs;
petioles 5-10 mm. long; inflorescences at the branchlets mostly
axillary, with a small number of flowers (1-4); peduncles 1.5 cm.
long, articulate, with 3 verticillate linear bracteoles only 2-3 mm.
long, densely ferrugineous tomentose with stellate hairs; sepals 3-5,
subcoriaceous, thick, ovate- triangular, about 14 mm. long, 6-8 mm.
broad, within pale, reddish, thin-pubescent, without densely fer-
rugineous-tomentose, united at the lower part; petal-cowl (or base)
suborbicular, 6-7 mm. long and broad, pubescent, strongly hooded
at the apex, the ligule obcordate or elliptical-rotundate, emarginate,
6 mm. long, to 7 mm. broad, red or purple, glabrous or scarcely
pilose, pediculate, the pedicel about 4 mm. long; staminal tube
1.6 mm. long, the purple membranous staminodia lanceolate, long-
acuminate, about 1 cm. long, 2 mm. broad, hirtulous; fertile filament
2 mm. long, thick, curved, the anthers 3-lobate, each part bicelled;:
ovary tomentose, hirsute, subgolobse, the glabrous subulate style
Flora of Peru 657
2 mm. long; fruit ovoid-ellipsoid, smooth, to 24 cm. long, 12 cm.
broad, with coriaceous hard pericarp covered with dense brown
tomentum. — Planted throughout the Amazonian basin, in Brazil,
Colombia, Venezuela, Peru. "Cupuassu" (Williams). The edible
pulp of the fruits is used for preparation of beverages and preserves.
F.M. Negs. 40706; 40705.
Loreto: Caballo-cocha on the Amazon River, Williams 2^01,
Theobroma obovatum Klotzch in Bernoulli, Uebersicht d. Th.
Arten 14. pi. 8. 1871.
Tree, 10-12 meters tall, with gray branchlets, the young ones
tomentulose becoming glabrous; leaves petiolate, pergaminaceous,
obovate-oblong or obovate-elliptic, very asymmetric and rounded
or obtuse at the base, shortly attenuate-cuspidate at apex, the
margin entire, almost smooth, 10-24 cm. long, 3.5-10 cm. broad,
pale green and glabrous above, the venation little conspicuous,
beneath pale cinereous, the midnerve very prominent, the secondary
nerves 5-6 each side, prominent, ascendant, curved and subdecurrent
toward the margin, the tertiary nerves transverse, very prominulous,
and the minor veins slender, reticulate; the nerves beneath glabrous
and marked with scattered small callous warts, the epidermic
surface covered with an appressed thin tomentum composed of tiny
intricate stellate hairs, the two lateral basilar nerves a little more
separated and in a more acute angle than the other pairs, the
nervation thus really pinnate rather than triplinerved at base;
petioles 5-10 mm. long, minutely tomentulose; stipules very nar-
rowly linear, tomentose, 3 mm. long, soon deciduous; flowers in
small few-flowered axillary cjmies, the slender stellate tomentose
pedicels 10-15 mm. long, articulate, with 3 very small (1 mm. long)
caducous bracts at the joint; buds globose, ochraceo- tomentose,
5-6 mm. in diameter; sepals ovate-oblong, acute, 6 mm. long, 3.4^
mm. broad, united for 1 mm. at base, pale red within, ochraceous
without; cowl or base of petals red, orbiculate-elliptic, 2.5 mm.
long, apex rounded-cucullate, 7-nerved, glabrous, the ligule orbicu-
late, 3.5 mm. long and broad, emarginate at apex, little cuneate at
base to a pedicel 2 mm. long, petaloid, deep red, scattered pilose;
staminal tube short (1 mm.), staminodia petaloid, obovate-oblong,
obtuse, 5-5.5 mm. long, 2.5 mm. broad, deep red, sparsely hirtulous,
chiefly at the margin; fertile filament 1 mm. long, rather thick,
curved; the anthers trimerous, each part bilocular; ovary tomentose,
the style glabrous; fruit obo void-ellipsoid, about 7 X 4 cm., with
a coriaceous pericarp densely covered with acute hard warts.
658 Field Museum of Natural History— Botany, Vol. XIII
Loreto: along Rio Itaya, Williams 161; 230. Lower Rio Hual-
laga, Puerto Arturo, Yurimaguas, Williams 5268. Balsapuerto,
forest, Klug 2983. Colombian Amazonas; Brazil. From Brazil are
registered as local names "cabeca de urubu" (Ducke), "cabeca de
umbu" (Krukoff), "cupuarana" (Frdes).
Theobroma speciosum Willd. ex Spreng. in L. Syst. Veg. ed.
16. 3: 332. 1826; Bernoulli, I.e. 8. pis. 2-3. T. quinquenervia Bern.
I.e. 8. pis. 2-3. 1871. T. speciosum var. quinquenervium K. Schum.
in Mart. Fl. Bras. 12, pt. 3: 75. 1886.
Tree, to 15 meters tall, trunk about 2 dm. in diameter, the lightj
gray bark smooth; branehlets brownish, more or less stellate
tomentulose; leaves firm-coriaceous, pale, short-petiolate, ovate oi
oblong-ovate, broadly rounded and very asymmetric at the base^
more or less attenuate toward the rotundate apex, abruptly cuspi-^
date, the margin entire and smooth, 3-4 dm. long, 14-18 cm. wide^
the upper side smooth, glabrous, only the principal nerves con-
spicuous, the lower pale-cinereous nervation subglabrous but th(
surface covered with a thin very compressed tomentum of minut
whitish stellate hairs; nerves usually 5, the central nerve much the
stronger, 7-8 pairs of secondary pinnate nerves starting a thirc
distant from the base, prominent, subcurved-ascending and sub-"
decurrent at the margin, the tertiary nerves transversal, also promi-j
nent, the minor veins rather obvious, reticulate; petioles tomentu-S
lose, thick, 1-1.5 cm. long; flowers long-pedicellate in delicate
panicles gathered and concentrated in many-flowered bunches borne
on the trunk, these massed inflorescences (to a dm. large or longer)
very showy, comprising even as many as 200 purple flowers; branch-
lets of the inflorescence 3-5 cm. long, very thinly and canescentl]
stellate-tomentulose, the filiform tomentulose pedicels to 1.5 cm.j
long; buds rotundate, oblong, reddish, 1 cm. long; calyx of 5 ovate
acute sepals 1 cm. long, 4 mm. broad, united one-third, purplishj
and glabrous within, reddish and covered with a pubescence of
stellate hairs without; petal-base red, obovate-oblong, 6 mm. long^
3 mm. broad, the apex rotundate, 3-nerved, scarcely puberulous,j
shortly unguiculate, pale red with purple nerves, the ligule obovat
spathulate, rotundate or truncate at apex, cuneate at base, sessile^
6-8 mm. long, 7 mm. broad, glabrous or subglabrous, deep red anc
venulose; staminal tube 2 mm. long, the purple staminodia oblong-
linear, long-acuminate, rather thick, densely papillose-hirtulous^
5 mm. long; fertile filaments 2 mm. long, flexuose, the anther
Flora of Peru 659
trimerous, lobules 2-celled; ovary obovoid, whitish-tomentose, 2
mm. long, the glabrous 5-parted style 2 mm. long; fruit globose-
ellipsoid, slightly pentagonous, about 10 X 8 cm., covered with
a tenuous tomentum; pulp white, sweet, inodorous (Ducke). — F.M.
Negs. 9640, 9639, 40703.
Loreto: middle Ucayali, Yarina Cocha, Tessmann 5398. Spread
and planted throughout Amazonia chiefly in Brazil, but also in
Bolivia, Ecuador, and Colombia. Brazilian names "cupuhy," "cacao
azedo," "cupurana," "cacaohy," "cacao azul."
Theobroma subincanum Mart, in Buchner, Repert. der
Pharm. 35: 23. 1830. Cacao sylvestris Aublet, Hist. PI. Guyan. franc.
2: 687. pi. 276. 1775. T. ferruginea Bern. Neue Denksch. Schweiz.
Ges. 24: 13. 1871. T. tessmannii Mildbr. Notizbl. Bot. Gard. Berlin
11:139.1931.
Tree rarely more than 15 meters tall, trunk 1.5 dm. in diameter,
with a gray bark; branchlets when young appressed-tomentose, later
glabrescent and rugulose; leaves firmly coriaceous, large, short-
petiolate, subobovate-oblong, rotundate at base and abruptly cuspi-
date, the margin entire or slightly sinuate toward the apex, usually
2-11 dm. long, 7-14 cm. broad, the upper side glabrous, principal
nerves very narrow and little conspicuous, the under side with
a very thick and prominent midnerve and 9-12 pairs of prominent
ascending parallel secondary nerves, those near the margin sub-
decurrent-curvate; tertiary veins transversal, prominent and reticu-
lately united with the smaller; tomentum double: on the ground
a whitish, dense, appressed tomentum formed by tiny slender
entangled hairs; on the veins comprised of larger and thicker,
reddish, stellate hairs; the density of this second indument is variable,
but it is always present; if scattered, the color of the leaves is more
ashy, as in the type; if more abundant, they cancel the ground color
and make the leaf appear brownish or ferrugineous; petioles 1-1.5
cm. long; flowers axillary or at branchlets in small 1-6-flowered
clusters; pedicels 4-5 mm. long, ferrugineous- tomentose, articulate
at the middle with 3 filiform tomentose deciduous bractlets 3 mm.
long; calyx of 5 sepals 7-8 mm. long, 3 mm. broad, one-third or half
united, ovate-oblong, acute, reddish and subglabrous within,
ferrugineous stellate-tomentose without; petals purple, the cowl
(base) obovoid, 5-6-nerved, the apex rotundate, hooded, 3-3.5 mm.
long, 2-2.5 mm. broad, puberulous; ligule obovate-spathulate, long-
pedicellate, at apex truncate or emarginate, base obtuse-cuneate,
660 Field Museum of Natural History— Botany, Vol. XIII
2 mm. broad, 1.5-2 mm. long, subglabrous, the margin puberulous,
the pedicel thin, 2 mm. long; staminal tube about 2 mm. long, the
membranaceous purple staminodia elliptic-oblong, subacute or
obtuse, 7 mm. long, 2 mm. broad, scarcely pilose; fertile filaments
1-1.5 mm. long, curved, the anthers 3-lobate, lobes 2-celled; ovary
obovoid, hirsute-tomentose, the acute glabrous style 2 mm. long;
fruit ellipsoid or elhpsoid-oblong, about 10 X 5-6 cm., with a short
thick pedicel (1x1 cm.), subrotundate or obtuse at both ends, the
coriaceous pericarp 2-4 mm. thick, with thin appressed stellate
tomentum; seeds ovoid or ovoid-oblong, more or less compressed,
22-32 mm. long, 12-16 mm. broad.— F.M. Negs 17942, 19645, 40704.
Loreto: Tierra Doble on the Rio Nanay, deep forest, Williams
1076; Alto Rio Itaya, forest of Paraiso, Williams 325 If. Santa Ana,
on the upper Rio Nanay, Williams 1233. Mishuyacu near Iquitos,
forest, Klug 87; Klug 857. Without definite locality, Tessmann s. n.
(NY-3717) : Brazil, Colombia. "Cacao-ceniza," "cumala," "uchpa-
cacao" (Williams). In Brazil "cupuhy," "cupui."
7. GUAZUMA Plum.
Trees with unequally dentate often oblique stellate pubescent
leaves and axillary shortly peduncled cymes of small flowers. Calyx
2-3-parted. Petals concave, inflexed, terminating in a linear bifid
appendage. Stamens united into a campanulate tube with 5 sterile
lobes and 5 stiped clusters of 4-6 anther cells. Ovary 5-celled with
many ovules; styles more or less connate. Capsules subglobose,
ligneous, tuberculate-echinate or setose.
Leaves firm; fruits muricate; inflorescence congested. .G. ulmifolia.
Leaves membranous; fruits densely long-setulose; inflorescence
open G. crinita.
Guazwma crinita Mart. Flora 20, pt. 2: Beibl. 95. 1837.
G. rosea Poeppig in Poepp. & Endl. Nov. Gen. & Sp. 3: 72. 1845.
Tall tree with usually subcordate ovate leaves much exceeded
by the copiously flowered tomentulose panicles of rose-tinted
blossoms, most of them sterile; branchlets erect, terete, deciduously
tomentulose; petioles to 12 mm. long; leaves obtuse, unequally
serrate, basally 5-nerved, tomentose beneath, minutely and sparsely
stellulate above, membranous, to 1 dm. long, half as wide; peduncles
about half as long, the unequal pedicels corymbose or cymose;
calyx 3-parted, the segments broadly ovate; petals cucullate, api-
Flora of Peru 661
cally 3-dentate, the ligule deeply biparted; anthers in 3's; ovules
numerous; capsules ovate, acute, ligneous, hirsute. — After Poeppig.
To over 30 meters tall with rather tall buttresses, the wood used for
sugar boxes and crates according to Williams, who gives anatomical
descriptions (also for G. ulmifolia), Field Mus. Bot. 15: 319. 1936.
Illustrated, Poeppig & Endlicher, I.e. pi. 283. F.M. Neg. 9637.
San Martin: Tarapoto, Williams 5698. Chazuta, Kliig Jt059.
— Loreto: Yurimaguas, Poeppig (type, G. rosea); Williams U221;
5015. Pongo de Manseriche, Mexia 6126. Balsapuerto, King 2971.
Lower Rio Nanay, Williams J^SU. Rio Itaya, WiUiams 151. Brazil.
"Atadija" (Poeppig), "bolaina" (Poeppig).
Guazuma ulmifolia Lam. Encycl. 3: 52. 1789. G. utilis
Poeppig in Poepp. & Endl. Nov. Gen. & Sp. 3: 72. 1845, fide Freytag.
G. tomentosa HBK. Nov. Gen. & Sp. 5: 320. 1823.
Tree, sometimes 30 meters tall, with long branchlets densely
stellate toward the tip, oblong or oblong-ovate acuminate 3-nerved
serrulate sparsely stellate leaves (both sides or canescent beneath),
short cjmies (about as long as petioles) of yellowish flowers; petioles
short or to 2.5 cm. long; leaves to 1.5 dm. long, 5 cm. wide, obtusely
acute and oblique at base; peduncles bifid; calyx ferrugineous
tomentose, the segments subrotund; fruits subglobose, 2-4 cm. thick,
densely roughened with short sharp tubercles, the many large hard
seeds surrounded by a little edible sweet pulp. — After Poeppig.
Leaves densely tomentose in the var. tomentella Schum. in Mart.
Fl. Bras. 12, pt. 3: 81. 1886, apparently G. tomentosa HBK. Some-
times 30 meters tall, and common in the Middle and Lower Huallaga
regions (Williams) ; the inner laminated bark furnishes a fiber used
for cordage. See Standley & Steyermark, Fl. of Guatemala (Fieldi-
ana: Bot. 24, pt. 6: 411-412. 1949), who list a truly remarkable
number of past or present attributes of this species in Central
America, including its food interest to bees, cattle, silk-worms and
children! Illustrated, Fawcett & Rendle, Fl. Jam. 5: 157, fig. 59.
F.M. Negs. 9638 (G. utilis); 35346.
Cajamarca: Rio Chotano Valley, Weherbaiier 7116 (var.); Rai-
mondi. — Lambayeque: Olmos, Weherhauer 7092 (var.). — San Martin:
Tarapoto, Ule 6ItSU; WiUiams 5675. — Junfn: La Merced, Weherhau£r
19It5; 6271 (var.). — Loreto: Yurimaguas, Poeppig ity\)e, G. utilis);
Williams k227; J^768 (det. Standley). Balsapuerto, Klug 2939 (det.
Standley). Florida, Klug 1995; 2018 (det. Standley). Rio Parana-
pura, Klug 3952 (det. Freytag; det. Standley, G. crinita). Puerto
i
662 Field Museum of Natural History— Botany, Vol. XIII
Mel^ndez, Tessmann ^7^.0. Yarina Cocha, Tessmann 314.62. To
the West Indies; Mexico. "Huashimo" (Raimondi), "hapayillo,"
"lluicho-vanilla" (Williams), "iumanasi," "hudsimo" or "hudsim"
(Weberbauer).
8. HELICTERES L.
Shrubs with stellate or branched pubescence, entire or serrate
leaves, and usually zygomorphic axillary flowers, solitary or in
clusters. Calyx tubular, 5-parted, often irregular. Petals 5,:
unguiculate, some or all frequently auriculate. Gynophore early
strongly curved, elongate, with 6-many slightly connate stamens,
the anthers linear. Ovary 5-lobed, 5-celled; styles 5, subulate.
Fruit composed of 5 ligneous commonly spirally twisted follicles
with many verruculose seeds. — R. E. Fries, Sv. Vet. Akad. Handl.
ser. 2. 42, no. 12: 21-22. 1908, gave a key to the actinomorphic
flowered species.
Leaves subsessile in a plane; petals included H. pentandra.
Leaves distinctly petioled; petals exserted H. ovata.
Helicteres ovata Lam. Encycl. 3: 88. 1789.
Fulvous stellate-tomentose including the calyces, or the leaves
in age greenish and more or less glabrate above; petioles 1-2 cm.
long, the caducous filiform stipules about as long; leaves broadly
ovate, acute or acuminate, often about 12 cm. long and more than
half as wide, unequally serrate; dichasia opposite the leaves or
subaxillary; calyx one third parted, about 2 cm. long, bilabiate,
3-5-lobate, subfloccose-tomentose, the spathulate red petals 23-27
mm. long, the processes near the base lacerate-dentate; gynophore
7-9 cm. long; fruit 4 cm. long, 1.5-2 cm. in diameter, finally gla-
brescent. — The Peruvian specimen was distributed as a new species
by Standley under the name of the country; the one flower has
coarser indument than described in Schumann's key to the species
of Brazil but matches closely some material so determined. Illus-
trated, Mart. Fl. Bras. 12, pt. 3: pi. 5. F.M. Neg. 35375.
San Martin : Juanjui, Klug Jf265. To Paraguay.
Helicteres pentandra L. Mant. 2: 294. 1771.
Strict shrub or little tree sometimes a few meters tall, with
widely spreading virgate rather coarsely stellate pubescent branches,
especially in the leafless lower part, closely leafy above, and sub-
Flora of Peru 663
regular flowers borne on small branchlets (dichasia) in their upper
leaf-axils, the subsessile ovate-oblong leaves disposed in a plane,
subcordate at base, acute, mostly about 1 dm. long, at least half as
wide, doubly serrulate, green, rugulose and scabrous-stellate above,
canescent-stellate beneath; bracts lanceolate, caducous; peduncles
4-5, pedicels 3-4 mm. long; calyx clavate-tubular, hirsute- tomentose,
about 2 cm. long, early yellowish becoming roseate, slightly longer
than the linear-spathulate white or roseate petals; gynophore to
4 cm. long; fruits 2-2.5 cm. long, 1 cm. in diameter, densely hirsute.
— ^Actually with 6 stamens, the only actinomorphic species with this
number. Grows on open sandy plains. Illustrated, Mart. Fl. Bras.
12, pt. 3: pi. S.
San Martin: Tarapoto, Woytkowski 350S5; Williams SU07; 590U.
In clearing, Juanjui, Klug It^05 (det. Standley). — Junin: La Merced,
5299; 5U86. Rio Peren^, Killip & Smith 2U969; ^5iii.— Amazonas:
Moyobamba, Mathews 1547 (det. Hochreutiner) . To the Guianas.
"Tornecillo" (Williams).
9. STERCULIA L.
Trees with simple or lobed leaves and unisexual or polygamous
flowers borne in axillary or terminal racemes or panicles. Calyx
5-cleft, usually colored. Petals none. Staminal tube with 10 or
15 anthers crowded apically. Carpels of the ovary 5, nearly free,
2-many ovuled, the styles peltate or lobed, the mature carpels
distinct, tardily dehiscent introrsely. Seeds 1-many, the endosperm
fleshy. — Flowers variable in size, and normally simple leaved species
may have lobed leaves on vegetative shoots (Ducke, Archiv. Jard.
Bot. Rio. Jan. 3: 213. 1922). Besides the following, S. roseiflora
Ducke, Archiv. Inst. Biol. Veg. Rio Jan. 2: 58. 1935, from as near
as Sao Paulo de Olivenga may occur within Amazonian Peru and
perhaps is unique in its rose-colored flowers. This is yet another
genus without a modem revision, and ex char, at least, many
proposed species are incompletely known.
Leaves of flowering branchlets simple.
Mature leaves covered beneath with a minute close indument.
S. solitudinis.
Mature leaves glabrous, glabrate or more or less loosely stellate
pubescent beneath.
Leaves quite glabrous at least in age, equally reticulate veined
both sides S. glabrifolia.
664 Field Museum of Natural History— Botany, Vol. XIII
Leaves at least slightly pubescent in the nerve axils or on the
nerves beneath in age.
Leaves smooth beneath, or reticulate, often about 2 dm.
long; fruits (known) about 4 cm. long.
S. Tessmannii; S. pruriens.
Leaves rugose-nerved, often ample; fruits about 1 dm. long.
S. speciosa.
Leaves of flowering branches 3-5-lobed, cordate S. apetala.
Sterculia apetala (Jacq.) Karst. Fl. Columb. 2: 35. pi. 118. 1869.
Helicteres apetala Jacq. Enum. PI. Carib. 30. 1760.
Tree sometimes attaining 30 meters and developing a tall straight
trunk, the densely leafy crown depressed and spreading; petioles to
2 dm. long, as the leaves beneath early more or less tomentose but
at least in Peru promptly glabrate, the stellate trichomes scattered
and inconspicuous unless on the nerves and veins; leaves often
3 dm. broad, deeply 3-5-lobed, the broad lobes rounded-obtuse or
acutish; panicles often equaling the petioles, apparently crowded
typically; calyx campanulate, about 2.5 cm. wide at the flaring top,
1.5-2 cm. long, yellowish with purple dots and lines; fruiting carpels
tomentulose without, hispid within, usually about a dm. long with
2-4 large black ellipsoid seeds. — Determinations by Standley.
S. elata Ducke, Archiv. Jard. Bot. Rio. Jan. 3: 211. 1922, may be,
possibly, the Peruvian tree; it was separated by the author from
S. Chicha St. Hil. of Goyaz, Brazil, which according to Schumann
differs in size of fruits and leaf-texture, while S. elata has gynophore
about one half instead of one third as long as calyx; the characters
appear to be relative.
San Martin: Juanjui, Klug U262. — Hudnuco: Monzon, Weber-
hauer 3652 (det. Ulbrich, S. Chicha). Tingo Maria, Burgos 3U. —
Loreto: Mishuyacu, Klug 2529. Puerto Mel^ndez, Tessmann 3913
(det. Ulbrich, S. Chicha). — Rio Acre: Mouth Rio Macauhdn, Kru-
koff 5^71 (distr. as S. elata Ducke?) . To Mexico and the West Indies.
"Sahote," "husyracaspi."
Sterculia glabrifolia Mildbr. Notizbl. Bot. Gart. Berlin 9:
1148. 1927.
Glabrous except for a stellulate puberulent indument in the
inflorescence; petioles 4-8 cm. long; leaves coriaceous, oval or
obovate-oval, rounded-obtuse at base, abruptly short-acuminate,
10-24, mostly about 17 cm. long, 4.5-13, mostly 8-10 cm. wide, with
■
Flora of Peru 665
&-10 lateral nerves, the reticulate veins equally manifest both sides;
flowers small, the sepals connate about 1.5 mm., the free portion
5 mm. long; fruit unknown. — Similar to S. frondosa Rich, but differs
in the entire absence of pubescence on the leaves and in the evident
and fine reticulation (Mildbraed) ; and S. frondosa at least as to type
has slightly retuse leaves. However any difference with S. pruriens
seems vague. F.M. Neg. 9623.
Loreto: Mouth of the Santiago, Tessmann WUl, tyv^-
Sterculia pruriens (Aubl.) Schum. in Mart. Fl. Bras. 12, pt. 3:
8. 1886. Ivira pruriens Aubl. PI. Guian. 2: 695. pi. 279. 1775.
Branchlets, petioles, these to 1 dm. long, and inflorescence
reddish-rusty tomentulose; leaves early pubescent above, tomentose
beneath, at maturity glabrate, subcoriaceous, oblongish, often
1.5-2.5 dm. long, 6-10 cm. wide, rounded or slightly or obscurely
cordate at base, distinctly but shortly acuminate at apex; calyx
12 mm. long; carpels rostrate, 4 cm. long, 3.5 cm. thick, borne on
a gynophore 2 cm. long. — Attains 40 meters (Krukoff). S. pilosa
Ducke, Archiv. Jard. Hot. Rio Jan. 3: 212. 1922, is rather similar
but seems to be well marked by its very long and abundant pilosity
and perhaps is rather allied to S. speciosa as the leaves are coarsely
bullate-rugose and lustrous above, the flowers pale reddish. S.
apeihophylla Ducke, Bol. T^n. Agron. Norte, Bel^m 4: 16. 1945, to
be expected, is similar according to the author but the opaque
leaves are smoother above, petals deep red, malodorous and the
fruits large as in S. speciosa.
Rio Acre: Mouth Rio Macauhdn, Krukoff 5421; 563 U (det. N. Y.
Bot. Gard.). To the Guianas.
Sterculia solitudinis Mildbr. Notizbl. Bot. Gart. Berlin 11:
138. 1931.
Type a sparsely branched tree 30 meters tall with trunk 9 dm. in
diameter, the specimen branch 12 mm. thick with glabrate rugulose
bark; petioles 6 or 7 cm. long; leaves ovate, little cordate at base,
subacute, about 2 dm. long, 14-18 cm. wide (or upper 9 X 12 cm.),
finally coriaceous, glabrous and the 7 primary veins scarcely promi-
nent above, beneath with the veins conspicuously reticulate and
compactly stellate-tomentose with dark ferrugineous indument;
panicle apparently few-branched, about 8 cm. long, tomentulose-
puberulent, including the stellately spreading male flowers, their
sepals 2 mm. connate, 7 mm. free, acuminate from the 2 mm. wide
666 Field Museum of Natural History — Botany, Vol. XIII
base with a tiny transverse callus within on the upper third, and
toward the base fasciculately long-pubescent; basally enlarged 10-
anthered androgynophore 5 mm. long; cupulate stamen tube gla-
brous as the rudimentary pistils. — Leaves densely red-rusty stellate
beneath in youth, later glabrate, rugose-reticulate; seems too near
the Colombian S. rugosa R. Br. but the author wrote: easily recog-
nizable by the dense reticulation of the leaves beneath and the
short red-brown felt that covers them so closely that a ten power
lens scarcely defines the stellate trichomes individually. F.M.
Neg. 17943.
Loreto: Rio Itaya, Tessmann 5190, type.
Sterculia speciosa Schum. in Mart. Fl. Bras. 12, pt. 3: 7. 1886.
Similar, apparently, to S. pruriens, but, according to Ducke, the
sterile branches at least with very large leaves often with a rounded
lateral lobe, fruits 12 X 10 cm. and flowers very malodorous; petioles
to 8 cm. long (probably longer); leaves oblong, obtuse, emarginate
and mucronate, rounded at base, those of flowering branches 2-2.5
dm. long, 9-14 cm. wide, rugose and glabrescent beneath, puberulent
in the axils; calyx variable in size, usually more than 15 mm. long,
the lanceolate acuminate segments tomentose both sides; gynophore
filiform, 8-9 mm. long, glandular-tomentose only at base, less than
half as long as the caljrx; stamen tube cupulate; fruits large (Ducke)
with 10 ellipsoid seeds (Martius). — Illustrated, Schumann, I.e. pi. 1.
Peru (probably). Upper Amazonian Brazil.
Sterculia Tessmannii Mildbr. Notizbl. Bot. Gart. Berlin 9:
1149. 1927.
Branchlet tips densely puberulent; petioles 4-11 cm. long; leaves
coriaceous, ovate-oval, rounded or obscurely cordulate, subabruptly
acuminate (acumen sometimes 2 cm. long), 1-2.5 dm. long, 6-14
cm. wide, glabrous above, sparsely stellate pubescent beneath,
lateral nerves about 8 and with transverse veins prominent beneath,
reticulation obvious above, obsolete beneath; calyx 9 mm. high,
connate about 1.5 mm.; otherwise no distinctions with related
species noted. — Type from a tree 18 meters high and according to
the author distinguished from S. pruriens by the much less dense
pubescence on the under surface of the leaves and the prominent
lateral and secondary venation beneath, apparently intangible
characters. Williams 2570 from La Victoria at the boundary and
referred here by me with query in herb, is doubtfully a Sterculia;
the specimen seen has only young buds. F.M. Neg. 9626.
I
Flora of Peru 667
Loreto: Flood-free woods at mouth of the Rfo Apaga below
Pongo de Manseriche, Tessmann S8U7, type. Mishuyacu near
Iquitos, KliLg 2525; 2512? (this with larger flowers proposed in herb,
by Standley as undescribed).
DILLENIACEAE
References: Eichler, Fl. Bras. 13, pt. 1: 65-120. 1863; Lanjouw
& Van Heerdt, in Pulle, Fl. Surinam 3: 386-408. 1941.
Trees or shrubs, sometimes scandent, rarely herbs, the alternate
simple leaves usually with harsh pubescence and with many parallel
lateral nerves. Stipules if present alate and adnate to petiole,
mostly deciduous. Flowers variously disposed, hermaphrodite, with
5 (rarely fewer or more) imbricate sepals and petals, the former
p)ersisting and frequently accrescent, the latter early deciduous and
often crinkled in bud. Stamens usually many, free or variously
connate at base; anthers dehiscent apically or longitudinally.
Gynoecium usually of several distinct pistils, ovary superior, 1-celled
and -carpelled, placentation parallel, sometimes appearing basal,
ovules 1 or more, each with 2 integuments, erect, the styles as many
as pistils, distinct. Fruit a follicle or berry-like, seeds with copious
endosperm, embryo minute and usually somewhat lacerated.
The types of the several species I have proposed in this family
were not available in the preparation of this final draft and con-
sequently their status, which I doubt now, has not been reviewed.
Sepals unequal in flower, the larger inner valvate and enclosing
racemose or paniculate fruits 1. Davilla.
Sepals nearly or quite alike, not accrescent.
Inflorescences usually solitary, terminal; aril lacerate.
2. Tetracera.
Inflorescences axillary or at leaf -scars, rarely in part terminal.
Flowers usually in axillary fascicles; carpels often 1; fruit
a berry or capsule 3. Doliocarpus.
Flowers in softly pubescent clusters at leaf-scars; carpels 2;
fruit a follicle 4. Curatella.
1. DAVILLA Vand.
Commonly lianas with entire or dentate estipulate leaves, the
trichomes always simple, and mostly terminal racemes or panicles
668 Field Museum of Natural History— Botany, Vol. XIII
of yellow flowers, or these axillary and reduced, sometimes greatly.
Sepals 5, the inner 2 much the larger, concave, after anthesis in-
durate. Petals 5 (1-6). Stamens many, the filaments thickened
apically, the cells of the globose anthers divaricate basally. Carpels
1-2 (3), 2-ovuled, the terminal style persisting, at maturity inde-
hiscent or regularly dehiscent, the 1-2 follicles within the persisting
stamens and the 2 inner sepals. Seeds black, lustrous, nearly
enclosed by the aril.
Branchlets glabrate to hirsute, not villous-tomentose.
Glabrate or glabrous even to inflorescence, the leaves smooth
above D. multiflora.
Pubescent, or at least leaves asperous above.
Leaves coarsely reticulate-veined beneath D. aspera.
Leaves finely veined beneath D. rugosa.
Branchlets rusty villous-tomentose D. densa.
Davilla aspera (Aubl.) Benoist, Bull. Soc. Bot. Fr. 60. 392. 1913.
Tigarea aspera Aubl. PI. Guian. 2: 918. pi. 350. 1775. D. lucida
Presl, Rel. Haenk. 2: 73. 1835. Tetracera Tigarea DC. Syst. Veg.
1:403. 1818.
Harshly asperous liana, especially the more or less elliptical
leaves above, these notably reticulate beneath by many veins that
are not at all or scarcely visible on the upper surface; lateral nerves
as sometimes tertiary impressed above, 8-24, prominent and usually
appressed hirsute beneath; petioles narrowly alate by the abruptly
decurrent leaf -blades, these sometimes abruptly and acutely apicu-
late, sometimes rounded and quite obtuse, variable in size as in
shape but usually about twice longer than wide, the larger about
1.5 dm. long, entire to minutely and remotely denticulate; panicles
upper-axillary and terminal sometimes hirsute but trichomes
ascending, usually merely asperous or glabrescent, several cm. to
rarely 1.5 dm. long and wide; pedicels slender, 3-7 (10) mm. long;
petals 3-4 mm. long, yellow, cucullate; outer sepals nearly minute,
the deeply concave imbricate inner 2 indurating and developing
with the enclosed fruit, this 5 mm. in diameter. — Compare D. rugosa,
to which Standley referred all the Peruvian specimens, perhaps
correctly. Illustrated, Mart. Fl. Bras. 13, pt. 1: 3?^. 27. F.M. Negs.
9487; 23906 (D. lucida); 38710.
Junin: Brushy river bluff, 5399 (det. KilHp). Puerto Yessup,
Killip & Smith 26337; 26273.— Loreto: Iquitos, Killip & Smith'
Flora of Peru 669
26956; Williams H88; IJtOS; S570; Klug 112. To the Guianas and
Central America.
Davilla densa Macbr. Candollea 6: 15. 1934.
Liana or perhaps only the elongate branches scandent, the
branchlets, petioles (1.5-2 cm. long), leaves beneath and the extreme-
ly compact inflorescences densely brownish villous unless in age;
leaves entire or most obscurely and remotely denticulate, drying
rigid-chartaceous, shortly asperulous-hirsutulous above, the mid-
nerve densely hirsute, nearly elliptic, more or less rounded and
abruptly alate-decurrent at base, acute or shortly acuminate, mostly
8 cm. long, 5 cm. wide, sometimes nearly 2 dm. long and half as wide;
pedicels numerous, 2-4 mm. long; sepals chartaceous, brownish
pilose, developing with the enclosed globose fruit, the whole scarcely
5 mm. thick, glabrous and lustrous within. — Except for the extra-
ordinary density of the inflorescence, apparently smaller flowers
and perhaps its more nearly entire leaves it is similar to D. rugosa
Poir. to which possibly it could be referred, at most as a variety
but not seen since publication, maybe not this genus.
Loreto: Yurimaguas, Williams S826, type; KiUip & Smith 275^8.
Davilla multiflora (DC.) St. Hil. PI. Usu. no. 22: 5. 1825.
Tetracera multiflora DC. Syst. Veg. 1: 400. 1818.
Glabrous and smooth or the exfoliating branches slightly asperous
and the ovate to elliptic usually rounded entire lustrous leaves and
many-flowered inflorescences sometimes slightly pilose, the former
usually on the 7-16 prominent lateral nerves beneath; veins more
or less conspicuous both sides; panicles ample, much-branched, the
apically enlarged pedicels 8-10 mm. long; outer sepals 2-2.5 mm.
long, the lustrous inner 6 or 7 mm. long and broad, exceeded by the
entire petals. — Actually reported as noted below from adjacent
Brazil and also on the Upper Amazon but perhaps not typical.
Illustrated, Delessert, Icon. Select. 1: 69.
Rio Acre: Ule 9611 (det. Mansfeld). Rio Macauhdn, Krukoff
5359? Brazil.
Davilla rugosa Poir. Encycl. Suppl. 2: 457. 1812. D. hrasiliana
DC. Syst. Veg. 1:405. 1818.
Typically distinct from D. aspera by the much finer veins, these
not rugosely reticulate beneath but usually obvious above, and also
by the hirsute spreading pubescence of the younger branches,
670 Field Museum of Natural History— Botany, Vol. XIII
petioles, leaf-nerves and panicles, but in Peru this indument little
obvious or lacking; also the slender but prominent lateral nerves
terminate more frequently into a spinulose cusp especially toward
the acute or acutish tips or the cusp may be from a vein as the
nerves often anastomose before the margin; fruit said to attain
8 mm, in diameter. — The characters in some specimens seem to be
intangible or possibly the glabrate Peruvian plant is distinct, at
least as a variant. Probably (ex char.) D. Lechleri Rusby, Mem.
Torrey Club 6: 2. 1896, type from Bolivia but said by author to be
the same as Lechler 2316 from Peru (probably from Puno, not seen),
belongs here. Determinations by Standley.
San Martin: Juanjui, Klug 1^22. Chazuta, Klug ^056. Tara-
poto, Williams 56SU', QUOk- — Junin: Puerto Bermudez, Killip &
Smith 26526. — Loreto: Yurimaguas, Williams JtOOS; U808. Central
and northern South America.
2. TETRACERA L.
Lianas, rarely trees, the leaves more or less roughened by a
stellate silicified indument. Flowers sometimes polygamous, rarely
dioecious, mostly in terminal often lax racemes. Sepals equal or
unequal, not accrescent, typically about 5 but sometimes many.
Anthers extrorse, the cells more or less divaricate. Carpels solitary
or usually 3-5, free, each with several ovules in 2-3 series. Fruits
often radially disposed, follicular, the aril of the 1-4 seeds lacerately
extended above the seed.
Sepals glabrous within; carpel 1 T. parviflora.
Sepals sericeous within; carpels 5 T. Williamsii.
Tetracera parviflora (Rusby) Sleumer, Repert. Sp. Nov. 39:
47. 1935. T. aspera (Aubl.) Willd. var. boliviana and var. peruviana
0. Ktze. Rev. Gen. 3, pt. 2: 2. 1898. Davilla parviflora Rusby, Mem.
Torrey Club 6:3. 1896.
Verruculose-asperous, the older bark exfoliating; petioles 5-10
mm. long, margined; leaves oval to oblong-obovate, abruptly con-
tracted at base, 6-12 cm. long, half as wide, entire to mucronately
sinuate-serrate especially toward the rounded or shortly and ob-
tusely acute tip, coriaceous, the 13-16 lateral nerves parallel andj
acutely prominent beneath as the reticulate veins; panicles about
4 cm. long, lax; bracts oblong, canescent, 1 mm. long; pedicels
angled, short; buds globose, 2.5 mm. in diameter; sepals minutely
Flora op Peru 671
stellate-puberulent-scabrous without, glabrous within, about equaled
by the white petals; carpel solitary. — Apparently this could be
interpreted as a variant of T. aspera, as indicated by Kuntze. F.M.
Neg. 15497 (var. peruviana).
San Martin: Juan Guerra, Williams 6896 (det. Sleumer). To-
cache, Poeppig. — Junin: In canyon thicket. La Merced, 5^9.
Rio Peren^, Killip & Smith 25019; ^5055.— Ayacucho: Estrella,
KiUip & Smith 22676 (det. Killip). Bolivia; Brazil.
Tetracera WilHamsii Macbr. Candollea 6: 14. 1934.
Liana, the slender flexuose minutely scabrous branches much
exfoliating in papyraceous sheets; leaves elliptic or oblong-elliptic,
rounded-cuneate at base and decurrent into the 1 cm. long petiole,
abruptly caudate-acuminate, 12 cm. long, 6 cm. wide, sometimes
to 17 cm. long, coarsely undulate dentate, chartaceous, scabrous
both sides and above especially on the midnerve, sparsely hir-
sutulous underneath, conspicuously appressed asperous on the 14
nerves; inflorescence moderately paniculate, the branchlets re-
motely 3-7-fiowered, somewhat hirsutulous, with stellate and sca-
brous trichomes intermixed; flowers about 8 mm. long, the broadly
elliptic sepals 5 mm. long, glabrous or slightly asperulous without,
densely sericeous villous within; petals obovate, minutely crenulate
apically, 6 mm. long; filaments 3 mm. long, barely connate at base
and long hirsute; carpels 5, attenuate into the glabrous styles. —
Apparently related to T. Breyniana Schlecht. because of character of
sepals but the leaves strongly serrate, a not impressive character
unless there are substantiating ones; cf. also T. volubilis L., sepals
pubescent both sides, but not seen in the preparation of this account.
However, voidance of the sepal character would destroy the tra-
ditional classification, which may be, of course, incorrect.
Loreto: Rio Itaya, Williams 5U, type; also Rio Mazdn, 8126;
JosS Schunke 168 (det. Standley, T. volubilis). "Paujilchaqui."
3. DOLIOCARPUS Rolander
Lianas or less frequently not scandent, rarely tree-like, the
coriaceous leaves estipulate, the indument simple, the flowers
usually in axillary umbelliform fascicles, rarely solitary. Sepals 5
(3-6), spreading, often unequal. Petals usually 3-4 (2-6). Fila-
ments dilated apically, the small anthers extrorse, dehiscing longi-
tudinally. Carpels 1 (2), 2 (1) -ovuled. Fruit baccate, dehiscing
672 Field Museum of Natural History— Botany, Vol. XIII
irregularly if at all or sometimes capsular, the seeds enclosed by the
membranous or fleshy aril. — The always extrorse anthers are often
used as a key character in contrast to those of Curatella. Several
of the species proposed recently seem to be based on characters of
questionable significance.
Flowers solitary or in fascicles of 2-6 or so; ovary villous or hirtel-
lous except D. amazonicus.
Leaves acutely acuminate.
Flowers about 1 cm. long or longer; pedicels often longer than
3 mm.
Flowers to 1 cm. long; leaves medium D. Rolandri.
Flowers about 2 cm. long; leaves ample D. magnificus.
Flowers about 5 mm. long; pedicels 3 mm. long.
Leaves ample, to 1 dm. wide D. amazonicus.
Leaves about 4.5 cm. wide D. curtipes.
Leaves obtuse or obtusely short acuminate.
Flowers 2-4, about 5 mm. long D. aptus.
Flowers solitary, about 1.5 cm. long D. pulcher.
Flowers in most fascicles 12-30 or more; ovary glabrous.
Leaves acutely acuminate, ovate-elliptic D. dentatus.
Leaves shortly or scarcely acute, oblong-elliptic D. densifiorus.
Doliocarpus amazonicus Sleumer, Repert. Sp. Nov. 39:
44. 1935.
Nearly glabrous shrub with subterete branchlets, ample lanceo-
late-oblong leaves and small very shortly pedicellate 3-6-fasciculate
flowers; petioles more or less hirsutulous, about 6 mm. long; leaves
attenuate to the subrotund or subtruncate base, acuminate, 1.5-
nearly 2.5 dm. long, 6- about 10 cm. wide, coarsely but irregularly
serrate (serrations about 2 mm. high, 1.5-2 cm. distant) or undulate,
chartaceous, type in herb, violet-brown above, suffused with reddish-
brown beneath, opaque, not all rugose, the 13-15 lateral nerves
little impressed above, conspicuous and pilosiusculous beneath, thej
subparallel veins rather prominent; pedicels scarcely 3 mm. long;j
sepals ovate, about 4.5 mm. long, 2.5 mm. wide, sparsely but
distinctly puberulent without, nearly glabrous within, the glabrous]
obovate petals little longer; ovary ovoid, glabrous, stigma peltate.
— Type a shrub with trunk diameter of 1 dm.; differs from DA
undulatiLS Eichler in the larger leaves and shorter pedicels (author) .\
Flora of Peru 673
Loreto: Mouth of the Capanahua, Rio Blanco, Tessviann S065,
type. Florida, Klug 205j^?
Doliocarpus aptus Macbr. Candollea 6: 13. 1934.
Scandent, the terete branches glabrous, the short branchlets
puberulous; petioles tjrpically about 2.5 cm. long; leaves elliptic-
obovate, broadly cuneate-acute at base, obtusely short-acuminate
or subobtuse, mostly about 1 dm. long, 6-7 cm. wide or smaller,
sometimes 1.5 dm. long, 8 cm. wide, entire, coriaceous, scarcely
lustrous, glabrous above, sparsely pilose-puberulent beneath (as the
petioles) with short and longer trichomes intermixed, the 12 or so
lateral nerves with the more or less reticulate veins prominent
beneath, scarcely visible above; pedicels 2-4-fasciculate, 2-2.5 mm.
long; petals greenish white, suboblong, 5 mm. long, pulverulent
without, glabrous within; filaments to 7 mm. long, little flexuose;
ovary as style densely brownish pilose, the stigma capitate-peltate.
— Var. curtus Macbr. I.e. is striking by the short (5-10 mm. long)
petioles, leaves mostly 7 cm. long, about 5 cm. wide, pedicels
scarcely 2 mm. long. The liana is evidently comparable to D.
dentosus Mart. (D. major Gmelin, fide Lanjouw) from which it
differs notably in its entire not at all acuminate leaves, pubescent
beneath, shorter pedicels and pubescent style, the indument of the
ovary much longer, but not seen in preparation of this final com-
pilation.
Loreto: Mishuyacu near Iquitos, Kliig 977, type; 714. (type, var.).
Doliocarpus curtipes Macbr. Candollea 6: 13. 1934.
Scandent, the branches sparsely pilose-hirsutulous; petioles 3-4
mm. long; leaves ovate-elliptic, acute at base, remotely and coarsely
mucronate-serrate toward the acuminate tip, mostly 1 dm. long,
4.5 cm. wide, membranous, minutely and sparsely lepidote both
sides, slightly pilose beneath on the nerves; flowers 1-4-fasciculate
in the axils, 5-6 mm. wide; pedicels 3 mm. long, densely short-
villous; sepals both sides, especially without, lightly sericeous, the
white petals densely so at base within, 5 mm. long, about equaled
by the filaments, the anthers oblong; ovary shortly villous, the style
glabrous. — Related to D. dentosus Mart. {D. major Gmelin, fide
Lanjouw) but pedicels much shorter, leaves definitely pubescent;
therefore, lacking any marked characteristic it could be treated as
a variant; however, not seen since publication.
Loreto: Mishuyacu near Iquitos, Williams 771, type.
674 Field Museum of Natural History— Botany, Vol. XIII
Doliocarpus densiflorus (Eichler) Steyerm. Fieldiana: Bot. 28:
367. 1952. D. brevipedicellatus Garcke var. densifloriis Eichler in
Mart. Fl. Bras. 13, pt. 1: 81. 1863.
Probably a shrub, glabrescent, the densely leafy flowering
branchlets nearly to tips almost filled continuously from node to
node with the compact fascicled numerous small flowers; indument
deciduous, early long-pilose on the stout canaliculate petioles, these
almost 1 cm. long, and leaf midnerve and 12 or so prominent lateral
nerves beneath; leaves ovate- or oblong-elliptic, shortly acute or
acutish, mostly 8-12 cm. long, about 4 cm. wide, coarsely and
remotely repand-serrate except toward base, slightly lustrous and
obviously veiny both sides, coriaceous, drying dark but paler
beneath; pedicels laxly puberulent-pilose, about 7 mm. long; sepals
medially puberulent, membranous in flower, the larger about 4 mm.
long and wide, little exceeded by the petals; filaments suberect in
bud, anthers ovoid; fruit unknown. — Gleason, Bull. Torrey Club
58: 388. 1931, avoided citing place of publication of species name
in giving Sprague & Williams as authority for above transfer and
therefore did not make it for them as indicated in Gray Herb. Index.
Actually both the author who published it and I had also used it
in herbaria.
San Martin: Tarapoto, Spruce U567, tjrpe; Klug 1551. To
Venezuela and Guiana.
Doliocarpus dentatus (Aubl.) Standley, Journ. Wash. Acad.
Sci. 15: 286. 1925. Tigarea dentata Aubl. PI. Guian. 2: 290. pi 351. {
1775. D. Rolandri in lit. not Gmelin, fide Sandwith. D. brevi-
pedicellatus Garcke in Linnaea 22: 47. 1849?
Shrub-tree or the few slender branches scandent, slightly
sericeous at the growing tips as the petioles (1-2.5 cm. long) anc
leaves, or these more or less continuously appressed pilose aboveJ
and especially on the 10-18 prominent lateral nerves beneathJ
ovate-elliptic or somewhat obovate, narrowed or acute at base,]
acutely acuminate, mostly 1-1.5 dm. long, about half as wide,]
chartaceous-coriaceous, slightly serrate above the middle, thej
nerves obscure above; flowers usually 15-30-fascicled, the slender
pedicels 6-10 mm. long or longer, puberulent as the 4 or 5 oblonj
to suborbicular, obtuse sepals, these to 5 mm. long or about as lonj
as the 2 or 3 obovate unguiculate petals; filaments early suberect^
anthers short; carpel subglobose, glabrous, style about 1 mm. longj
berry 6-7 mm. in diameter, reddish, the 1 or 2 seeds lustrousJ
Flora of Peru 675
fuscous. — The numerous small flowers and the glabrous carpels
suggest that Garcke's species, if not distinct, is a part of D. denkUus
rather than D. Rolandri.
San Martin: Zepelacio, Klttg 87 U8 (det. Standley). — Hudnuco:
Monz6n, Weberbauer Sj^90. — Junfn: La Merced, 5578 (det. Sprague).
— Loreto: On the Ucayali, Tessmann SJt.U9. Rio Nanay, WiUiams
975; 1210. To Central America and the West Indies.
Doliocarpus magnificus Sleumer, Repert. Sp. Nov. 39: 45.
1935.
Beautiful liana with ample smooth foliage and large solitary or
few-fasciculate whitish flowers, the obovate petals to 2 cm. long and
half as wide; petioles obviously canaliculate, about 1 cm. long, 2 mm.
thick; leaves coarsely and irregularly undulate-serrate or merely
-mucronulate, obovate-oblong, long-cuneate to base, acutely acu-
minate (acumen to 2 cm. long), 8-11 cm. wide or wider, 2.5-3.5
dm. long, chartaceous, not at all rugose, minutely lepidulose beneath
(nearly soft to touch), the 17 or so prominent (beneath) lateral
nerves pilose-puberulent both sides, the vein-reticulation beneath
notably elevated; pedicels about 3 mm. long, 1.5 mm. thick; sepals
orbicular-obovate, densely appressed sericeous both sides, 12 mm.
long, 7 mm. wide; filaments nearly 1 cm. long; ovary conical, densely
yellowish sericeous. — Allied to D. grandiflorus Eichl. (sect. Othlis)
but exceptional in size of leaves and flowers. Probably will be
found in adjacent Peru.
Rfo Acre: Seringal Auristella, Ule 9610, type. Bolivia.
Doliocarpus pulcher Sleumer, Repert. Sp. Nov. 39: 46. 1935.
Smooth (to touch) liana with terete glabrate branches, pubescent
branchlets, large solitary flowers and ample elliptic or ovate-elliptic
chartaceous leaves, these glabrous above, pilose-puberulent on the
9 or so prominent lateral nerves beneath, entire or undulate, some-
what rounded or broadly cuneate at base, obtuse, the petioles
2.5-4.5 cm. long; pedicels 12 mm. long, 2 mm. thick, densely ap-
pressed gray strigose; sepals obovate-oblong, sericeous only without,
13-15 (-20) mm. long, 7-9 mm. wide, the obovate-spathulate green-
ish-white petals about 1.5 cm. long, 8-10 mm. wide; filaments
glabrous, to 5 mm. long; ovary as style densely hirsute as the
globose fruit. — Distinguished from D. aptus by the larger fewer-
nerved leaves and three times as large flowers (author), the type
to 20 meters tall with trunk to 3.5 dm. in diameter.
i
676 Field Museum of Natural History — Botany, Vol. XIII
Loreto: Flood-free wood at mouth of the Santiago, Tessmann
4128, type; also 4^18.
Doliocarpus Rolandri Gmelin, Syst. Nat. 2: 805. 1791. D.
major Gmelin, I.e. D. dentosus Mart, in Flora 24, App. 2: 65. 1844,
fide Sandwith, Kew Bull. 171. 1931.
Branchlet tips and petioles early puberulent, the branches
asperulous, exfoliating and glabrescent in age; petioles 4-11 mm.
long, sulcate above, narrowly alate by the decurrent limb, this more
or less elliptic, often spinulosely serrate toward the acutely acu-
minate apex, 5-13 cm. long, 2-6 cm. wide, sometimes merely repand
or even entire, coriaceous, above sparsely, beneath densely lepidote
(scales early with a central trichome), the midrib and 6-10 lateral
nerves prominent both sides, the latter (as the former also above)
appressed pilose beneath; pedicels 2-6, usually 3-8 mm. long, densely
puberulent as the buds and suborbicular sepals both sides, these
to 7 mm. long, 5 mm. wide; petals 3 or 4, broadly obovate, to 9 mm.
long and wide, puberulent and glabrescent without; filaments
recurved in bud, anthers oblong; carpel 1, hirtellous as the 11 mm.
thick fruit, the 2 seeds compressed, to 7 mm. long, 5 mm. wide,
nearly black. — After Lanjouw who questioned the identity of
Garcke's plant, which ex char, seems to be nearer D. dentatus
(i.e. D. Rolandri in lit.). Apropos this latter name, misinterpretation
of a name's identity is no reason for discarding it. Var. parous
Macbr. has leaves to 2 dm. long, nearly 1,5 dm. wide, sparsely
undulate-denticulate, softly pilose beneath, the petioles 2-3 cm.
long, the 2-5-fasciculate pedicels 5-8 mm. long, differences probably
not particularly significant, the type according to the collectors
a tree 10 meters high. Not seen since pubHcation; probably, ex
char., goes to D. dentatus. F.M. Negs. 19165; 19166 (var.).
Loreto: Yurimaguas, Killip & Smith 2754-5 (type, var.). Central
and South America.
4. GURATELLA L.
Shrubs or small trees, the estipulate leaves scabrous with stellate ,
silicified trichomes, the short panicles of white flowers typically
on older branchlets at the leaf-scars. Sepals imbricate, 4 or 5,
spreading, subequal, the petals as many. Stamens persisting, the;
filiform filaments apically dilated, the anthers usually introrse,]
dehiscing longitudinally. Carpels 2, cohering basally within.
Flora of Peru 677
Fruits 2 follicles in each flower, each with 2 seeds subenclosed in the
membranous striate aril.
Curatella americana L. Sp. PI. ed. 2. 748. 1762.
Stout shrub or tree to 12 meters, with rough exfoliating brown
bark, gnarled densely scaly branches, the young branchlets early
pubescent, and short panicles of fragrant white flowers commonly
from the axils of fallen leaves; j>etioles (alate, rarely 1 cm. long) and
leaves coated, the latter especially above, with almost minute harsh
stellate trichomes; leaves variably elliptic but always decurrent
from rounded or emarginate base and rounded-emarginate or obtuse
at apex, often a dm. or two long, about half as wide, the 8-18 lateral
nerves terminating in short teeth, prominent beneath; panicles 3-7
cm. long, closely pubescent including the pedicels (5-8 mm. long)
and the 4 (or 5) roundish sepals both sides, these to 6 mm. long,
reflexing; petals obovate, 6-7 mm. long, often denticulate, deciduous;
carpels pubescent with long erect trichomes, the hispid globose
follicles to 8 mm. thick. — Illustrated, St. Hilaire, PI. Usu. pi. 24,
(as C. Cambaiba) and F*flanzenfam. ed. 2. 21: 19.
The sandpaper-like leaves are used for polishing (Weberbauer) ;
abundant, sometimes forming thickets on the sandy plain of Tara-
poto (Williams).
San Martin: Tarapoto, Ule 61*24 (det. Perkins); WiUiams 5677.
Moyobamba, Weberbauer 4578; 290. — Hudnuco: Chicoplaya, Ruiz
& Pavdn. To the West Indies and Central America. "Ractapanga"
(Weberbauer).
ACTINIDIACEAE
Much like Dilleniaceae but the pistil solitary, placentation
axillary, ovules with a single integument, many in each cell, the
styles usually as many as carpels, generally persistent; fruit a tardily
dehiscent berry or leathery capsule, the seeds non-arillate with
usually large embryos and abundant endosperm. — There is probably
no gain from the standpoint of usefulness in separating this family;
the unit as a whole is said probably to be related to the Ranuncu-
laceae; the segregate has been also called Saurauiaceae; cf. Lawrence,
Taxonomy Vase. PI. 599. 1951.
SAURAUIA Willd.
Palaua R. & P. Prodr. 100. pi, 22. 1794. Apatelia DC. Prodr.
1: 526. 1824.
678 Field Museum of Natural History— Botany, Vol. XIII
References: Buscalioni, Malpighia 25-30. 1912-1927; key, 25:
218-225; index, 30: 436-444; 25 & 26 by Buscalioni & Muscatello;
Gilg & Werdermann, Pflanzenfam. ed. 2. 21: 44. 1925, after Diels,
Bot. Jahrb. 57:441. 1922.
Character of the family: The most recent account of these
strikingly attractive plants is lacking in organization and pre-
sentation, but one may admire the author's inherent ability and
interest, which prompted his studies, probably under difficulties.
In any case specific characters to this day are not understood and
a satisfactory treatment will probably be possible only with cyto-
logical as well as morphological data. Very apropos are the following
observations of A. C. Smith, Joum. Arnold Arb. 22: 502. 1941,
prompted by his studies of Papuasian species, which may be sum-
marized here since they apply well to the Peruvian plants and may
serve as a guide for field notes of collectors.
Smith agrees with Burtt, Hook. Icon. 34: pi. 3316. 1936, that
position and branching of inflorescence, degree of bract development,
pubescence of leaves beneath and other characters are not funda-
mental but for expediency are at present more or less usable, while
probably the type of indument may prove to be basic in indicating
true relationship; characters which are fairly stable (i.e., type,
direction and length of hairs, shape of scales, size of flowers, number
of stamens, their arrangement as that of styles, etc.) have not
always been adequately described. Smith further notes that the
abundant and obvious fusiform crystal-bearing cells (containing
both raphides and styloids) in leaves, petals and ovaries of some
species may be helpful in differentiating specific character. — The
name is after one von Saurau and written Saurauia by Willdenow
in his herbarium. There is therefore no justification for the spelling
Saurauja by some authors.
Besides the following, some of the Colombian species proposed
by R. E. Schultes, Caldasia 2, no. 6: 27^5. 1943, may occur in
Peru, especially those from Putumayo, and several of the glabrate
Ecuadorian species, similar to the earlier described Peruvian S.
glabrata, sens, lat., are to be expected. Also omitted from the key
is S. isoxanthotricha Busc, origin unknown.
Branches (flowering) glabrous or soon glabrate, the very short setae
or indument sparse.
Indument lacking or nearly on mature leaves or confined to
nerves.
Leaves ample, obovate.
Flora of Peru 679
Indument, except on younger parts, nearly lacking.
S. glabrata.
Indument sparse but subpersisting S. Weberbaueri.
Leaves oblanceolate, about 3 cm. wide S. Briqueti.
Indument setulose-puberulent to tomentose on mature leaves
beneath.
Leaves simply or minutely pubescent beneath.
Inflorescence densely spreading setulose.
Leaves setulose beneath S. scabriuscula.
Leaves except nerves, puberulent furfuraceous beneath.
S. pseudoscabra.
Inflorescence minutely appressed setulose S. peruviana.
Leaves tomentose-stellate beneath S. Raimondiana.
Branches notably rather persistently long-setose-barbate, some-
times laxly, sometimes also densely scurfy-tuberculate.
Indument of leaves confined to the nerves.
Leaves ample, petioles elongate; flowers 7 mm. wide.
S. natalicia, S. Loeseneriana.
Leaves medium, petioles 1 cm. long; flowers 5 mm. wide.
S. solitaria.
Indument of leaves sparse, setulose on nerves and veins, lax on
upper stems S. Prainiana.
Indument of leaves not restricted to veins beneath, scabrous-
setulose above.
Stamens 30-40, perhaps more, but at most 80.
Leaves moderately bullate above only in age, setulose.
iS". biserrata.
Leaves very bullate and conic-tuberculate above, reticulate
beneath S. formosa.
Stamens 130-140; indument of S. biserrata S. roseotinda.
Saurauia biserrata (R. & P.) Spreng. Syst. 4. Cur. Post 211.
1827. Palaua biserrata R. & P. Syst. Veg. 181. 1798. Apatelia
biserrata (R. & P.) DC. Prodr. 1: 526. 1824. P. lanceolata R. & P.
I.e. A. lanceolata (R. & P.) DC. I.e., not S. lanceolata DC. Mem.
Soc. Phys. Hist. Nat. Gendve 1: 421. 1822. S. Ruiziana Steud.
Nom. end. ed. 2. 2: 516. 1841, nomen. A. lanceolata (R. & P.) DC.
var. peduncularis DC. I.e. S. scabra Poepp. var. Prainiana Busc.
680 Field Museum of Natural History — Botany, Vol. XIII
Malpighia 27: 493. 1916. S. Ruiziana Steud. var. peduncularis (DC.)
Choisy, Mem. Soc. Phys. Hist. Nat. Geneve 14: 116. 1855.
A 5 meter shrub, the branchlets, petioles and peduncles long-
setose bearded; leaves oval-lanceolate-oblong, attenuate at both
ends, subacute to acuminate, about 1 to sometimes 3.5 dm. long and
5-12 cm. wide, rather densely appressed setulose above, a little
rough but scarcely scabrous to the touch, densely crisply villous
beneath; peduncle axillary, 3-4 times longer than the petiole,
villous-setose; racemes short; flowers crowded, large, pedicellate;
sepals glabrous within, obovate, obtuse; petals pilose within at base
as also the filaments; ovary glabrous; styles filiform. — S. pseudo-
ruitziana Busc, Malpighia 30: 244. 1927, is probably included in the
above description. It is based on a specimen by Pearce, Herb. Kew,
of doubtful origin, and by the author is distinguished from true
5. Ruiziana by the 80 stamens, setulose upper leaf surfaces, by
the 25 instead of 23-26 nerves, shorter (1 cm.) inflorescence branches,
their setae 4-5 mm. long instead of 8 mm. F.M. Negs 23915; 23920
(S. lanceolata);2S9S4:.
Hudnuco: Pillao and Mufia, Ruiz & Pav6n, type; Woytkowski
3U178. Huacachi near Mufia, U81. Cuchero, Poeppig 1607. —
Ayacucho: Choimacota Valley, Weberhauer 75J^5.
Saurauia Briqueti Busc. Malpighia 30: 140. 1927.
Branchlets reddish, glabrous or the tip minutely tuberculate and
puberulent; petioles 1-1.5 cm. long, glabrate or tuberculate-setulose
under a lens; leaves oblanceolate, about 9 cm. long, 2.5-3 cm. wide,
scabrous-granulose above but glabrous, similarly scabrous beneath
and minutely puberulent; secondary nerves about 15, scarcely
visible above, fine and slightly reticulate beneath; inflorescences
about as long as leaves, glabrous or the branchlets minutely and
sparsely puberulent-setulose; peduncle 3-4 cm. long, glabrous or
nearly; pedicels slender, short; bracts 2-4 mm. long, linear; flowers
8-10 mm. broad, calyx lobes about 4 mm. long, petals much longer;
stamens 20 or fewer, styles short. — Allied by author to S. Weber-
baueri but there is no more difference in leaves than shown in type
specimens of S. scabriv^cula. A 4 meter shrub. Keyed, I.e. 25:
224. 1912. Illustrated, I.e. 28. pi 2, fig. 26. F.M. Neg. 5750.
Junin: Huacapistana, 2,500 meters, Weberbauer 2177.
Saurauia formosa Sleumer, Notizbl. Bot. Gart. Berlin 12: 144.
1934. S. Ruiziana Steud. var. Weberbaueri Busc. Malpighia 26:
6. 1913.
Flora of Peru 681
Indument of the stout branchlets, lateral leaf-nerves, these about
25, and ample axillary inflorescences densely femigineous strigose
the trichomes to 3 or 4 mm. long and rigid; petioles 5-11 cm. long;
leaves obovate-elliptic, cuneate at base, gradually acuminate, 3-4
dm. long or longer, nearly half as wide, densely conic-tuberculate
above (asp)erous), beneath especially on the densely reticulate
elevated veins closely stellate, soft to touch, the thickened incurved
margin minutely mucronulate, the incurved mucro 1 mm. high;
panicles elongate, cymosely divided, to at least 3 dm. long (peduncles
to 1.5 dm. long, 5-6 mm. thick); bracts densely strigillose, long-
deltoid, 6-8 mm. long; flowers 22 mm. broad; calyx lobes 5, obovate-
elliptic, obviously strigose, 7-8 mm. long, 4 mm. wide, the oblong-
spatula te petals ciliate, about 11 mm. long, 4 mm. wide; interstaminal
trichomes rufous; ovary glabrous. — Number of stamens unknown.
A beautiful species, 4 meters tall, of the Section Strigosae Busc., in
vicinity of S. Ruiziana (author). Illustrated, Busc. I.e. 25. pi. 5,
fig. 7 (his var. of 5. Ruiziana).
Junin: Rio Portachuelo near Calabaza, 2,800 meters, Weberbauer
66S7, type. Above Huacapistana, Sandwith U563; Weberbauer 20U0
(tjrpe, S. Ruiziana var. Weberbaueri), near San Ram6n, Hodge 6252.
— Huancayo: Oxapampa, Soukup 2JtSS? — Hudnuco: Pampayacu,
Sawada 53.
Saurauia glabra ta (R. & P.) Choisy, M^m. Soc. Phys. Hist.
Nat. Geneve 14: 119. 1855. Palaua glabra R. & P. Syst. Veg. 182.
1798.
Completely glabrous (type) except the branchlet tips, petioles
(to 3 cm. long), and the large inflorescences, these with peduncles
to 4 cm. long or longer, open, few-branched, minutely scurfy puberu-
lent; leaves elliptic-obovate, somewhat narrowed to base, acutely
short-acuminate or cuspidate, finely serrulate, to about 2 dm. long,
7 cm. wide; pedicels 4-6 mm. long; flowers about 1 cm. across;
stamens apparently about 20. — In all probability several of the
segregate species (as, in Peru, S. Briqu£ti, S. Weberbaueri) including
also S. pseudoparviflora Busc. from Colombia and Bolivia and S.
Rusbyi Britt. of Bolivia should be compared here as local develop-
ments if not variants. F.M. Neg. 23918.
Hudnuco(?) : Ruiz & Pav6n, type.
Saurauia isoxanthotricha Busc. Malpighia 25: 410. pi. 5,
fig. 6. 1913.
682 Field Museum of Natural History— Botany, Vol. XIII
Branchlets fistulose-sulcate, clothed toward the apex with very-
long reddish setae; petioles to 3.5 cm. long; leaves coriaceous, softly
setose both sides, especially on the 18 nerves, lanceolate-obovate,
rounded-acute at base, acute or acuminate, 13-15 cm. long, 5 cm.
wide, the slightly irregular serrations setulose; panicle branched,
setose-scabrous, about equaling the leaves, the linear bracts 1 cm.
long; flowers subsessile; calyx pubescent within and without; styles
obsolete; ovary glabrous. — Unfortunate that the locality of any of
the earlier names among several close forms should be unknown;
from the illustration it seems to be near S. Prainiana among Peruvian
species.
Peru(?): Without locality, (without collector, 1216, Herb. Kew).
Saurauia Loeseneriana Busc. Malpighia 25: 399. 1913.
Smooth except for the rather sparse straggling long setae on the
stems and petioles and the short appressed ones on the leaf -nerves
beneath; leaves membranous, paler beneath, except for the midrib
glabrous or nearly so above, long obovate-lanceolate, cuneate at
base, shortly acuminate, 3 dm. long or longer, 8-11 cm. wide,
mucronulate-denticulate; lateral nerves 25-30, about 1 cm. distant;
inflorescence densely long-setose, few-flowered, little if at all ex-
ceeding the 3-4 cm. long petioles; bracts lanceolate, the upper
triangular; flowers subsessile, medium; calyx 3 mm. long, puberulent
throughout; stamens pilose at base; ovary glabrous. — Illustrated,
I.e. 28. pi. 2, fig. 28. F.M. Neg. 5752.
Cajamarca: Tambillo, Jelski 259. Arenales, Prov. Cutervo, Stork
& Horton 10161 (det. Standley, S. Ruiziana).
Saurauia natalicia Sleumer, Notizbl. Bot. Gart. Berlin 12:
144. 1934.
Branchlets laxly pubescent with ferrugineous setae, the indument
of the stout petioles (1-2.5 cm. long), leaf midnerve especially
beneath, and axillary cymose-paniculate inflorescence laxly (or the
last appressed) strigose; leaves elliptic-oblong, long-attenuate to
base and more or less so to apex, 9-15 cm. long, 3.5-4 cm. wide,
glabrous (except midnerve), membranous, somewhat and irregularly
serrate-dentate, the lateral nerves 16-24; peduncles 5-8 cm. long;
flowers 1.5 cm. broad; bracts minute; calyx lobes 5, obovate-oblong,
4 mm. long, nearly half as wide, glabrous; petals white, obovate-
elliptic, 7 mm. long, 3.5 mm. wide; ovary glabrous. — There is a light
pilosity at insertion of stamens; next to S. Loeseneriana Busc. with
larger leaves and flowers, more strigose inflorescence (author).
Flora of Peru 683
Ayacucho: Choimacota Valley, Weberbauer 7567; 7567a. — Cuzco:
Near Chile-Chile, Valle de Marcapata, 2,500 meters, Weberbatier
7867, type.
Saurauia peruviana Busc. Malpighia 27: 319. 1916. S. flori-
bunda Benth. var. peruviana Busc. Malpighia 28: 119. 1917?
Branchlets more or less appressed setulose and stellulate
pubescent, the 2-5 cm. long petioles tuberculate and stellate-puberu-
lent; leaves chartaceous, glabrate but lightly scabrous above, setu-
lose-puberulent beneath, obovate or oval-lanceolate, acute or
acuminate, the base cuneate, mucronately and setosely serrulate,
2-3 dm. long, 7-10 cm. wide; inflorescence shorter than the leaves,
few-flowered, the slender peduncles and branchlets furfuraceous and
minutely appressed setulose; bracts minute; calyx rufescent, gla-
brous within, little shorter than the petals; stamens 25-30; styles
elongate. — The variety proposed for the Ecuadorian species of
Bentham seems better placed here, ex char., stems, petioles and
inflorescence puberulent-tuberculate, leaves granulose above, mi-
nutely setulose and stellulate beneath. Illustrated, I.e. 25. pi. 8,
fig. 15. F.M. Neg. 19162.
Amazonas: Chachapoyas, {Mathews, type, S. floribunda var.).
Without locality, Haenke, type; Mathews.
Saurauia Prainiana Busc. Malpighia 25: 248. pi. 6, fig. 10. 1912.
Branchlets, petioles and inflorescence barbate-setose, not at all
or very obscurely puberulent; leaves oval or little obovate, about
2 dm. long, 6-10 cm. wide, usually obtuse at base, acute or shortly
acuminate, chartaceous, minutely and sparsely scabrous above and
setulose on nerves, paler and lightly setulose on all the veins be-
neath, very minutely mucronate-denticulate; lateral nerves 16-18,
about a cm. distant; axillary racemes sometimes a dm. long; bracts
linear-lanceolate, 4-5 mm. long; flowers subsessile, about 7 mm.
wide, the oval sepals setulose without, the slightly longer oblongish
petals about 7 mm, long; stamens about 25; ovary glabrous. — A 5-
10 meter tree. F.M. Neg. 5753.
Loreto: Cerro de Escaler, Ule 6530, type. Pumayacu, Balsa-
puerto to Moyobamba, Klug 3176 (det. Standley).
Saurauia pseudoscabra Busc. Malpighia 28: 1. 1917.
Stem tips and petioles (1.5-2.5 cm. long) spreading setulose with
trichomes in part 3-5 mm. long; leaves finely serrulate, coriaceous,
684 Field Museum of Natural History— Botany, Vol. XIII
decurrent to base, acute or acuminate, 1-2 dm. long, 4-6 cm. wide,
minutely scabrous-setulose above, beneath the 13-14 prominent
nerves setulose, the surface somewhat furfuraceous or softly setose;
panicles many-flowered, subequaling the leaves, the stout peduncles
7-10 cm. long, branches 1-2.5 cm. long, bracts 3-6 mm. long;
flowers subsessile; calyx lobes 3-4 mm. long, glabrate or minutely
and sparsely setulose; stamens 25; styles 5, long. — May be a part
of S. scabriuscula but there seem to be distinctions of possible
significance; the stem pubescence suggests that of the more northern
S. tomentosa HBK. with large flowers. F.M. Neg. 5756.
Puno : Sandia to Cuyocuy o, 2,300 meters, ( Weberbauer 8Jf2, type) .
Saurauia Raitnondiana Sleumer, Notizbl. Bot. Gart. Berlin
12: 145. 1934.
Characterized especially by the presence of three types of in-
dument on the leaves, a tomentosity of very short branched tri-
chomes on petioles (3.5 cm. long), midrib and the 16-19 lateral
nerves beneath, ferrugineous stellate trichomes on the leaf -surfaces
apart from the principal nerves, but lanuginose-barbate in their
angles with the midrib; leaves cuneate-oblong, gradually acuminate,
1.5-2.5 dm. long, 4.5-8 cm. wide, chartaceous, densely and obtusely
serrulate; panicles many-flowered, about half as long as the leaves,
more or less densely pubescent with shortly branched trichomes;
flowers about 1.5 cm. broad; calyx lobes 5, ovate-elliptic, 5 mm.
long, 3 mm. wide, sparsely stellate-puberulent dorsally; petals
nearly subrotund, 7 mm. long, 6 mm. wide; stamens many, 5 mm.
long; ovary glabrous. — On account of the tomentose indument
ought to belong to the Tomentosae Busc. of Section Ditrichae Busc.
but the presence of the stellate trichomes places it at a distance
(author). Var. caxamarcensis Sleumer, I.e. 146, has leaves 12 cm.
long, 3.5^ cm. wide, tomentose as panicle, this as long and the
flowers apparently a little smaller.
Cajamarca: Montana de Nancho, Raimondi U758, type; also
66U; 5SH (type, var.), 7331.
Saurauia roseotincta R. E. Schultes, Bot. Mus. Leafl. Harvard,
16: 83. 1953.
Slender open tree, the leaves — these only on the upper part —
pink-red pubescent beneath, the white flowers not abundant; petioles
stout, to 3.5 cm. long, 4-5 mm. thick, coarsely as the inflorescence,
this about 2 dm. long, rufous- or ferrugineous-setose; leaves elliptic,
Flora of Peru 685
minutely denticulate, cuneate basally, subacute, 21-34 cm. long,
7-11 cm. wide, subcoriaceous, bullate and asperous-setose above,
especially toward the nerves, softly and minutely white stellate-
pilose all over beneath and all the nerves densely roseate stellate-
pilose; bracts setose-pilose, subulate, mostly 5 mm. long; flowers
13 or 14 mm. across, the rounded subentire petals 6-7 mm. long,
&-6 mm. wide; sepals 5, the inner rhomboid-ovate, 9-10 mm. long
and wide, setose-carinate without, puberulent within toward obtuse
apex as the thicker subacute outer, these strongly golden without,
5-6 mm. wide; filaments about 130, rufous setose below, 23 mm.
long, the anthers small; ovary globose, 4 mm. in diameter, glabrous,
the 5 styles to 1.5 mm. long. — Near S. biserrata (R. & P.) Spreng.
{S. Ruiziana) and relatives with only a few stamens (author).
Well-named, the type about 3 meters high.
Hudnuco: Chaglla, S652, tjrpe.
Saurauia scabriuscula Macbr., nom. nov. S. scabra Poepp.
ex Choisy, M^m. Soc. Phys. Hist. Nat. Geneve 14: 118. 1855,
not HBK.
Branchlets, petioles and panicles harshly pubescent and reddish
tomentulose; petioles 1.5-3 cm. long, stout; leaves coriaceous or
chartaceous, green but scabrous and minutely setulose above, rather
densely reddish setulose beneath and rather softly so, obovate-
lanceolate or oblanceolate, mostly 7-16 cm. long, (3) 4.5 (-6) cm.
wide, barely acute, cuneate at base, most minutely denticulate;
flowers tiny, about 2.5 mm. wide; calyx glabrous; ovary glabrous;
styles long; stamens 20-25, the anthers terminally dehiscent but by
a cleft instead of the usual pore (Gilg). — Reluctantly I give this
a new name because of the mechanical necessity; there is doubtless,
sens, lat., a synonym which will eventually supply the need but
ex char, none of the Peruvian species seem to be quite the same and
until the group is revised it seems better not to apply a name already
in use, not positively applicable. F.M. Negs. 5754; 23294.
Hudnuco: Cuchero, Poeppig 16H. Chinchao, Sawada 97.
Saurauia solitaria Sleumer, Notizbl. Bot. Gart. Berlin 12:
147. 1934.
Densely leafy branchlets strigose with trichomes to 3 mm. long;
petioles 1.5 cm. long; leaves cuneate-oblong, shortly acuminate, 16-
23 cm. long, 5-6.5 cm. wide, chartaceous, glabrous except the strigose-
pilose midnerve both sides and the 14-16 lateral nerves, these
686 Field Museum of Natural History— Botany, Vol. XIII
sparsely strigose beneath; marginal serrations elongate- triangular,
strigillose at tip, about 2 mm. high; panicles about a third as long
as the leaves, rather densely many-flowered, the peduncles 7 cm.
long; flowers 8 or 9 mm. across; bracts minute; pedicels (as pe-
duncles) closely ferrugineous strigose; calyx lobes ovate-elliptic,
3 mm. long, 1.8 mm, wide, glabrous; petals roseate, elliptic-rotund,
4-4.5 mm. long, 2.5 mm. wide; ovary glabrous. — Section Strigosae
Busc, without apparent relationship (author). Type a shrub 2
meters high.
Libertad: Valley of the Mixiollo, Prov. Pataz, about 1,800
meters, Weherhauer 7067, type.
Saurauia Weberbaueri Busc. Malpighia 30: 148. 1927.
Allied to S. Briqueti but distinctive in its much larger cuneate-
obovate leaves, 1.5-2 dm. long, 6-8 cm. wide, on long petioles, ample
(nearly 2 dm. long) inflorescence with pubescent peduncle, evident
linear bracts, and 4 instead of 5 styles. — As to this character the
number of styles in some species apparently varies; it seems probable
that it is a part of S. glabrata if not also of S. Briqueti. A shrub.
Illustrated, I.e. 25. pi. 5, jig. 8. F.M. Neg. 5762.
Amazonas: Chachapoyas, 2,600 meters, Weberbauer JfJfSJt.
OCHNACEAE
References: Gilg, Pflanzenfam. ed. 2. 21: 53-87. 1925; Engler
in Mart. Fl. Bras. 12, pt. 2: 297-366. 1876; 13, pt. 1: 397^20. 1871.
Rarely herbs or half-shrubs, commonly simple-leaved plants,
the leaves always stipulate (stipules often deciduous), alternate,
glabrous and usually with numerous pinnate nerves. Flowers
hermaphrodite, variously diposed, sometimes solitary, the sepals
and petals free, imbricate or rarely contorted, especially the latter,
these 5 (4-10), the former 4-5 (rarely 10). Stamens inserted at
base or above a disk or torus, few to many, distinct, sometimes
accompanied by free or connate staminodia, these subulate or
petaloid, the filaments persisting; anthers linear, basifixed, opening
by a pore or longitudinally. Ovary entire to deeply lobed, 1-10-
celled, the 1-many ovules variously attached; style 1, rarely parted
apically. Fruit an elongated capsule and septicidal or drupaceous,
the carpels connate or finally often distinct on the enlarged torus.
Seeds 1-many, with or without endosperm, this character serving
as the basic division of the family; however, the following artificial
key seems to distinguish the few Peruvian genera.
Flora of Peru 687
The last 3 genera in the following key have sessile radiating
stigmas (Dwyer) and as in Godoya stamens are in one row.
Suffrutescent herbs or slender stemmed shrubs with notably pecti-
nate-ciliate stipules 2. Sauvagesia.
Shrubs or shrub-trees, the stipules deciduous or inconspicuous.
Leaves oblong-lanceolate or -elliptic, sometimes basally acute, but
not narrowed to base unless also to apex 1. Ouratea.
Leaves obovate or spathulate, distinctly narrowed to base,
broadest above the middle or rounded-obtuse.
Leaves long-spatulate, several times longer than wide; stamens
many; stipules persisting 3. Cespedesia.
Leaves obovate, about twice longer than wide; stamens 10;
stipules deciduous.
Flowers about 2 cm. long; leaves crenate; sepals glandular
within 4. Godoya.
Flowers about 1 cm. long; leaves minutely appressed denticu-
late; sepals not glandular 5. Krukoviella.
1. OURATEA Aublet
Glabrous shrubs or trees with evergreen lustrous serrulate or
entire leaves, the small stipules deciduous, and pretty yellow or
sometimes reddish tinged flowers usually in bracteate terminal
or axillary panicles or racemes. Sepals and petals 5, the former
often coherent and persisting, the latter little longer, ordinarily
obovate and clawed. Stamens 10, the very short filaments from the
base of a disk, the rugose anthers opening from a terminal pore.
Ovary 5-10-lobed ; styles united ; ovules solitary. Drupes 5 or fewer
on the expanded fleshy disk, the seed erect.
The standing of the species proposed in this large group which
has had no revision for a long time is in many cases open to question
and the key naturally is only suggestive; the traditional separation
of species on the basis of leaf-venation seems often intangible or
possible of diverse interpretation; under these circumstances it has
not been within the scope of this work for me to try evaluation of
the species proposed for Peru particularly as regards their relation-
ship to extra Peruvian entities, nor to dispose of all collections.
Veins innumerable, mostly parallel, little joining; flowers 5-7 mm.
long or if larger inflorescence pendulous or racemose.
I
i
688 Field Museum of Natural History— Botany, Vol. XIII j
Petals about 9 mm. long; inflorescence elongate, pendulous, simple.
0. phaeophylla, 0. pendula.
Petals 5-7 mm. long; inflorescence rather erect or suberect, some-
times more or less paniculate.
Inflorescence racemose or racemiform.
Leaves chartaceous-coriaceous, mostly or all several cm. wide
or if narrower very opaque.
Leaves about twice as long as wide, obtuse or acute.
0. tarapotensis.
Leaves, at least mostly, about three times longer than wide,
acutely long-acuminate 0. iquitosensis.
Leaves membranous, lustrous, about 2.5 cm. wide.
0. Williamsii.
Inflorescence a pyramidal panicle 0. Weberhaueri.
Veins many, but more or less reticulate; flowers about 1 cm. long or
longer or the inflorescence more or less pyramidal-paniculate
not racemose or pendulous.
Leaves broadest medially, about 3.5 dm. long, 9-12 cm. wide.
0. amplifolia.
Leaves oblong, about 2 dm. long, 6-8 cm. wide.
Leaf-veins elevated 0. aromatica.
Leaf -veins obscure 0. oleosa.
Ouratea amplifolia Sleumer, Notizbl. Bot. Gart. Berlin 13:
349. 1936.
Glabrous slender tree, the trunk only 5 cm. thick, 4 meters tall,
the very stout branchlets reddish-brown (herb.) ; petioles to 12 mm.
long, canaliculate above; leaves broadest medially, subrotund
basally, shortly and acutely acuminate apically, about 3.5 dm. long,
9-12 cm. wide, rigid-coriaceous, somewhat lustrous above, paler
beneath, coarsely serrate (with about 4 per centimeter, 1-1.5 mm.
high) or subsinuate-dentate toward the tips, midnerve prominent
both sides, the 25-30 primary lateral ones moderately raised, the
numerous subhorizontal secondary ones faint; panicles erect, about
12 cm. long, dense; rachis and pedicels rugulose, the latter thick,
8-10 mm. long; sepals oblong, 8-9 mm. long, 5-6 mm. wide, green,
the inner 3 lutescent margined, about equaling the yellow petals
and stamens; filiform style 7 mm. long. — Distinguished by author
from 0. gigantophyUa (Erh.) Engler by the much more numerous
raised instead of impressed veins and shorter pedicels.
Flora of Peru 689
Loreto: Mouth of the Santiago, flood-free woods, {Tessmann
m9, type).
Ouratea aromatica Macbr., sp. nov.
Fniticosa glaberrima; petiolis circa 5 mm. longis, 2-3 mm.
crassis foliis flexile coriaceis, supra nitidulis oblongo-vel interdum
ovato-ellipticis, basi rotundatis, ad apicem late vel paullo angustatis,
subobtuse acutis, plerumque 12-15 cm. longis, 5-6.5 cm. latis vel
circa 9 cm. longis, 4-4.5 cm. latis, minute denseque serrulatis basin
versus integris, utrinque nervis lateralibus et venis numerosis laxe
reticulars conspicue elevatis: paniculae laxiflorae circa 2 dm. longae
vel interdum solum 3-5 cm. longae, ramis inferioribus 1.5-8 cm. longis;
pedicellis laevibus 10-13 mm. longis, sepalis anguste ovato-lanceo-
latis margine conspicue scariosis, 1 cm. longis; petalis obovatis apice
rotundatis circa 12 mm. longis, staminibus circa 6 mm. longis. —
Plant 5 meters tall, stem 2 dm. in circumference, flowers ochre
yellow, aromatic (Schunke); suggests 0. odora Poeppig ex Engler
in Mart. Fl. Bras. 12, pt. 2: 334. 1876, with acutely acuminate
leaves, longer petioles, and 0. Spruceana Engler in the inflorescence
but the nerves and veins not impressed, rather elevated.
Loreto: Gamitanacocha, Rio Mazdn, Jos^ Schunke S3, type.
Ouratea iquitosensis Macbr. Candollea 5: 382. 1934.
Quite glabrous shrub, the stout branchlets somewhat flexuose,
the petioles deeply sulcate, 3.5-5 mm. long; leaves oblong-lanceolate
or sometimes elliptic-oblong, rounded to the shortly acute base,
gradually and acutely acuminate, mostly about 12 cm. long, 3-4
cm. wide, or to 15 cm. long, 5.5 cm, wide, lightly serrulate nearly to
base, firm, opaque, the strongly arcuate lateral nerves finally ascend-
ing, the horizontal veins numerous, both rather prominent beneath;
panicles nearly simple or racemiform, dense, 3-4 cm. long; pedicels
slightly angled, smooth, slender, 7-9 mm. long, subequaling the
conic-ovate buds; sepals oblong-lanceolate, acute; petals oblong-
obovate, obscurely clawed, nearly 7 mm. long, scarcely 3 mm. wide;
gjniophore and ovary subequal, the filiform style much longer. — By
typographical error petals described as nearly 10 mm. long. With
some of the characters of 0. olivaeformis (St. Hil.) Engler but
differing essentially in its dull leaves with distinctive venation;
maybe comparable too to 0. garcinioides Ule, Notizbl. Bot. Gart.
Berlin 6: 338. 1915, with oblong barely and shortly acuminate leaves;
as some other species it always grows, apparently, on inundated
banks.
690 Field Museum of Natural History— Botany, Vol. XIII
Loreto: Iquitos, Williams 1^98, type; also 1^91; 1^92; 1j^96;
im; 1U99; 1503.
Ouratea oleosa Macbr., sp. nov.
Arbor, ramulis subteretibus rubro-brunneis, nitidulis; petiolis
8-10 mm. longis, 3.5-5 mm. crassis; foliis fere oblongo-ellipticis,
basi subrotundatis, apice subabrupte breviterque acuminatis, circa
2 dm. longis, 6-8 cm. latis, subcoriaceis, supra nitidulis, plus minusve
remote serrulatis; nervis lateralibus tenuibus numeroissimis obscuris
vel vix prominulis; paniculae crassae, 1.5 dm. longae, dense granu-
losae, ramis circa 2 cm. longis; pedicellis 1 cm. longis, vix 1 mm.
crassis; sepalis ovato-lanceolatis, 1 cm. longis; petalis obovato-
oblongis, circa 1 cm. longis; staminibus 7 mm. longis; stylo circa
8 mm. longo. — Apparently, as 0. amplifolia, in the general group
of the Brazilian 0. gigantophylla (Ehr.) Engler; certainly has nothing
in common with 0. tarapotensis, to which Standley referred it. At
least in the herbarium the leaves have an oily sheen.
San Martin : Juanjui, Klug ^295, type.
Ouratea pendula (Poeppig) Engler in Mart. Fl. Bras. 12, pt. 2:
339. 1876. 0. Killipii Macbr. Candollea 5: 383. 1934, fide Sleumer.
Small tree with oblanceolate chartaceous leaves the serratures
small, not mucronate, the veins prominent, especially above, and
narrow racemiform panicles sometimes nearly a dm. long; petioles
4-7 mm. long; leaves entire toward the cuneate base, acuminate,
1.5-2 dm. long, 5-6 cm. wide, the 15 or so arcuate primary nerves
forming an angle of about 80° (the lesser at right angles) with the
well-defined midrib; pedicels 1-3 fascicled, 5-6 mm. long; buds
oblong, acute, 7 mm. long; sepals lanceolate, acute, broadly scarious
margined, 8-9 mm. long, equaled by the petals; carpels 5. — I
distinguished my proposed species principally on the comparative
size of the petals (obovate, nearly 5 mm. long) and sepals (to 3 mm.
long); actually they are somewhat longer. F.M. Neg. 9684.
San Martin: Tocache, Poeppig 2027, type. — Loreto: Iquitos,
Killip & Smith 27169 (type, 0. Killipii). Yurimaguas, Williams
3812; mO; Killip & Smith 27972. Surinam?
Ouratea phaeophylla Sleumer, Notizbl. Bot. Gart. Berlin 13;
351. 1936.
Type a shrub 1 meter high with slender terete branchlets and!
chartaceous densely and regularly serrulate lanceolate-oblong leaves;
Flora of Peru 691
petioles 5-6 mm. long; leaves broadly cuneate at base, more or less
shortly acuminate, the tip itself acute, subcurved, drying deep
castaneous above, pale brown beneath, 12-15 cm. long, 3-4.5 cm.
wide, serrations very acute, about 6 per centimeter and 1 mm. high,
the midnerve prominent both sides, lateral nerves obscure, primary
curved-ascending, secondary more or less horizontal; racemes simple,
3.5-6.5 cm. long, pedicels 3-4 mm. long; sepals ovate-lanceolate,
acuminate, 6-7 mm. long, 2 mm. wide below, the apically rounded
petals twice as wide, about 9 mm. long; style 5 mm. long. — Com-
parable to 0. pendula but has smaller thinner sharply serrate leaves,
thicker rachis, shorter pedicels (Slejimer). The Killip & Smith
specimen has leaves to 5.5 cm. wide, pedicels to 1 cm. long, fruits
about 6 mm. in diameter.
Junin: Cahuapanas, Rio Pichis, Killip & Smith 26718? (fruit).
— Loreto: Yurimaguas, Killip & Smith 27703? (fruit) ; 28200? (fruit).
Rio Nanay, Williams 768. — Cuzco(?) : Rain forest near Rio Apuri-
mac, 600 meters, (Weberbauer 5629, type).
Ouratea tarapotensis Macbr. Candollea 5: 384. 1934.
Small essentially glabrous shrub, the strongly sulcate petioles
mostly 3, Occasionally 4 mm. long; leaves elliptic or oblong-elliptic,
narrowed and acute to rounded or even emarginate at base, rounded,
obtuse or shortly acute at tip, commonly 10-12 cm. long, 4-5 cm.
wide, minutely but densely serrulate or sometimes mucronate-
serrate except toward base, chartaceous-coriaceous, scarcely lustrous,
the little prominent lateral nerves arcuate-ascending, the numerous
veins subhorizontal; panicles dense, racemiform, about 3 cm. long;
pedicels slender, smooth or obscurely tuberculate, 5-8 mm. long,
often distinctly longer than the ovoid glabrous buds; sepals narrowly
ovate-lanceolate, 5 mm. long, only slightly exceeded by the oblong
or narrowly oblong-obovate obtuse scarcely basally narrowed petals;
gynophore and ovary subequal, the former in fruit obovoid, 5-7
mm. long; drupes subclavate-ovoid, to nearly 1 cm. long, 7 mm.
thick, olivaceous, lustrous, more or less veiny. — Apparently peculiar
to the Tarapoto region and no comparable species discovered by me.
San Martin: Tarapoto, Williams 6720 (flowers); 57/^8, type
sheets; also 66 W; 66U: 6908.— Loreto: Yurimaguas, Williams ItSOl?
Ouratea Weberbaueri Sleumer, Notizbl. Bot. Gart. Berlin 13:
355. 1936.
Sparsely branched shrub or small tree to 5 meters tall with
shortly petioled oblong subcoriaceous leaves and lax panicles, about
692 Field Museum of Natural History— Botany, Vol. XIII
1 dm. long, of small (5 mm. long) flowers; leaves attenuate both
ends, subacute, drying grayish-green above, pale olive beneath,
13-21 cm. long, 5-7.5 cm. wide, regularly denticulate (teeth obtuse,
about 5 per centimeter, scarcely 1 mm. high), midnerve plane above,
prominent beneath, primary lateral nerves about 10, lightly but
distinctly impressed, prominent beneath, arcuate, not at all joining,
secondary lateral nerves or veins slightly elevated; lowest panicle
branches 3-4 cm. long, progressively shorter, the uppermost hardly
1 cm. long; rachis slender, the pedicels 3-5 mm. long, elongating
after anthesis; sepals lanceolate, acute, 5-6 mm. long, 1.8 mm. wide,
the similar petals deep yellow; style 3 mm. long. — Habit of 0.
olivaeformis (St. Hil.) Engler, the flowers much smaller (Sleumer);
in the inflorescence and small flowers suggests 0. aquatica (HBK.)
Engler.
San Martin: Zepelacio, Klug SJ^SJt; 3517; 3720.— Loreto: South
of Moyobamba, 1,200 meters, Weberhauer ^683, type.
Ouratea Williamsii Macbr. Candollea 5: 384. 1934.
Graceful slenderly branched shrub; petioles 2-3 mm. long,
canaliculate above; leaves oblong-lanceolate, cuneately narrowed
both ends, acute, mostly about 7 cm. long, about 2.5 cm. wide, or
sometimes 11 cm. long, subentire or slightly serrulate especially
toward the base, nearly translucent membranous, lustrous, the
approximate very fine nerves and veins obscure on both surfaces;
panicles terminal, racemiform, 3.5-7 cm. long; pedicels subalate
with a solitary fugacious bract, slender, about 5 mm. long, equaling
the ovoid buds, or in fruit to 12 mm. long; sepals narrowly ovate,
acutish; petals scarcely acute, probably little longer than the calyx;
gynophore oblong-ellipsoid; drupes unknown.— Resembles 0. lucens
(HBK.) Engler but differs particularly in the much longer slenderer
pedicels; the thinner leaves, essentially racemose flowers seem to
distinguish it from 0. aquatica (HBK.) Engler.
San Martin: Tarapoto, Williams 6707, type; also 6090.^
Hudnuco: Above Cayumba, Mexia 8306 A (det. Standley, 0^
iquitosensis) .
2. SAUVAGESIA L.
Herbs or more or less suffrutescent below with mostly small
often callus-margined leaves and pectinate-ciliate stipules. Flowers
pink or white, axillary or terminal and bracted by reduced leaves.
Sepals and petals 5, the former persisting, appressed to the septicidal
Flora of Peru 693
3-valved capsules, the latter convolute, deciduous. Fertile stamens,
alternating with staminodia, the latter usually in 2 series, the outer
filiform, the inner opposite the petals, scale-like, or petaloid. Anthers
linear, laterally dehiscent. Ovary 3-celled at base, 1-celled above,
the cells with many ovules in 2 rows, the numerous seeds reticulate.
Leaves and stipules imbricate; pedicels about 5 mm. long.
S. fruticosa.
Leaves and stipules remote; pedicels soon about 1 cm. long.
S. erecta.
Sauvagesia erecta L. Sp. PI. 203. 1753.
Slender semiprostrate to erect reddish brown stems more or less
ligneous toward the base, often 1-several dm. high; stipules lanceo-
late-linear, long-ciliate, to 1 cm. long; leaves oblong-lanceolate,
attenuate to very short petiole, acute, to 2 cm. long or somewhat
longer, usually 5 (-8) mm, wide, membranous, callous-serrate, the
primary nerves more prominent above; flowers solitary or few in
axillary cymes; pedicels about 1 cm. long; sepals oblong, acute,
mucronate, to 6 mm. long, the somewhat longer obovate petals
white or pink and darker at base; outer corona uninterrupted, the
inner completely elliptical staminodia; capsules ovoid, acute, scarcely
exserted. — S. rubiginosa St. Hil., treated as a variety by Eichler,
may extend from Amazonian Brazil; the upper leaves are somewhat
reduced resulting in racemiform inflorescences sometimes several
cm. long and suggests S. Sprengelii as interpreted by Dwyer in herb.
Usually along trails and in clearings; only a few of many collections
seen cited.
Ruiz and Pav6n found an infusion being used for "sickness of
the chest."
San Martin: Zepelacio, Kliig SUOU- San Roque, Williams 711 W,
7650. Lamas, Williams 6363. Near Moyobamba, Woytkowski
SSSJtS. — Hudnuco: Chacahuasi, Ruiz & Pavdn. Cuchero, Chinchao
and Pillao, Ruiz & Pavdn. Grassy clay banks, Pampayacu, SOj^O. —
Junin: Chanchamayo Valley, Schunke U91; 1^79. — Cuzco: Shrub
wood and grass steppe formation, 1,700 meters, Weherhau£T 7828.
— Loreto: Below Yurimaguas, AUard 22J^7 (det. Lyman Smith).
Mishuyacu, Klug U55. Pongo de Manseriche, Dennis 291 U2.
Covering wet beaches, Rio Santiago, Mezia 6159. Rio Nanay,
WiUiams 860. Widely distributed in the Tropics. "Yerba de San
Martin" (Ruiz & Pav6n).
694 Field Museum of Natural History — Botany, Vol. XIII
Sauvagesia fruticosa Mart. Nov. Gen. & Sp. 1: 38. 1824. S.
rosacea Ule, Notizbl. Bot. Gart. Berlin 6: 344. 1915.
More or less suffrutescent, subverticillately much branched, the
older branches often covered with the persisting lanceolate pectinate
3-6 mm. long stipules; leaves subsessile, somewhat spreading,
crowded, oblong- or linear-lanceolate, repandly glandular-serrate,
narrowed to base, acute, 1-2 cm. long, (1) 3-4 mm. wide, lustrous
green above, paler beneath; flowers 2-4, pedicels 3-5 mm. long;
sepals subequal, oblong-lanceolate, acute, 3-4 mm. long; petals
roseate, oblong, obtuse, 4 mm. long or somewhat longer; outer
staminodia lacking or filiform, in pairs or solitary, capitately en-
larged apically and alternating with the attenuate purple inner
series; capsules 5 mm. long, exserted; seeds brown, ellipsoid, scrobi-
culate, 1 mm. long. — The flower clusters are mostly toward the
branchlet tips and thus often form a leafy raceme. Description
drawn to include Ule's characterization which by the presence of
some outer staminodia caused him to ally his species to S. deflexifolia
Gardner of Brazil, and Dwyer to S. Sprengelii St. Hil., M^m. Mus.
Hist. Nat. Paris 11: 98. 1824, of Brazil with the staminodia in a con-
tinuous series as in S. erecta; how much dependability can be placed
on this apparently variable character is a subject for investigation.
The related but distinct S. ramosissima Spruce of the upper Amazon
has 2 mm. long flowers in apical pseudo-umbels, the staminodia
fasciculate and alternating. F.M. Neg. 9664.
San Martin: Tarapoto, Spruce ^■^AA (det. Eichler); Williams
5956. Cerro de Escalero, Ule 6352 (type, S. rosacea). Amazonian
Brazil.
3. CESPEDESIA Goudot
Reference: Dwyer, Lloydia 9: 54-61. 1946.
Fine glabrous trees with crowded ample coriaceous crenate
leaves and large terminal ebracteate panicles of yellow flowers.
Stipules scale-like on petioles often filamentose within. Sepals and
petals 5, the former equal, small, deciduous, the latter convolute,
much larger. Stamens 10-many, inserted at base of truncate torus
in about 3 series; anthers curved, linear, with 2 apical pores or clefts.
Ovary 4-5-celled, sometimes imperfectly, the stigma sessile, simple.
Capsules septicidally 5-valved with numerous minute seeds.—
Named for M. C^spedes, cleric and professor of botany at Bogota;
there seems to be no reason for changing the original spelling to
Cespedezia.
I
Flora of Peru 695
Cespedesia spathulata (R. & P.) Planch., Hook. Lond. Journ.
Bot. 5: 647. 1846. Godoya spathulata R. & P. Prodr. 58. 1794.
C. Sprucei van Tiegh. Ann. Sci. Nat. s^r. 8. 19: 51. 1904. C. amazon-
ica Huber, Bol. Mus. Goeldi 7: 302. 1913.
Stipules conspicuous, to 6 cm. long, 1.5 cm. wide at base, obtuse,
often splitting marginally or cleft at apex; petioles stout, flat above,
to 8 mm. wide; leaves obovate-spatulate, 4-10 dm. long, 12-18 cm.
wide, tapering toward truncate-obtuse apex, sinuate margined, the
lateral nerves ridge-like above, at the middle 7-15 (20) mm. distant;
flowers mostly in threes, pedicels and fleshy petals at anthesis 13-20
mm. long; sepals to 3 mm. long; capsules often falcate, to about
7 cm. long, 4 mm. wide, the oblong seeds 1 mm. long, the wings
8-15 mm. long. — After Dwyer, as also synonymy. C. repanda
(HBK.) van Tiegh. of Colombia, doubtfully in Peru, seems to be
a variant, the nerves 1.5-2.5 cm. distant, pedicels 7-12 mm. long,
petals 1-1.5 cm. long. A 6-25 meter tree with golden yellow flowers.
F.M. Negs. 96J^3; 21 H6 (both C. Sprucei).
San Martin : Zepelacio, Klug 3^8. San Roque, Williams 7675;
Lamas, 6^60. Monte Campana, Spruce 1^831 (type, C. Sprucei). —
Hudnuco: Cuchero, Ruiz & Pavdn, type; Poeppig 1287. — Junin:
La Merced, Killip & Smith 23888.— Loreto: Balsapuerto, Klug 2851.
Mishuyacu, Klug 12j^. Rio Santiago, Tessmann ^1 78. To Colombia
and British Guiana.
4. GODOYA R. & P.
Reference: Dwyer, Lloydia 9: 47-50. 1946.
Tall trees with lenticellate annulate branches, golden yellow
flowers in axillary and terminal racemes or panicles and simple
coriaceous sessile or subsessile crenate leaves. Sepals 5, erect,
unequal, fimbrillate-ciliate at base within, deciduous. Petals 5,
yellow, obovate, convolute. Disk inconspicuous. Stamens 10 in
a single whorl (staminodia none), declinate, the subsessile linear
anthers apically 2-pored. Ovary excentric, shortly stiped, linear-
oblong, rostrate, 5-celled as the septicidal 5-valved capsule, the
compressed seeds numerous. — Commemorates D. Manuel de Godoy,
professor of botany at Madrid. Both G. oblonga and obovata are
known by the name of "laupe" and their wood is valued for dura-
bility and resistance (Ruiz & Pav6n).
Godoya obovata R. & P. Syst. Veg. 101. 1798. G. oblonga
R. & P. Prodr. 58. pi. 11. 1794.
696 Field Museum of Natural History— Botany, Vol. XIII
Branches with scattered lenticels and conspicuous leaf-scars,
these sometimes 4.5 mm. wide; leaves obovate, cuneate at base,
triangular or obtuse at apex, 6.5-13 cm. long, 3-6.5 cm. wide,
minutely serrate, the secondary veins 3-10 mm. distant, drying
lustrous and brown or darker above; buds glandular-glossy, to 2.5
cm. long, 11 mm. wide, the racemiform inflorescences to 2 dm. long,
with whorled imbricate striate oblongish obtuse or retuse bracts
to 2 cm. long, the subannular scars to 3 cm. distant; pedicels ligneous,
0.5-2 cm. long, articulation 4-8 mm. long; sepals to 1.5 cm. long,
3-5 mm. wide, entire, obtuse, retuse or even deeply cleft; petals
unequal, obovate-elliptic, 2-3 cm. long, to 2 cm. wide, retuse;
anthers smooth, apparently inflated; ovary linear-fusiform. — After
Dwyer. To 25 meters high (Sawada). Illustrated, R. & P. Fl.
Peruv. 4: pi. 378; Dwyer, I.e. 49 (flowers).
Huanuco: Macora and Cuchero, Ruiz & Pavon, type; Poeppig.
Chinchao, Sawada 92. Bolivia. "Laupe."
5. KRUKOVIELLA A. C. Smith
Reference: John D. Dwyer, Torreya 45: 69-72. 1945.
Planchonella van Tiegh. Ann. Sci. Nat. s^r. 8. 19: 39. 1904, not
Pierre, Not. Bot. Sapot. 34. 1890.
Shrub, possibly scandent, lenticels evanescent, stipules glandular-
appendaged on inner margin, petioles short, the serrulate margins
of the leaf-blades minutely denticulate, the secondary veins con-
spicuous, subplanate. Inflorescence paniculate, the elongate rachis
much branched; pedicels shortly articulate. Sepals and petals 5,
the former without glandular appendages, the latter often bilobed.
Stamens 10 in one series, dehiscing by a single terminal pore. Pistils
solitary, 5-carpellate, the very short style with 5 radial sessile
stigmas; ovules imbricate in several rows on 5 intrusive T-shaped
placentae. F^its capsular, fusiform-falcate, the seeds short winged.
— After Dwyer. Commemorates the enthusiastic botanist B. A.
Krukoff; as pointed out by Dwyer the genus must rest on less
character than originally thought but is a convenient and meritori-
ously named segregate.
Krukoviella disticha (van Tiegh.) Dwyer, Torreya 45: 71. 1945.
Planchonella disticha van Tiegh. Ann. Sci. Nat. s^r. 8. 9: 39. 1904.
K. scandens A. C. Smith, Journ. Arnold Arb. 20: 296. 1939. Godoya
disticha (van Tiegh.) Ule, Notizbl. Bot. Gart. Berlin 6: 340. 1915.
Flora of Peru 697
Glabrous, branched; stipules thin-coriaceous, 8 mm. long;
petioles stout, 5-8 mm. long; leaves obovate, obsoletely and re-
motely serrulate, little revolute, 8-13 cm. long, 4-6.5 cm. wide, the
obscure serrations white-tipped, the 10-18 lateral nerves prominent
beneath; panicles terminal, to 2 dm. long, and often with 1 or
several flowers on lateral branchlets, their f>edicels to 1 cm. long;
bracts deciduous; sepals subequal, elliptic, concave, about 4-6 mm.
long; petals about 12 mm. long, 8 mm. wide; stamens 10, smooth,
the stout filaments 1 mm. long, anthers 5.5 mm. long; capsules
subsessile, attenuate both ends, 3-4 cm. long, 4 mm. thick, the
ellipsoid seeds 0.75 mm. long. — Shrub 1-4 meters tall, the slender
branches with dark gray-brown bark. The petioled leaves with
different serration and nervature and the smaller flowers distinguish
the species from G. obovcUa (Ule). Type of K. scandens scandent
with adventive roots in trees; flowers deep yellow; from Tabatinga
at the boundary. Illustrated, Torreya 45: 70 (flowers, fruits).
F.M. Negs. 9645; 23884.
San Martin: Tarapoto, Spruce UOOS, t5rpe; Williams 5988. Cerro
de Isco, 1,000 meters, Ule 6716. Amazonian Brazil.
CARYOCARACEAE. Souari Family
Reference: L. Williams, Trop. Woods 42: 1-18. 1935.
Erect, ligneous, with opposite or alternate digitately 3-5-foliolate
leaves and hermaphrodite flowers borne in terminal ebracteate
racemes. Calyx as petals 5-6, imbricate, the latter free or cohering
above; stamens rather ferrugineous in 5-6 series or basally connate,
the early geniculate filaments sometimes sterile, the outer always
with 2-celled anthers. Ovary 4-20-celled, ovules solitary. Fruit
with a ligneous muricate or spinescent endocarp finally dehiscing
into 1-seeded cocci; seeds often reniform, endosperm thin, embryo
with spirally twisted radicle, the small cotyledons hamately in-
flexed.
Souari fruits, particularly those of Caryocar nuciferum L. of
South America, are of some economic importance as the mesocarp
is rich in oil and the nuts are edible; the woods are hard and durable,
useful for purposes requiring strength and resistance to dampness
(Williams).
Leaves opposite; caljrx deeply parted 1. Caryocar.
Leaves alternate; calyx truncately dentate 2. ArUhodiscus.
698 Field Museum of Natural History — Botany, Vol. XIII
1. CARYOCAR L.
References: Lanjouw & Van Heerdt, in Pulle, Fl. Surinam 3:
366-372. 1941; L. Williams, Trop. Woods 42: 1-18. 1935.
Trees with opposite coriaceous leaves, the leaflets often dentate,
the stipules, if developed, promptly caducous. Petals free or basally
subconnate with the stamens, the inner of these without anthers.
Ovary 4-6-celled, the 4-6 filiform styles greatly elongate. Seeds
large, subreniform, radicle fleshy, arcuate.
The edible nuts of C. nuciferum L. of the Guianas provide
a cuHnary oil; for the morphology of this species with many figures
see Fritz Blank, Ber. Schweiz. Bot. Ges. 49: 438-494. 1939. Wil-
Hams' account, besides technical wood descriptions and economic
data, contains keys without species descriptions, but with useful
notes concerning habitats and dimensions. Possibly cultivated
(or native?) — a nut was found by Kuhlman on the Rio Branco,
Brazil (Ducke) — it may be known by its large (to 8 cm. long) flowers
and fruits (to 12 cm. long) covered with smooth rounded projections.
The differences observed between some of the Peruvian collec-
tions and assumed to indicate specific entities may doubtfully be
found to carry that import when the trees are well-known.
Stipules persisting; stamens about 150; fruit spines 2-4 mm. long.
C. microcarpum.
Stipules caducous or none; stamens about 300; fruit spines (known)
5-12 mm. long. ^
Leaflets entire or merely undulate.
Leaflets glabrous C glabrum.
Leaflets pubescent in nerve axils beneath C. coccineum.
Leaflets obviously, sometimes remotely, crenulate . C. Tessmannii.
Caryocar coccineum Pilger, Notizbl. Bot. Gart. Berlin 10:
127. 1927. j
Nearly glabrous tree, similar to C glabrum but the coriaceous
petiolulate leaflets with fasciculate trichomes in the axils of the
nerves beneath, these prominent, about 10, peduncles about 1 cm.
long, calyx (with pediform base) 1 cm. long, and petals, at least
in type, scarlet, to 2.5 cm. long, the dark carmine stamens about
twice as long. — Type a tree of flood-free terrain, 20 meters tall,
trunk 5 dm. in diameter; Williams -4-4^5, often 30 meters, small
buttresses, flowers garnet to yellow, fruit 2.5 cm. thick, kernel edible.
Flora of Peru 699
according to Williams, who describes wood, Field Mus. Bot. 15:
332. 1936, heart wood pale reddish or russet brown. F.M. Neg.
9696.
Loreto: Mouth of the Santiago, Tessmann 1,092, type. Fortaleza
on the Huallaga, lowland, Williams jU25. "Almendro" (Williams).
Caryocar glabrum (Aubl.) Pers. Syn. PI. 2: 84. 1806. Saouari
glabra Aubl. PI. Guian. 1: 599. pi. 2^0. 1775.
Branches scarred by the promptly deciduous lanceolate stipules,
these to 3 mm. long; petioles 3-9 cm. long; stipules caducous, minute,
the petiolules 2-7 mm. long; leaflets 3, oblong- sometimes ovate-
elliptic, obtuse and rounded or subcordate at base, obtusely short-
acuminate, 8-15 cm. long, 3.5-6 cm. wide, entire (subundulate at
margins), glabrous, chartaceous or subcoriaceous; peduncles 2-6.5
cm. long, raceme axis 1-2 cm. long with usually 15-25 flowers;
pedicels rather stout, 1.5-3 cm. long; calyx lobes suborbicular,
fimbriate-pilose, about 4 mm. wide, 2-3 mm. long; petals 2 cm. long
or somewhat longer, at least half as wide, very thick, fimbriate-denti-
culate; stamens about 350, the shortly united outer series 3.5-4 cm.
long, the inner on annulus 5 mm. high only 1.5-2 cm. long; anthers
oblong; ovary cells and styles 4, the latter 4-5 cm. long; fruits
ellipsoid-globose, 5-6 cm. long, 5-8 cm. broad, 4 cm. thick (Ducke),
exocarp as mesocarp thick, fleshy, outer soft reddish layer, endocarp
soon deciduous and scarcely covering the ligneous sharp spines of
the inner layer, these 5-12 mm. long, the subreniform (usually 1 or 2)
seeds about 5 cm. long, 3-4 cm. thick, rich in oil and, according to
Ducke, savory. — After Lanjouw & Van Heerdt (see for synonymy),
who have followed the interpretation of Ducke, Archiv. Jard. Bot.
Rio Jan. 4: 133. 1925, and Pilger, Pflanzenfam. ed. 2. 21: 90-93.
1925, in the use of Aublet's names, which is the business-like solution
to the nomenclatorial problem. Usually a 20 or 35 rarely 50 meter
tree with brown heartwood of flood-free woods (Ducke); Tessmann's
tree was 15 meters high, trunk 58 cm. in diameter, petals pale yellow,
filaments roseate; Ducke in a letter to Pilger, kindly shared with me,
suggested that Rio Itaya specimens were probably C microcarpum,
so the Tessmann material may have been an earlier determination.
C. Krukovii Gilly, Trop. Woods 72: 17. 1942, from Sao Paulo and
perhaps extending into Peru, has minute calyx lobes (1-1.5 mm.
long), stamens and roseate petals 10 mm. long. C. parviflorum A. C.
Smith, Joum. Arnold Arb. 20: 298. 1939, of Amazon region, has
flowers only 18-25 mm. long, with filaments 3 cm. instead of about
700 Field Museum of Natural History— Botany, Vol. XIII
6 as for C. glabrum (Smith). Illustrated, Ducke, Archiv. Jard. Bot.
Rio Jan. 5: pi 23. fig. 55 (flower).
Loreto: Soledad, Tessmann 5299 (det. Pilger). To the West
Indies. "Almendra" (Tessmann), "piquia-rana" (Brazil).
Caryocar microcarpum Ducke, Archiv. Jard. Bot. Rio Jan.
4: 133. 1925.
Resembles C. glabrum but stipels persisting, stamens hardly
half as many, fruit 3-4 cm. broad, the spines of unequal length, often
fasciculate and only 2-4 mm. long; moreover the petals are greenish-
white (as stamens) or roseate or yellow, fruit 2-3 cm. long and thick,
with rather thick and fleshy exocarp, mesocarp and outer endocarp,
and the 2-4 seeds 2-4 cm. long, 10-18 mm. in diameter. — A smaller
tree than C. glabrum, wood nearly uniform, and found on inundated
terrain; the leaves and bark rubbed in water produce a foam used
for soap; the fruits are not edible (Ducke). Williams, Field Mus.
Bot. 15: 333. 1936, describes the wood (heart-wood grayish brown)
from his specimen under C. glabrum to which his general remarks
apply in part. Illustrated, Ducke, Archiv. Jard. Bot. Rio Jan. 5:
Vl. 23, fig. 56 (flower). F.M. Neg. 9699.
Loreto: Timbuchi, Rio Nanay, Williams 1000. Mishuyacu near
Iquitos, Klug 689. Rio Mazan, Jose Schunke 167 (det. Standley,
C. Tessmannii) . To Guianas. "Almendro de bajo" (Williams).
Caryocar Tessmannii Pilger, Notizbl. Bot. Gart. Berlin 10:
126. 1927. C. amydaliforme (sic) R. & P. ex G. Don, Gen. Syst. 1:
654. 1831. C. amygdaliforme (sic) R. & P. ex Walpers, Repert.
Bot. 1:410.1842.
Glabrous, the flowers with ochre yellow petals and carmine
filaments, the former 2.5- nearly 3 cm. long, the latter 5 cm. long;
petioles about 1 dm. long, petiolules 7-9, or terminal to 13 mm. long;
leaflets subequal, thin-coriaceous, nearly elliptic or obovate-elliptic
and cuneately narrowed to base, caudate-acuminate, often 15-17
cm. long, 7 cm. wide, irregularly and remotely crenate or undulate-
crenulate, the 11-13 lateral nerves prominent only beneath; stipels
none; racemes 4-5 cm. long; peduncles 1-flowered, stout, 13-25
mm. long; pedicels with decurrent calyx 1 cm. long, the rounded
calyx lobes ciliolate; fruit according to collector spiny, the inner
part edible. — C. amygdaliferum Mutis of Colombia has distinctly
serrate leaflets with stipels; it is probably the tree referred to the
species of Mutis by Ruiz & Pavon and published, essentially as
Flora of Peru 701
a nomen nudum, under the names cited above, which, as Pilger
suggests, are to be cancelled. Williams specimen doubtful; heart-
wood pale yellow or light brown; tree 30 meters, trunk 2 dm. in
'diameter, flowers red, drupe round, brown speckled, pulp rich in oil
used for cooking and medicinally, kernel also edible according to
Williams, who gives anatomy of wood. Field Mus. Bot. 15: 333. 1936.
C. Tessmannii, somewhat similar to C pallidum A. C. Smith of the
upper Amazon, has scarlet filaments, less distinctly serrate leaves,
axillary hair tufts, no stipels (Smith, Joum. Arnold. Arb. 20: 297.
1939). F.M. Neg. 9700.
San Martin: Sandy loam along forest trail, Tarapoto, Williams
6106? Juanjui, Klu^ 3907 (det. Standley). — Hudnuco: Chicoplaya,
Ruiz & Pavdn (det. Pilger). — Loreto: Pongo de Manseriche, Tess-
Tnann 432^, type. "Almendra," "almendr6n" (Williams).
2. ANTHODISCUS G. F. W. Meyer
Shrubs or trees with alternate leaves, the 3 leaflets entire or
crenate and the flowers with short cupulate calyx, the petals co-
hering above and calyptrately deciduous. Stamens at least in bud
annulately connate basally, all fertile. Ovary cells and styles 8-20,
the latter shortly linear-filiform. Seeds small; radicle greatly
elongating, spirally contorted.
Leaflets obtusely acuminate A. trifoliatus.
Leaflets rounded apically.
Pedicels soon to 1 cm. long; leaves nearly glabrous. A. peruanus.
Pedicels 5 mm. long; leaves opaque above, pilosulous beneath on
nerves A. glaucescens.
Pedicels about 1 cm. long; leaves lustrous above, pilose beneath.
A. pilosus.
Anthodiscus glaucescens Macbr. Candollea 5: 385. 1934.
Branchlets puberulent, petioles (2-2.5 cm. long), leaf-nerves
beneath and racemes sparsely pilosulous, otherwise glabrous;
leaflets similar but the lateral somewhat smaller, the central to 1 dm.
long, 4 cm. wide, all oblong-obovate, rounded-apiculate apically,
nearly sessile or cuneate-attenuate to short petiolule, crenate-
serrulate, subcoriaceous, opaque and glaucous above, paler beneath,
the nerves and veins rather prominent on the surfaces; peduncles
3 cm. long; racemes densely flowered, 5 cm. long, pedicels 5 mm.
702 Field Museum of Natural History— Botany, Vol. XIII
long; calyx subcoriaceous, cupulate, obsoletely or lightly crenate;
flowers (immature) 7 cm. long, the thick petals glabrous; filaments
before anthesis more or less incurved, strongly unequal, the longer
about 5 mm. long, smooth; ovary ovoid; styles 10, erect subulate
attenuate apically, about 2 mm. long. — Perhaps should be included
in A. peruanus, but at the time described I wrote: no described
species in so far as I have been able to determine is similar. The
type about 2 meters high at edge of forest.
San Martin: San Roque, Williams 7491, type.
Anthodiscus peruanus Baillon, Adansonia 10: 241. 1872.
Rather robust tree, quite glabrous, the leaflets of the ample
leaves unequal, the lateral much smaller, the middle one to about
1 dm. long, 4 cm. wide, all long-obovate or subspathulate, notably
narrowed to base, obtuse, irregularly crenate and finally unevenly
reticulate veined; racemes to 1.5 dm. long, the pedicels basally and
conspicuously articulate; calyx subequally 5-crenate, coriaceous as
the strongly imbricate petals; stamens soon separated into 5 pha-
langes, the solitary central one much the longer, all the anthers
finally erect; ovary bordered at base with a somewhat unequal disk,
cells as many as 12, the radiating style branches rather fleshy,
apically glandular-stigmatose and scarcely sulcate within. — Species
name as above by author; in Pflanzenfam. ed. 2. 21: 93. 1925 given
as peruvianus, perhaps unintentionally. Tall tree of non-inundated
forests, the Tessmann example 18 meters high, trunk 6 dm. in
diameter, flowers bright golden yellow, fragrant. A. amazonicus
Gleason & Smith, Brittonia 2: 156. 1936, has smaller subemarginate
leaflets, pedicels not inserted in pits. F.M. Negs. 29274; 35283.
Loreto: Pongo de Manseriche, Tessmann 1^856 (det. Pilger).
Without data, Ruiz & Pavdn, type; River o. Amazonian Brazil.
Anthodiscus pilosus Ducke, Trop. Woods 90: 23. 1947.
Branchlets lenticellate, the younger as the petioles, these 4-5
cm. long, leaves beneath and racemes including the calyces densely
canescent pilose; terminal petiolule 1-2 cm. long, lateral 3-10 mm.
long, the terminal leaflet 7-10 cm., usually 4-6 cm. wide, the lateral
rarely much smaller, all obovate, cuneately decurrent, rounded and
minutely apiculate (sometimes subretuse) at apex, revolute and
distinctly crenate, chartaceous, more lustrous above than beneath,
the 9-12 lateral nerves and fine reticulate veins rather conspicuous;
racemes 11-14 cm. long, the upper half floriferous; calyx 4-5 mm.
Flora of Peru 703
broad, 2-2.5 mm. high, subobsoletely 5-dentate; petals intense
yellow, to 1 cm. long, oblong, nearly glabrous; stamens to 8 mm.
long, filaments slightly tuberculate; ovary glabrous. — Unique in
the abundant indument; discovered in a high dry forest among
several species of the "caatinga" of the upper Rio Negro, for example
Leptothyrsa SpriLcei (Ducke).
Loreto: In white sandy soil, San Juan Nuevo near Iquitos,
{Ducke 1855, type).
Anthodiscus trifoliatus G. F. W. Mey. Prim. Fl. Essequeb.
194. 1818.
Glabrous tree with terete branchlets, the leaflets of the rather
large leaves subequal, the flowers greenish, about a cm. across and
very fragrant; petioles 2-3 cm. long, petiolules rarely 5 mm. long;
leaflets oblong-lanceolate, acute, shortly attenuate to base, 6-12
cm. long, 3-5.5 cm. wide, crenate-serrate, lustrous above; racemes
terminal, simple, many-flowered, the pedicels only 2 or 3 mm. long
and with a pair of ovate acute thin-margined bracts below the
middle; calyx obsoletely 5-lobed, to 4 mm. broad; petals green,
oblong, acuminate; filaments (apparantly, ex icone) coarsely tuber-
culate; ovary depressed medially, the 14-20 styles compressed-
subulate. — Fruit simulating that of Malva rotundifolia (author).
Standley proposed an herbarium name after the collector for the
Peruvian tree, which when better known may prove to be distinct
from the type of Meyer, which was about 12 meters tall from the
Rio Essequibo. Illustrated, Trans. Linn. Soc. 18: pi. 20.
San Martin: Pongo de Cainarachi, Klug 2682 (det. Standley
with query). British Guiana.
MARCGRAVIACEAE
Scandent or epiphytic, rarely arborescent, with alternate
coriaceous entire leaves (fertile stems) and terminal racemes, these
often short and umbelliform, the bracts (or nectaries) more or less
deformed and lobed, often sac-like. Sepals and petals 5, at least
the former strongly imbricate, sometimes calyptrate and falling
as a unit. Anthers basifixed. Ovary several-celled. Fruit usually
indehiscent with numerous seeds; endosp)erm wanting, radicle
slightly curved, much longer than the small cotyledons. — The
German "Nektarbehalter" is a good concise name for the often
bizarre foliose nectar-containing bracts.
704 Field Museum of Natural History— Botany, Vol. XIII
Inflorescence in part sterile (the central flowers); sepals 4; petals
calyptrately connate 1. Marcgravia.
Inflorescence entirely fertile; sepals 5; petals free or basally connate.
Bracts solid or nearly, usually globose; ovary 2-celled; racemes
long; stamens 5 2. Ruyschia.
Bracts sac-like, with 2-parted hollow spur; ovary 5-celled; racemes
rather long; stamens 5 3. Sourouhea.
Bracts spoon-shaped or sac-like, but not 2-parted; ovary 3-5-
celled; stamens often many; flowers often subumbellate.
4. Norantea.
1. MARCGRAVIA L.
Racemes usually much shortened and umbelliform, the sac-like
bracts in the middle of the umbels stiped, free from pedicels. Petals
calyptrately deciduous as an undivided unit. Stamens 12-40, mostly
lightly connate basally, free from corolla. Anthers linear. Stigmas
obscurely radiate. Fruits globose, somewhat fleshy. — Leaves of
creeping sterile branches often small, submembranous.
Leaves apparently veinless or nearly.
Leaves obtusish, sometimes cusped M. macrocarpa.
Leaves acuminate or distinctly acute.
Leaves eglandular or 2 glands near base beneath . . M. trinitatis.
Leaves with about 8 glands beneath M. Williamsii.
Leaves obviously veined, at least beneath.
Leaves to about 3.5 cm. wide, subsessile M. crenata.
Leaves distinctly wider unless M. Weberbaueri, shortly but ob-
viously petioled.
Leaf -veins scarcely if at all visible above M. Weberbaueri.
Leaf -veins more or less obvious both sides.
Pedicels enlarged to 3 mm. thick below flowers.
M. macrophylla.
Pedicels slender, about 2 mm. thick below flowers.
Pedicels about 2 cm. long.
Leaves often subelliptic, 1-1.5 dm. long, often nearly
half as wide M. parviflora.
Leaves oblong, 2-2.5 dm. long, about a third as wide.
M. UmgifoUa.
Pedicels about 5 cm. long M. strenua.
Flora of Peru 705
Marcgravia crenata Poeppig ex Wittm. in Mart. Fl. Bras. 12,
pt. 1: 223. 1878. M. polyarUha Delf. var. ocddentcUis Wittm. I.e.
224 as to Peru.
Leaves subsessile, oblong-ovate, rounded-obtuse at base, cuspi-
date or obtuse, 8-10 cm. long, about 3 cm. wide, membranous,
nervose, marginally glandular crenate, the adult with a few patelli-
form glands mostly toward the apex; flowers 15-23, the slender
pedicels about 3, rarely 4 cm. long, suberect, crowded; bracts shortly
petiolate, erect, 4-4.5 cm. long, elongate tubular-cucullate, slightly
clavate apically, the orifice obliquely open; corolla ovoid-conic,
obtuse, 8 mm. high, 5 mm. thick; stamens about 30.
San Martin: Vitoc, Ruiz & Pav6n. — Hudnuco: Chihuamula and
Cuchero, Poeppig 1U77, type. — Junln: Chanchamayo Valley,
Schunke 6US. — Loreto: Caballo-Cocha, WiUiams 2307. Brazil.
Marcgravia longifolia Macbr. Candollea 5: 386. 1934.
Resembles M. macrophylla but leaves 2-2.5 dm. long and bracts
strongly clavate but scarcely 6 mm. thick at apex, 2 cm. long;
peduncles to 1.5 dm. long; pedicels about 30, sparsely rufous tuber-
culate, 2 cm. long, 2 mm. thick below the flower. — This and M.
strenua may be better treated as variants of the similar species but
the three forms are recognizable on characters accepted as specific
elsewhere in the genus.
Loreto: Mishuyacu, near Iquitos, Klitg 92U, type; 398.
Marcgravia macrocarpa [R. & P.] Sleumer ex Diels, Bibl. Bot.
116: 107. 1937. Af. oblongifolia Pav6n ex Wittm. in Mart. Fl. Bras.
12, pt. 1: 223. 1878, fide Diels, not M. oblongifolia Pav6n in synony-
my.
Leaves subsessile, oblong or narrowly oblong, obliquely obtuse
at base, rounded-obtuse at tip, usually 8 or 9 cm. long, about 2.5
cm. wide, very heavy, notably with a series down the middle each
side of midnerve of large obliquely poriform glands and sometimes
a few between the margin and the series; umbels few-flowered,
subsessile, the rather stout pedicels 6 cm. long; bractlets as sepals
large (expanded calyx 12 mm. across), orbiculate, spreading; corolla
oblong-conic, lightly umbonate, the point recurved; stamens about
32; stigma mammiform. — In the Ecuadorian specimen the stigma
is subsessile as in M. flageUaris Poeppig, which is scarcely distinct
(Diels) ; leaves dark green above, brighter beneath, veins invisible.
Illustrated, Wittmark, I.e. pi. J^l, fig. 2 (leaf, fruit and flower).
706 Field Museum of Natural History— Botany, Vol. XIII
Fruits edible, known as "Purum-hijos" from their resemblance
shape and taste to figs (Ruiz & Pavon).
San Martin: Vitoc, Ruiz {& Pavdn) 1790, type, Herb. Dahlem.
— Junin : Chanchamayo, Isern 205If.. Ecuador.
Marcgravia macrophylla (Wittm.) Gilg, Bot. Jahrb. 25: Beibl.
60: 27. 1898. M. rectiflora Tr. & PI. var. macrophylla Wittm. in
Mart. Fl. Bras. 12, pt. 1: 222. 1878.
Subterete flowering branches, pedicels and bracts often strongly
tuberculate; leaves shortly petioled, oblong-elliptic, acutely cuspi-
date or sometimes subfalcately peduncled, almost 1.5-2 dm. long,
6-8 cm. wide, nervose, with 2 large glands toward the base and pore-
like ones beneath; pedicels thick, to about 5 cm. long, two or three
times longer than the straight, shortly stiped, strongly clavate
bracts, these with orbicular orifice; bractlets large, sepaloid; corolla
cylindric-conic; stamens 36. — Most hke M. polyantha Delp. (Gilg).
Illustrated, Wittmark, I.e. pi. W, fig- 2 (leaf and bract). F.M. Neg.
9725.
San Martin: Tocache, Poeppig 2005, type. — Loreto: Paraiso,
upper Rio Itaya, Williams SSI 8. Ecuador.
Marcgravia parviflora Richard ex Wittm. in Mart. Fl. Bras.
12, pt. 1: 227, pi. k2, fig. S. 1878.
Branchlets densely rugulose-lenticellate; petioles 4-8 cm. long;
leaves oblong-elliptic or subobovate, acute at base, shortly and
obtusely acuminate, mostly about 1-1.5 dm. long, 4-6 cm. wide,
coriaceous, rather lustrous especially above, glanduliferous at margin
but hypophyllous glands none, the 8-12 lateral nerves prominent
both sides, not obviously parallel; inflorescence umbelliform, some-
times 30-flowered, the peduncle long, the shortly stiped central bracts
tubular-saccate, inflated above and with a small sterile flower;
pedicels notably tuberculate, 2-2.5 cm. long, bearing the flowers
perpendicularly, densely puberulent as the broad sepals, these with
upturned edge; corolla typically conical-ovoid, rugulose, puberulent-
pilose; stamens 11 or 12, the lanceolate anthers about as long as the
flat filaments; ovary sulcate, 8-celled, the short style with broad
flat stigma; fruit about 8 mm. thick, the many seeds foveolate. —
The var. macrophylla Wittm. I.e. 228, has, for the genus, gigantic,
nearly eglandular leaves, shorter peduncle enlarged at apex, bract-
lets free from calyx, corolla subglobose; it is known from adjacent
Brazil.
Flora of Peru 707
Peru (probably; see note above). To French Guiana, Panama
and Brazil.
Marcgravia strenua Macbr. Candollea 5: 386. 1934.
A vigorous liana similar to M. macropkylla; petioles stout, 5 mm.
long; leaves oblong-elliptic, broadly cuneate at base, caudate-
acuminate, to 1.5 dm. long, 6 cm. wide, lightly and laxly reticulate-
veined both sides, subrigid-coriaceous and with 2 large glands at
base; pedicels about 5 cm. long, slender, conspicuously verruculose-
tuberculate; bracts unknown; ovary about 6 mm. high. — Dis-
tinguished readily from the similar species in spite of the incom-
pleteness of the material, by the very rough long slender pedicels.
Loreto: Yurimaguas, Williams 3809, type; also Jt536; Killip &
Smith 29032.
Marcgravia trinitatis Presl, Bot. Bemerk. 122. 1844. M.
flagellaris Poeppig ex Gilg & Werdermann, Pflanzenfam. ed. 2. 21:
105. 1925. M. rectiflora Tr. & PI. and var. flagellaris (Poeppig)
Wittm. in Mart. Fl. Bras. 12, pt. 1: 222. 1878; M. flagellaris Poeppig
in s3nnonymy.
Leaves (fertile) narrowly oblong, rounded at base, gradually
acuminate, (8) 11-12 cm. long, (2) 3-3.5 cm. wide (typically),
.nearly veinless, glandular margined, the surface glands lacking
except 2 near base; umbels few-flowered, the 10 or so pedicels about
4 cm. long, the tubular-cucullate bracts clavate apically, usually
shorter; sepals suborbicular; corolla calyptriform; cylindric-conical,
obtuse; stamens 36-40, filaments elongate; ovary globose, 8-10-
celled, the mammiform stigma obscurely lobed; fruits large, globose,
with an edible pulp at maturity. — Near Leticia on the boundary
Williams found a variation, apparently, with leaves 8-9.5 cm. long,
17-22 mm. wide (var. leticiana Macbr., comb. nov. M. flagellaris
var. leticiana Macbr. Candollea 5: 387. 1934). With the carelessness
I have pointed out as unnecessary my varietal name was made
a part of an unpublished one; and since M. trinitatis Presl, 1844, is
the earliest of the several si>ecies names listed by G. & W., it seems
impossible, granted that this plant is distinct, to accept their listing
as publication as the key character applies equally to all the species
named. Therefore, particularly since Urban & Gilg in 1897 referred
without question Poeppig 2026 to M. trinitatis, that name sens. lat.
seems the available one for our plant, which at most is probably
only a variety. F.M. Neg. 32237 (Poeppig) .
708 Field Museum of Natural History — Botany, Vol. XIII
Loreto: Near Leticia, Williams 31 ^9 (type, var. leticiana). Yuri-
maguas, Poeppig 2026 (type, M. flagellaris); Williams 4-^97. — Rio
Acre: Ule 9612 (det. Gilg). Brazil.
Marcgravia Weberbaueri Gilg, Bot. Jahrb. 42: 124. 1908.
M. crenata Poepp. var, roquensis Macbr. Candollea 5: 387. 1934.
Younger branches 2.5 mm. thick, densely lenticellate, the leaf
internodes about 3 cm.; petioles 5-7 mm. long; leaves oval or oval-
oblong, narrowly cuneate to base and the acute apex, 7-11 cm. long,
nearly 3 or 4-5 cm. wide, opaque both sides, the many parallel
lateral nerves long percurrent near the margin, inconspicuous above,
slightly so beneath, with many small glands generally scattered and
a few larger ones at the margin; rachis 6 or 7 mm. long, with about
30 flowers, the densely transversely rugose pedicels 2.2-2.4 cm. long;
5 green bracts at apex of rachis, about 7 mm. long, stiped, long-
tubular, 3-4 cm. long, at apex 7-8 mm. thick, below apex 4 mm.
thick, the rotund enlarged throat about 8 mm. across; buds oblong,
7 mm. long, 4-5 mm. thick; sepals about 3 mm. wide, only 1 mm.
long. — Elegant distinct species allied to M. umbellata L. (author).
F.M. Neg. 9735.
San Martin: Near Moyobamba, 1,300 meters, Weberbauer J^667,
type; 289. San Roque, Williams 7388 (type, M. crenata var.
roquensis).
Marcgravia Williamsii Macbr. Candollea 5: 386. 1934.
Petioles 3-4 mm. long; leaves oblong or oblong-elliptic, rounded
at base, nearly caudate-acuminate (acumen to 1.5 cm. long), mostly
about 1 dm. long, 3 cm. wide, subcoriaceous, seemingly quite vein-
less, entire but strongly undulate margined and with only about
8 punctiform glands beneath; pedicels 2-2.5 cm. long, little enlarged
below the flowers; bracts to 7.5 cm. long, narrowly tubular-cucullate,
little incurved apically, rather clavate, orifice suborbicular, margin-
ally expanded with subapical sterile flower; corolla ovoid-rotund. —
Comparable and related to M. Eichleriana Wittm. because of the
extended rachis that characterizes^ that species and M. coriacea Vahl
but remarkable in the length of the bracts. Except for these, in
general aspect suggestive of M. crenata (with longer pedicels) and
M. Weberbaueri, Rare vine, darker inner bark deep chocolate
brown, branches pendent from tall trees, according to Williams,
who gives wood anatomy. Field Mus. Bot. 15: 334. 1936.
Loreto: Forest of Paraiso, Alto Rfo Itaya, WiUiams 3209, type.
"Murcuhuasca" (Williams).
Flora of Peru 709
2. RUYSCHIA Jacq.
Much like Norantea but petals always slightly connate at base,
stamens 5, the anthers affixed dorsally, ovary 2-celled and bracts
3-lobed, adnate to pedicels, globose or rather (rarely) spoon-shaped
but always nearly or quite solid, and fruits dehiscent.
Ruyschia sphaeradenia Delpino, Atti Soc. Ital. Sci. Nat.
Milano 12: 180, 195. 1869; Nuovo Giom. Bot. Ital. 1: 257. 1869.
Slender flowering branchlets greenish-gray; petioles 5-10 mm.
long; leaves oblong or obovate, attenuate to base, subemarginate
and minutely mucronulate, 5-8 cm. long, 2-3 cm. wide, with nu-
merous seriate glands toward the subvalvate margin; racemes
elongate, many-flowered, pedicels slender, 7-10 mm. long; bracts
inserted about medially, pendulous, 1-2 mm. thick; subsessile,
globose; buds globose-cylindrical, petals oblong; ovary subglobose
after an thesis, 2-celled. — The flowers exhale a very agreeable fra-
grance (Ruiz & Pav6n). My collection was from a scrub-tree in
rocks. Illustrated, Mart. Fl. Bras. 12, pt. 1. pi. U9, jig. 2. F.M.
Neg. 9717.
Cajamarca: Cutervo, Raimondi. — Hudnuco: Chaglla, Ruiz &
Pavdn, type. Yanano, S776.
3. SOUROUBEA Aublet
Habit and foliage as in other genera but flowers racemose, bracts
3-partite, the middle lobe narrow, hollow, the lateral more or less
parallel with the pedicel. Bractlets simulating outer sepals. Petals
more or less connate basally, reflexing in anthesis and deciduous as
a unit. Stamens 5, joined to p)etals at base; anthers affixed dorsally.
Ovary 4-6-celled, the sessile peltate stigma 4-6-lobed. Fruits
globose, seeds few.
Pedicels slender, 1- scarcely 2 mm. thick medially; leaf -nerves
slender, completely obvious beneath, rarely faint . . S. guianensis.
Pedicels stout, at least 2 mm. thick even in flower medially; leaf-
nerves if obvious beneath often incompletely and coarse.
Glands in part seriate; nerves scarcely prominent.
S. suaveolens, S. didyma.
Glands all scattered; nerves prominent beneath. .S. pachyphylla.
Souroubea didyma [Poeppig] Gilg, Bot. Jahrb. 25: Beibl. 60:
33. 1898. S. crassipes (Tr. & PI.) Wittm. var. didyma [Poeppig]
Wittm. in Mart. Fl. Bras. 12, pt. 1: 254. 1878.
L
710 Field Museum of Natural History— Botany, Vol. XIII
Leaves oval-obovate or often broadly obovate, rounded at both
ends or shortly apiculate at tip, thick-coriaceous, the nerves both
sides scarcely obvious, 8-12 cm. long, 3-5 (-7) cm. wide; pedicels
very thick, the flowers large and the 3-parted bracts small, the spur
short, thick, spreading, the wing rather fleshy, lanceolate, rounded.
—Illustrated, Wittmark, I.e. pi. 51, fig. 3. F.M. Negs. 9719; 32235.
Hudnuco: Pampayacu, Kanehira 193; 198. Near Cuchero,
Poeppig, t3rpe. — Loreto: Puerto Mel^ndez, Tessmann J^71i.7 (det.
Gilg, afline).
Souroubea guianensis Aublet, PI. Guian. 1: 244. pi. 97. 1775.
Epiphytic or climbing, sometimes a shrub or small tree; petioles
4-8 mm. long; leaves more or less obovate, usually somewhat obhque
toward the cuneate or slightly rounded base, the rounded or obtuse
apex mucronate, ordinarily about a dm. long, half as wide, typically
lighter colored and rough above with a few hypophyllous glands
scattered more or less remotely from the margin, the fine lateral
nerves obscure above, obvious beneath; racemes a dm. or two long,
many-flowered, the rachis stout; bracts borne at tip of the long
(typically 1.5-2 cm., -4 cm. in fruit) pedicels, red, 3-parted, the
middle lobe a hollow narrow sac-like spur about 1 cm. long, clavate
at tip, the lateral lobes lanceolate-spathulate with upturned margins,
early recurved, later paralleling the pedicel; sepals suborbicular,
about 3 mm. long, 4 mm. wide, ciliolate; petals connate nearly
medially, obtuse, fleshy, yellow or orange, 8-9 mm. long; fruits
rugulose, 5-ridged, 5-celled; seeds arcuate, linear, 3-5 mm. long. —
Tessmann ^560 with very broad subrotund leaves was designated in
herb, as a variety by Gilg. Illustrated, Mart. Fl. Bras. 12, pt. 1:
pi 50.
Hudnuco: Cayumba, Mexia 8310 (det. Standley). — Loreto:
Yurimaguas, Poeppig 2879. Puerto Mel^ndez, Tessmann J!f798;
U560. Rio Mazdn, Jos^ Schunke 20 (det. Standley). — Puno: Tatan-
ara, Lechler 2659; Gay. To the Guianas and Central America.
"Laupo" (Mexia).
Souroubea pachyphylla Gilg, Bot. Jahrb. 25: Beibl. 60: 33.
1898.
Allied to S. didyma Gilg; petioles 7-9 mm. long; leaves broadly
obovate, narrowed to the thick petiole, rounded or shortly apiculate,
about 14 cm. long and half as wide, rigid coriaceous, opaque both
sides, the strictly parallel nerves prominent beneath, veins incon-
Flora of Peru 711
spicuous, small glands unevenly dispersed; racemes to 1.5 dm. long,
the pedicels 13-14 mm. long, 2 mm. thick, glabrous; bracts 12-13
mm. long, 3-parted, the bifurcate fleshy base 5 mm. long, 2-2.5
mm. wide, the spur 7-8 mm. long, 2.5 mm. thick at tip; sepals about
3 mm. across; petals forming bud 7-8#mm. high, 5 mm. thick. —
F.M. Neg. 9720.
San Martin: Shrub formation near Moyobamba, Weberhauer
J^6S6; 289. Tarapoto, Spruce U69, type; Ule 6S20 (det. Pilger).
San Roque, Williams 7011.
Souroubea suaveolens Gilg, Bot. Jahrb. 42: 128. 1908.
A 3 meter shrub, the brown 4-5 mm. thick branches with a
readily exfoliating whitish bark; petioles 7-8 mm. long; leaves
broadly obovate, rounded at tip, cuneate to base, 10-14 cm. long,
4-8 cm. wide, rigid coriaceous, opaque both sides, with a nearly
regular series of broad open glands about 2 cm. from the midrib
and some others scattered; lateral nerves many, about 2 mm. from
the margin, joined, only beneath prominent, the veins not at all;
racemes terminal, lax, 9-11-flowered, rachis 4-7 cm. long, pedicels
3 cm. long; bracts below the flowers, green, then dull yellow, finally
brunnescent, spur 7-8 mm. long, 3 mm. thick, the alate parts about
as long, lanceolate, acute; bractlets sepaloid, the sepals themselves
4 mm. long, 5 mm. wide; buds obovoid, rounded, dull yellowish
color, 9 mm. long, 6 mm. thick. — Beautiful species affine 5. pachy-
phylla (author). F.M. Neg. 9721.
Hudnuco: Near Monz6n, 1,600 meters, Weberbauer 3508, type;
255.
4. NORANTEA Aublet
Reference: Gilg & Werdermann, Pflanzenfam. ed. 2. 21: 100-101.
1925.
Racemes terminal, elongate. Bracts connate to base or middle
of pedicels, cucullate or sac-like. Petals lightly cohering basally
or free. Anthers linear or oblong. Ovary 3-5-celled, the sessile
stigma 3-5-radiate. Fruit fleshy-coriaceous. — Since the Museum
specimens of this group were lent abroad, without advising me until
just before completing this number, it has been impossible, without
delaying publication unduly, to revise my tentative synopsis com-
piled many years ago, or to ask the borrower to contribute the genus.
However, the few Peruvian species have been clearly discriminated
by Gilg and Werdermann, I.e. and in acknowledging my indebted-
712 Field Museum of Natural History— Botany, Vol. XIII
ness I record my pleasant remembrances of two personalities,
gracious as able.
Omitted as essentially nomina nuda are N. cordachida G. Don
and N. obovata (R. & P.) G. Don, Gen. Syst. 1: 625. 1831.
Bracts mostly sessile.
Bracts cylindrical, the length several times exceeding the diameter.
Inflorescence more or less clustered.
Pedicels about 5 mm. thick N. macroscypha.
Pedicels slender.
Leaf glands in 4 irregular series N. sandiensis.
Leaf glands in 2 series N. Eoetvoesorum.
Inflorescence umbelliform.
Leaves broadly obovate with 1 row each side of midrib of
open glands N. Pardoana.
Leaves about oblong, with seriate glands near margin or
scattered.
Glands few, 1-seriate.
Leaves scarcely acuminate N. peduncularis.
Leaves acuminate N. curva.
Glands many, scattered N. Uleana.
Bracts not or scarcely longer than thick N. magnifica.
Bracts mostly long-stiped (unless in N. anomala). ■
Flowers sessile.
Inflorescence leaves to 9 dm. long, lateral nerves prominent.
N, cacabifera.
Inflorescence leaves smaller, lateral nerves inconspicuous.
N. anomala.
Flowers shortly pedicellate.
Bracts at pedicel base; petioles short. ..... .N. haematoscypha.
Bracts above the middle; petioles elongate N. japurensis.
Norantea anomala HBK. Nov. Gen. & Sp. 7: 218. pi. 6A7h\s.
1825.
Shrubby, with oblong or more or less obovate or lanceolate,
leaves 6-10 cm. long, scarcely 3 cm. wide (petioles to 1 cm. long),
shortly acuminate or obtuse, with numerous small glands and a few
large open ones; spikes or racemes to 3 dm. long, the flowers sessile F
Flora of Peru 718
or subsessile; bracts hemispheric-galeate, subsessile or usually stiped,
above base of pedicel or below flower, 10-12 mm. long (including
stipe); corolla to 2.5 mm. in diameter; stamens 5-10, the anthers
cordate; ovary 3-celled. — Illustrated, Mart. Fl. Bras. 12, pt. 1:
pi. Jt8, jig. S (bract and flowers).
Peru (probably). Bolivia; Ecuador to Central America.
Norantea cacabifera [R. & P.] G. Don, Gen. Syst. 1: 625. 1831.
N. Caccabion Delp. Atti Soc. Ital. Sci. Nat. Milano 12: 182, 205.
1869; fide Wittmark.
Striking by the large oblong leaves, to 2 dm. long, 7 or 8 cm.
wide, prominently nerved and almost eglandular (petioles 1-1.5
cm. long), and the sphaeroid-obovoid much inflated bracts, rather
obliquely pyriform, 7-11 mm. long, 5-8 mm. thick, on slender stipes
6-11 mm. long, the aperture linear-oblong; flowers sessile, rather
large; stamens 6-8, the anthers ovate-cordate; ovary globose, ovules
few.— Illustrated, Mart. Fl. Bras. 12, pt. 1: pi. J^7, fig. 2 (bracts and
anthers).
Peru (probably). Adjacent Ecuador.
Norantea curva Macbr. Candollea 5: 385. 1934.
Resembles N. peduncularis; branchlets flexuose; leaves oblong-
or elliptic-lanceolate, acute or obtusely acuminate, about 13 cm.
long, 3-4.5 cm. wide, flexible but chartaceo vis-coriaceous; glands
mostly seriate and about 5 mm. distant or some approximate;
pedicels flexuose, slender, 5.5 cm. long; bracts 1.5-2 cm. long; calyx
scarcely 5 mm. high; flowers subovoid, exserted 5 mm. — Except as
noted apparently very much like N. peduncularis with obovate-
oblong thicker leaves, little if at all pointed, stouter pedicels, larger
calyx; the somewhat similar N. Uleana has tapering leaves with
much more approximate glands.
San Martin: San Roque, Williams 6999, type; also 7j^0.
Norantea Eoetvoesorum Richter ex Gilg & Werderm. Pflan-
zenfam. ed. 2. 21: 101. 1925.
Branches terete, 4-5 mm. thick, the canescent-brown bark not
exfoliating; petioles stout, 4-5 mm. long; leaves ovate-oblong,
unequally cuneate to base or more or less rounded on one side,
narrowed to the rounded, obtusish or retuse apex, to 14 cm. long,
5 cm. wide, coriaceous, green or brownish above, cinnamon-brownish
and the lateral nerves prominent beneath, the veins obscure on both
714 Field Museum of Natural History— Botany, Vol. XIII
opaque surfaces; glands 0.4-0.7 mm. broad, subseriate; racemes
terminal, dense, about 14-flowered, the stout pedicels 3 cm. long;
bracts 4-5 mm. stiped, narrowly tubular, 12-14 mm. long, 4-5 mm.
broad; sepals orbicular, entire and subscarious, 4-10 mm. wide;
buds subglobose. — Named for the family Eotvos, Hungarian
scientists.
San Martin: Vitoc, Ruiz & Pavdn, type.
Norantea haematoscypha Gilg, Bot. Jahrb. 42: 127. 1908.
Branches unequally striolate, 3 mm. thick; petioles 5 mm. long,
merging into the basally attenuate oblanceolate leaves, their apex
subrounded but acutely and minutely apiculate, mostly 5-7 cm.
long, to 17 mm. wide, with a few rather large glands on the upper
part obsoletely seriate, 2-3 mm. from the margin, the lateral nerves
rather obvious only beneath; racemes 10-14 cm. long, subspicate,
very dense, the spreading or subevolute pedicels 5-7 mm. long, the
bracts at base on stipes 17 mm. long, cochleate, blood-red, 8 or 9
mm. long, 5 mm. thick, the orifice little enlarged; bractlets ovate,
a little below the calyx; sepals 3-4 mm. wide; buds rounded, 5-6
mm. long, 3 mm. thick, green. — Elegant species allied to N. oxystylis
Baillon (Gilg). F.M. Neg. 9706.
Puno: Shrub formation, Sandia, 2,200 meters, Weberbauer 1057,
type; 237; Metcalf 30590 (det. Leonard). Sandia to Chunchusmayo,
Weberbauer 1156; 242.
Norantea japurensis Mart. Nov. Gen. & Sp. 3: 179. pi. 295.
1832.
Well-marked by the large obovate or oblong leaves decurrent
at base into petioles 2-2.5 cm. long, the blades 1-1.5 dm. long, 5-6
cm. wide, glands few, lateral nerves and veins weak; racemes dense,
subsecund, often several dm. (to 1 meter) long; bracts ample, often
roseate, ovoid or obovoid, 12-17 mm. long, 8 mm. in diameter,
borne above the middle of the 4-5 mm. long pedicels on stipes 5-8
mm. long, the aperture oblong; corolla only 3 mm. in diameter;
stamens about 30; ovary ovoid.
Peru (probably). Upper Amazonian Brazil. "Juri" (Martius).
Norantea macroscypha Gilg, Bot. Jahrb. 25: Beibl. 60: 31.
1898.
Flowering branches extremely thick or subfleshy; petioles about
5 mm. long and thick; leaves obovate, rounded both ends, the apex
Flora of Peru 715
apparently always emarginate, 12-16 cm. long, 5-9 cm. wide, mid-
nerve little prominent both sides, secondary nerves few, immersed
above, obvious beneath, veins not at all, the numerous mostly
small or minute glands subparallelly seriate at margin; racemes
elongate, the pedicels 4.5-5.5 cm. long, 6-8 mm. thick, erect-
spreading; bracts sessile, pendulous on lower third of pedicel, 5 cm.
long, medially 7 or 8 mm. thick, the orifice as wide, narrowed to
both ends; sepals 11-13 mm. long, 14-15 mm. broad. — Affine
probably A^. peduncularis; perhaps N. obovata (R. & P.) G. Don or
N. macrocarpa (R. & P.) G. Don but those with mucronulate leaves
and descriptions quite lacking in characterization (author) ; actually
the former is the earlier name and the latter is Marcgravia macro-
carpa.
Hudnuco: Chacahuasi, Ruiz & Pavdn, type. "Purumhigo."
Norantea magnifica Gilg, Bot. Jahrb. 42: 127. 1908.
Branches 4 mm. thick, striate; leaves obovate-oblong, long-
narrowed to subsessile base (or very short broad petiole), rounded
at apex but this acutely apiculate, opaque, rigid-coriaceous, 9-11
cm. long, 3-5 cm. wide, the large open glands on the upper part
often obsoletely seriate, about 1 cm. from margin, the lateral nerves
and veins inconspicuous; racemes thyrsoid, 10-12 cm. long, pedicels
4-5 cm. long, gradually enlarged above, erect-spreading; bracts
ferrugineous on lower quarter of pedicel, suberect, galeate, about
3 mm. stiped, 7 or 8 mm. long, 4 mm. thick; sepals 6-7 mm. long,
about as wide; buds 9-11 mm. long, 8 mm. thick, green. — Beautiful
2 meter shrub aflfine N. brasiliensis Choisy and N. costaricensis Gilg
(author). F.M. Neg. 9708.
Hudnuco: Near Monzon, 2,000 meters, Weberbaiier S528, t5T5e;
265.
Norantea Pardoana Weberb. & Gilg, Bot. Jahrb. 42: 125. 1908.
Branches 3 mm. thick, the pale papyraceous bark exfoliating
unequally, the leaves 6 or 7 mm. distant, obovate, cuneately nar-
rowed to the petiole, this to 3 mm. long and nearly as wide, rounded
at apex, 5-8 cm. long, 2.5-4 cm. wide, opaque both sides, sometimes
rigid, glands and nerves obvious £is in N. sandiensis; rachis scarcely
1 cm. long, 5-8-flowered, the erect pedicels 4.5-5 cm. long; bracts
tubular-saccate on lower third of pedicels (stipes 3 mm. long), little
enlarged above, 6 mm. thick, 2.5-3.5 cm. long, the suborbicular
orifice about 6 mm. across; sepals 6 mm. broad; buds 8-9 mm. long,
716 Field Museum of Natural History— Botany, Vol. XIII
5-6 mm. broad, the oblong petals greenish; fruits smooth, about
12 mm. thick, 9 mm. high, the persisting style 1 mm. long. — A 2
meter shrub apparently allied to N. mixta Tr. & PI. and N. macro-
stoma Gilg (authors) ; commemorates a president of Peru who aided
the earlier collecting of the senior author.
Junin: Above Huacapistana toward Palca, 1,900 meters, Weher-
bauer 1976, type; 247.
Norantea peduncularis Poeppig ex Wittm. in Mart. Fl. Bras.
12, pt. 1:238. 1878.
Flowering branches stout; petioles 4-5 mm. long; leaves obovate-
oblong or oblong, obtusish at base, rounded-subemarginate, 7-8
cm. long, 3-3.5 cm. wide, heavy coriaceous, with 8-10 sometimes
seriate glands; umbellate pedicels to 13, erect-arcuate, about 6 cm.
long, 1.5-2 mm. thick; bracts pendulous, tubular-saccate, ventrally
somewhat incurved, the large orifice nearly orbicular, the free part
sublingulate, 2.5 cm. long, 5 or 6 mm. across, the spurs sessile;
bractlets appressed, sepaloid; corolla drying yellowish; petals
strongly imbricate, oblong, the outer narrower; stamens about 25,
anthers linear; ovary 6-celled, style stout, stigma indistinctly lobed.
Feru: Poeppig 1^78. Colombia.
Norantea sandiensis Gilg, Bot. Jahrb. 42: 126. 1908.
Older branches whitish, smooth, unequally angled, the leaf
internodes 1.5-2 cm. long; leaves obovate or obovate-oblong,
cuneately narrowed to the very broad petiole, this about 5 mm.
long, rounded or often retuse, 7-10 cm. long, 2.5-4 cm. wide, some-
times rigid-coriaceous and with many lenticelliform glands mostly
in 2 series about 1 cm. from the margin, lateral nerves mostly
obscure both sides; racemes 1.5-2 cm. long, 3-8-flowered; pedicels
erect, 5-6 cm. long; bracts green below, reddish brown above,
tubular-saccate, clavately enlarged, 1-2 mm. long-stipitate, 5-7
mm. thick at apex, 4-4.5 cm. long, the suborbicular orifice about
1 cm. across; sepals 7-8 mm. in diameter; buds globose-ovoid, 10-12
mm. long, 7-8 mm. thick, broadly rounded; corolla large, the green
petals strongly imbricate.
Puno: Open shrub-formation, Azalaya, Sandla to Chunchusmayo,
1,500 meters, Weberbauer 13^3, type; 278. Sandia, Raimondi.
"Alchuyhuaturo" (Raimondi).
Norantea Uleana Pilger, Verb. Bot. Ver. Brandenb. 47: 155.
1905.
Flora of Peru 717
Flowering branchlets and subsessile leaves fleshy coriaceous, the
latter elliptic, little oblique at base, obtuse and shortly subcuspidate,
10-11 cm. long, 3.5-4.5 cm. wide, the nerves lightly impressed above,
beneath rather prominent, the lateral fine and borne at an acute
angle, the small but well-marked glands crowded toward the mar-
gins; flowers greenish, congested, subumbellate, peduncle nearly
none, pedicels about 16, around 6 cm. long; bracts on the lower third
of the pedicels, curved, reflexed, clavately enlarged apically, 3 cm.
long, the orifice wide; sepals and petals fleshy, the first broadly
rounded, the second obovate-elliptic, obtuse, 12 mm. long, half as
wide, stamens about 30. — Related to N. peduncularis Poeppig with
more obovate-oblong leaves, broader at apex, coarser nerves
(author). F.M. Neg. 9715.
Loreto: Cumbaso, Ule 6S36, type.
OUIINACEAE
Smooth shrubs or small trees closely resembling the Guttiferae
but the entire or dentate sometimes pinnate leaves stipulate often
conspicuously, free styles filiform, ovary cells biovulate, fruit inde-
hiscent, berry-like, the cotyledons distinct, thick or thin, the radicle
minute.
Sepals at least lightly adnate at base; leaves multipinnate, with
crystal sclereids (Foster) 1. Touroulia.
Sepals quite free; leaves unless the younger simple, crystal sclereids
none (Foster).
Leaves usually opposite; ovary 1-3-celled, styles 2 3. Quiina.
Leaves always verticillate (Peru) ; ovary 6-12-celled, styles several.
2. Lacunaria.
1. TOUROULIA Aublet
Trees, the imparipinnate leaves opposite, the stipules inter-
petiolate, and male or hermaphrodite flowers in terminal racemose
or paniculate cymes. Calyx lobes of male flowers imbricate, the
5 petals contorted, the stamens numerous; female sepals united only
at base, petals 5 or more; ovary 7-8-celled (styles 7 or 8) with 1-2
ovules in each cell. Fruit baccate, striate, 7-celled, with as many
villous or tomentose seeds. — Froesia Pires, Bol. T^n. Inst. Agron.
Norte, Bel^m 15: 22-25. 1948, has the divided leaves of this genus
but free sepals, fruit of 3 follicles, glabrous seeds, collected as near
i
718 Field Museum of Natural History— Botany, Vol. XIII
Peru as Sao Paulo de Olivenga; the parallel lateral nerves of the
13-15 leaflets are rib-like on the lower surface.
Adriance S. Foster, Amer. Journ. Bot. 37: 848-862. 1950, has
studied the venation of Touroulia and Froesia Pires, showing the
retention of the patterns of seedling leaf-venation in the terminal
part of adult leaflets of the former genus and the presence of crystali-
ferous sclereids in both genera (lacking otherwise in this family)
similar to those in the Ochnaceae and certain Linaceae.
Touroulia amazonica Pires & Foster, Bol. T^cn. Inst. Agron.
Norte, Bel^m 20: 49. 1950.
Small tree, the terete branches glabrescent, the branchlets and
inflorescences reddish tomentose including the sepals without;
leaves 2-5 dm. long, 13-17-foliolate, the petioles 5-15 cm. long,
petiolules 2 mm. long, the stipules 2, subulate, 3-5 mm. long;
leaflets oblong-lanceolate, obtuse or lightly attenuate at base,
acuminate, obscurely glandular-serrulate (with lens), coriaceous, the
lateral nerves about 20, acutely prominent beneath, the undulate
veins subparallel; sepals unequal, 5-6 mm. long, obtuse or rounded;
petals persisting, not resinous, glabrous, 10-12 mm. long, 5 mm.
wide; stamens about 50; fruiting (hermaphrodite) inflorescence 9
cm. long, pedicels (not articulate) 6-8 mm. long; fruits (immature)
globose, 11 mm. long, 9 mm. thick, 8-celled, 8-styled, the seeds
rusty tomentose. — Differs from T. guianensis Aubl. in the larger
hermaphrodite flowers, persisting not resinous petals, leaflets
coriaceous and veined as in species of Froesia, that is, directly cross-
paralleled between the lateral nerves (authors). Type an 8 meter
tree with trunk diameter of 5 cm. Found near Peru, as Sao Paulo de
Oliven^a. Illustrated, Pires & Foster, I.e. pis. 12 (plant), 13, lU
(leaf venation).
Peru (probably). Amazonian Brazil.
2. LACUNARIA Ducke
Reference: Ducke, Archiv. Inst. Biol. Veg. Rio Jan. 2: 169-171.
1935.
Trees with simple verticillate (in Peru) glabrous leaves and
rather large dioecious flowers in often slightly compound racemes
that are solitary or 2-3 in the axils. Sepals as petals imbricate, the
former 4, the latter 4-8. Stamens 20-40 or 70-170; ovary un-
developed in male flowers. Ovary 6-12-celled, the styles as many as
Flora of Peru 719
the 2-3-ovulate cells. Fruit a &-12-celled berry with well-developed
pericarpal cavities; seeds 1-3 in each cell with thin cotyledons,
copious endosperm.— See A. S. Foster, Bull. Torrey Club 78: 382-400.
1951, for an important contribution to an understanding of the
"Heterophylly and Foliar Venation in Ldcunaria," in which he
shows that early leaves of seedlings in two species are deeply pin-
natifid and traces the transition (and venation) to the simple mature
leaves. Ducke's work consists of a key which, however, includes
ranges as well as considerable characterization in contrasting clearly
each species and this with my appreciation is summarized here.
Petals 8 (7) ; stamens about 100; leaves with 30-40 secondary nerves,
puberulent beneath L. pukhrinervis.
Petals 4 or 5; stamens about 35 or fewer (known); leaves glabrous,
secondary nerves 8-25.
Secondary nerves 8-12 L. acreana.
Secondary nerves 15-25 L. Jenmani.
Lacunaria acreana Ducke, Archiv. Jard. Bot. Rio Jan. 4:
141. 1925.
Glabrous unless the branchlet tips and the racemes, these
rufo- or ferrugineous tomentose including the orbicular outer sepals;
stipules linear-subulate, acuminate or setaceous, 0.5-1.5 cm. long;
petioles 1.5-2 cm. long or longer; leaves oblong, rarely somewhat
obovate, long-attenuate to base, obtuse or more or less and shortly
acuminate, 1-2 dm. long, 4-6 cm. wide, subcoriaceous, the 9-12
slender nerves rather prominent beneath; male inflorescence un-
known; rachis to 2 cm. long, pedicels finally often defiexed, to 2 cm.
long; sepals 4, 4-5 mm. long, inner more elliptic, 7 mm. long, tomen-
tose only at base; petals whitish, 4-5, obovate-orbicular, 6 mm.
long; ovary strongly striolate, cells as styles 12, ovules 2. — Type
5-6 meters high. Close to or possibly conspecific with L. macro-
stachya (Tul.) A. C. Smith, Trop. Woods 58: 31. 1939, from the
Solimoes region (not Peru, Tulasne, Ann. Sci. Nat. s6r. 3. 11: 162.
1849). To be expected in Peru, as also L. coriacea A. C. Smith, I.e.,
with somewhat smaller leaves, pedicels (male) 4-5 mm. long. F.M.
Neg. 9755. Illustrated, Mart. Fl. Bras. 12, pt. 1: pi. 110 {Quiina
macrostdchya).
Rio Acre: Nova Olinda to S5o Luiz, (Kuhlmann, type, Herb.
Jard. Bot. Rio Jan.).
720 Field Museum of Natural History— Botany, Vol. XIII
Lacunaria Jentnani (Oliver) Ducke, Archiv. Jard. Bot. Rio
Jan. 5: 171. 1930. Touroulia Jenmani Oliver in Hook. Icon. pi. 1998.
1891.
Branchlets and petioles (to 4 cm. long) early rufous puberulent,
becoming glabrate; stipules linear-subulate, rigid, to 1 cm. long;
leaves more or less obovate or oblong-elliptic, cuneate or acutely-
narrowed at base, acutely or subobtusely short-acuminate, often
1.5-3 dm. long, 5(4)-10(12) cm. wide, rarely remotely denticulate,
coriaceous, somewhat lustrous, the 15-25 lateral nerves notably
prominent only beneath; inflorescences rufous tomentose compound
racemes, often about 1.5 dm. long, with short verticillate lateral
branchlets, the male more densely flowered, the pedicels only 4-5
mm. long, the female pedicels 1-2.5 cm. long, accrescent in fruit;
sepals 4, concave, the 2 interior to 5 mm. long, sparsely pilose
marginally toward tip (male) or to 7 mm. long, sparsely pilose
(female); petals 5-7, obovate-oblong, 5 and 8 (female) mm. long;
stamens 20-35, slender filaments 3-6 mm. long, anthers subglobose;
ovary subcylindric, sulcate, cells as styles 10-13, the latter 2 mm.
long, broadened into the plumose stigmas; fruit key-shaped or
subglobose, to 12 cm. long, 9 cm. thick, the pericarp 12 mm. thick
with as many cavities as fissures in the resiniferous exocarp; seeds
3 to a cell, ovoid, 1-1.5 cm. long, covered with long rufous pu-
bescence.
Loreto: Iquitos (fide Ducke). Amazonian Brazil.
Lacunaria pulchrinervis Ducke, Archiv. Inst. Biol. Veg. Rio]
Jan. 2: 169. 1935.
Small tree, the slender branchlets, conspicuous secondary paral-
lel leaf nerves beneath (33-43) and racemiform male inflorescence
(female unknown) reddish puberulent or tomentulose; petioles 4-6
cm. long; leaves 2.5-4 dm. long, 9-15 cm. wide, lanceolate elliptic
or oblong-obovate, attenuate to petiole or obtuse basally, shortly
acuminate, subentire, coriaceous, subconcolorous but rather bright
deep green between the nerves beneath; verticils of 1-2 flowers, the
pedicels to 3 cm. long; sepals 4, coriaceous, 5-6 mm. long; petals 8,
white, strongly reflexed after an thesis, to 1.5 cm. long, 8-11 mm.
wide, distinctly nerved, glabrous except ciliate margined but notably
lustrous with a resinous substance; stamens white, 105-110 (counted
in 3 flowers). — After author, who compared it with L. grandiflora
Ducke of the Rio Negro with opaque narrower petals. Since the
Flora of Peru 721
type (one collection by Ducke) was from Sao Paulo de Oliven^a,
Brazil, the species in all probability is also within Peru.
Peru (probably). Amazonian Brazil.
3. OUIINA Aublet
Leaves opposite or rarely verticillate, often elegantly and finely
lined by the transverse veins between the pinnate nerves. Stipules
narrow, rigid, often geminate. Flowers polygamo-dioecious, small,
disposed in short axillary panicles or terminal racemiform or di-
varicate-racemose inflorescences or fascicles. Sepals and petals 4 or
5 or the latter 8, both imbricate. Stamens many, connate only at
base if at all; filaments filiform, anthers small, oblong or globose-
didjnnous. Ovary 2-3-celled with as many apically stigmatose
distinct styles; ovules ascending. Fruit baccate, the endocarp
fulvous, the 2-4 ovoid or globose seeds tomentose, the cotyledons
thick. — Ducke, Archiv. Jard. Bot. Rio Jan. 5: 172. 1930, pointed
out that most if not all species are polygamo-dioecious and therefore,
as noted by A. C. Smith, Trop. Woods 58: 25. 1939, the primary
divisions of the keys of Tulasne and Engler are not dependable.
As usual, it has been impossible to consider more than in a general
way the extra-Peruvian species that may be found to antedate some
described from Peru which will be disclosed only by a revision of
the group.
Leaves verticillate, long-spatulate, those of new shoots pinnatifid.
Q. pteridophylla.
Leaves opposite, all entire or not pinnatifid.
Inflorescences sessile; pedicels to 1 cm. long, leaves to 4 dm. long.
Q. grandifolia.
Inflorescences not sessile; pedicels and leaves shorter.
Inflorescences many-flowered, crowded with the subequal
stipules Q. Tessmannii.
Inflorescences elongating or reduced to 1-few flowers.
Leaves abruptly caudate, the acumen 1-2 cm. long, 1-3 mm.
wide Q. amazonica.
Leaves if acuminate not caudate as above.
Principal lateral nerves often to 15 or more subequally
marked and distant.
Leaves narrowed to base; flowers small. . . .Q. peruviana.
722 Field Museum of Natural History— Botany, Vol. XIII
Leaves rounded to acute base; flowers 5-6 mm. long.
Q. Klugii.
Principal lateral nerves often about 10 or in any case
unequally marked and distant.
Leaves glabrous.
Leaves very lustrous above, about elliptic. . .Q. nitens.
Leaves slightly lustrous, ovate- or oblong-lanceolate.
Q. rhytidopus.
Leaves puberulent beneath, mostly even in age.
Q. pubescens.
Ouiina amazonica A. C. Smith, Trop. Woods 58: 27. 1939.
Glabrous except sometimes the branchlet tips, petioles (2-5 mm.
long) and sepals a little puberulent, the leaves slenderly caudate-
acuminate, the slender inflorescences hermaphrodite, usually 2.5-5
cm. long; stipules subulate-setaceous, 3-5 mm. long; leaves elliptic,
acute, obtuse or rounded at base, the abrupt acumen obtuse or
subacute, 1-2 cm. long, 1-3 mm. wide, obscurely if at all crenate,
chartaceous, 6-14 cm. long, 2.5-5.5 cm. wide, the 9-12 lateral nerves
acute elevated above, very conspicuous as the veins beneath; flowers
mostly solitary, pedicels 4-5 mm. long, articulate toward the little
enlarged base; sepals 3 or 4; deltoid-ovate or suborbicular, about
1 mm. wide; petals 3-6, membranous, entire, 1.5-1.7 mm. long and
wide; stamens about 9, uniseriate, styles 2, little longer than ovary
(male inflorescences and flowers similar to above description but
ovary none, stamens 15-17 fasciculate); fruits obscurely striate,
11-21 mm. long, 7-12 mm. wide, mucronulate, the oblong-ellipsoid
seeds minutely villosulous, 4-10 mm. long, 3-5 mm. broad. — Allied
to Q. Poeppigiana Tul.; not uncommon in the upper Amazon Basin.
Loreto: Sierra del Pongo, Mex 
