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Full text of "Flora of Peru"

FLORA OF PERU 

BY 

MILDRED E. MATHIAS 

ASSOCIATE PROFESSOR OP BOTANY 
UNIVERSITY OP CALIFORNIA, LOS ANGELES 

AND 

LINCOLN CONSTANCE 

PROFESSOR OF BOTANY 
TJNIVERSITY OF CALIFORNIA, BERKELEY 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART V-A, NUMBER 1 

JULY 31, 1962 
PUBLICATION 953 



Library of Congress Catalog Card Number: 36-10^26 



PRINTED IN THE UNITED STATES OF AMERICA 
BY CHICAGO NATURAL HISTORY MUSEUM PRESS 



t.5-4 
5. / 



FLORA OF PERU 



MILDRED E. MATHIAS AND LINCOLN CONSTANCE 1 



UMBELLIFERAE. Carrot Family 

References: 0. Drude, Umbelliferae in Engl. & Prantl, Pflanzenf. 
3 (8) : 63-250. 1897. H. Wolff, Umbelliferae-Saniculoideae in Engl. 
Pflanzenr. 61 (IV. 228) : 1-305. 1913. H. Wolff, Umbelliferae-Api- 
oideae, 43 (IV. 228) : 1-214. 1910. H. Wolff, Umbelliferae-Apiodeae, 
90 (IV. 228): 1-398. 1927. Mathias & Constance, Umbelliferae, 
North American Flora 28B: 43-374. 1944-45. 

Plants usually herbaceous, rarely woody at base, acaulescent 
or caulescent, annual or perennial, with commonly hollow stems. 
Leaves alternate, or rarely opposite, or basal, compound or some- 
times simple, usually much incised or divided, with usually sheathing 
petioles. Flowers small, regular, in simple or compound umbels, 
or the umbels sometimes proliferous or capitate; rays sometimes 
subtended by bracts forming an involucre; umbellets usually sub- 
tended by bractlets forming an involucel. Calyx tube wholly adnate 
to ovary; calyx teeth obsolete or small. Petals 5, usually with an 
inflexed tip. Stamens 5, inserted on an epigynous disk. Ovary 
inferior, bilocular, with one anatropous ovule in each locule; styles 2, 
sometimes swollen at base, forming a stylopodium. Fruit consisting 
of 2 carpels united by their faces (commissure), flattened or com- 
pressed dorsally (parallel to commissure), laterally (at right angles 
to commissure), or terete, each carpel with 5 primary ribs, one 
dorsal, two lateral (on the edges of commissure) and two between 
dorsal and lateral ribs (intermediate ribs), and rarely with secondary 
ribs, the ribs filiform to broadly winged, thin or corky; vittae (oil 
tubes) obsolete or present in the intervals (spaces between ribs) 
and on commissural surface, rarely also in pericarp; carpels 1-seeded, 
splitting apart at maturity, usually suspended from summit of a 
slender prolongation of the axis (carpophore). Embryo small; endo- 
sperm cartilaginous. 

Umbelliferae are widely distributed in temperate and moun- 
tainous areas of the world, especially in the northern hemisphere. 

1 The authors wish to acknowledge partial support from grant G-13393 by the 
National Science Foundation. 



4 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

The family includes a number of common foods and condiments 
such as carrot, parsnip, celery, parsley, anise, caraway, dill, chervil, 
fennel, and lovage. Many of these have escaped from cultivation 
and have become weedy components of floras in various parts of 
the world. In Peru about 20 per cent or 17 species of the family 
are these escaped cultivars and other widespread weedy species. 
The incidence of endemism in Peruvian Umbelliferae is amazingly 
high, some 40 per cent of the species being known only from that 
country. Only five of the endemic species and varieties and only 
five widely ranging species are of northern-hemisphere affinity. The 
others belong mostly to Andean genera. Twenty-nine species extend 
outside of Peru and are referred to genera widely distributed in 
the southern hemisphere. 

The high endemism may be in large part a reflection of our 
still inadequate knowledge of the flora, many of the species being 
known from single collections. Range extensions are common and 
the discovery of new species frequent as a new area or a new collec- 
tion is studied. Certainly the umbelliferous flora of Peru is still 
poorly known and comprehensive collections will continue to im- 
prove our knowledge. It is hoped that this preliminary revision 
of the family will encourage collectors and Peruvian students of 
the family. 

Umbels all simple, lax and open to capitate, variously arranged. 
Cushion plants of high altitudes. 

Umbels capitate; flowers sessile; fruit scaly Eryngium. 

Umbels lax; flowers pedicellate; fruit glabrous or pubescent. 
Fruit with distinct vittae in the intervals but without a 
woody endocarp; carpophore present; leaves pinnatisect. 

Oreomyrrhis. 

Fruit lacking vittae in the intervals but with a prominent 
woody endocarp; carpophore lacking; leaves not pinnat- 
isect Azorella. 

Erect, ascending, or creeping perennials, biennials or annuals, 

not conspicuously cespitose or matted to form cushions. 
Umbels capitate (flowers stipitate in Sanicula); fruit promi- 
nently scaly or uncinate-prickly (Saniculoideae) . 
Fruit variously covered with scales or tubercles; flowers all 
perfect and sessile; leaves neither palmately lobed nor 
parted Eryngium. 



FLORA OF PERU 5 

Fruit uncinate-prickly; flowers either perfect or staminate 
in the same umbellet, at least the latter stipitate; leaves 
palmately 3-5-parted Sanicula. 

Umbels lax; fruit glabrous, pubescent or glochidiate. 

Foliage of septate linear phyllodes; fruit with prominent aeri- 
form tissue Lilaeopsis. 

Foliage various, but always with an evident leaf blade; fruit 
lacking prominent aeriform tissue. 

Stipules or stipular processes evident at base of petiole. 

Foliage and inflorescence stellate-pubescent or stellate- 
glochidiate Bowlesia. 

Foliage and inflorescence glabrous or pubescent, but nei- 
ther stellate-pubescent nor stellate-glochidiate. 

Erect annuals; cauline leaves opposite Spananthe. 

Creeping perennials; leaves basal or alternate. 

Hydrocotyle. 

Stipules lacking, the petioles sheathing at base. 
Plants prominently caulescent, glabrous and glaucous; 
petals with a prominent gland; involucral bracts 
free or largely so; carpophore lacking. 
Annuals; petal apex merely acute, not prominently 
inflexed; stylopodium obsolete Domeykoa. 

Perennials; petal apex long-inflexed; stylopodium evi- 
dent Eremocharis. 

Plants acaulescent or subcaulescent with no or few cau- 
line leaves; petals glandless; involucral bracts con- 
nate below; carpophore present Oreomyrrhis . 

Umbels regularly compound (or some umbels, especially the lateral, 
occasionally simple). 

Fruit armed with bristles, prickles, or glochids (see also Sanicula). 

Fruits caudate at base, several times longer than broad; leaf- 
lets ovate, 15-40 mm. long Osmorhiza. 

Fruits rounded or truncate, neither caudate at base nor more 
than 2-3 times as long as broad; leaflets filiform to lance- 
olate, 2-12 mm. long. 

Involucral bracts prominent, dissected; rays prominent; car- 
pels flattened dorsally; leaflets linear to lanceolate. 

Daucus. 



6 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Involucral bracts entire or lacking; rays abortive; carpels 

flattened laterally; leaflets filiform Torilis. 

Fruits unarmed, glabrous or pubescent. 

Flowers yellow, greenish-yellow, maroon, deep red or purple, 
but never white. 

Leaves decompound with filiform ultimate divisions. 

Perennial or biennial; petals obtuse, rounded at apex; car- 
pophore divided to base; petioles wholly sheathing. 

Foeniculum. 

Annual; petals shallowly emarginate; carpophore merely 
bifid; petioles sheathing only below Ridolfia. 

Leaves variously divided but with mostly oblong or lanceolate 
to orbicular leaflets. 

Perennial; involucel longer than flowers and sometimes 
longer than fruit. 

Fruit ellipsoid-cordate; involucel of conspicuous filiform 
bractlets to 3 cm. long, exceeding mature fruit. 

Neonelsonia. 

Fruit oblong to ovoid; involucel of linear to obovate 
bractlets to 1.5 cm. long, shorter than fruit. 

Arracacia. 

Annual or biennial; involucel lacking or shorter than 
flowers. 

Leaves ternate-pinnately or pinnately decompound, the 
leaflets linear; fruit 2-4 mm. long, compressed later- 
ally, all ribs filiform; involucel present. 

Petroselinum. 

Leaves pinnately compound, the leaflets oblong to ovate, 
to 8 cm. broad; fruit 5-6 mm. long, flattened dor- 
sally, the lateral ribs winged; involucel lacking. 

Pastinaca. 

Flowers white, or sometimes greenish or rosy or with a colored 
mid vein. 

Petals unequal, the marginal petals radiant; ripe carpels co- 
herent, not separating readily Coriandrum. 

Petals subequal; ripe carpels separating from carpophore. 

Rays and pedicels borne on discoid receptacles, rigidly 
contracted in fruit . . . . Ammi. 



FLORA OF PERU 7 

Neither rays nor pedicels borne on discoid receptacles and 
not contracted in fruit. 

Both involucre and involucel lacking; umbels all sessile 
or short-pedunculate Apium. 

Involucel, and often also an involucre, present; umbels 
(at least the terminal) prominently pedunculate. 

Stout introduced weedy annuals with purple-spotted 
stems; fruit ribs prominently undulate-winged. 

Conium. 

Native perennials, without conspicuously spotted 
stems; fruit ribs normally un winged and plane. 

Carpels compressed dorsally; involucel bractlets 
shorter than pedicels Paraselinum. 

Carpels compressed laterally, or terete; involucel 
bractlets longer than pedicels, sometimes slightly 
exceeding flowers and fruit. 

Petals purple, maroon, or greenish (rarely whitish), 
with a conspicuously narrower inflexed apex; 
styles slender; seed face sulcate. . .Arracacia. 

Petals white (often with a colored midvein), ob- 
tuse or acute, but without a prominent in- 
flexed apex; styles short; seed face shallowly 
concave Niphogeton. 

AMMI L. 

Slender, erect, herbaceous, caulescent, branching, essentially gla- 
brous annuals, biennials, or perennials, from slender taproots. 
Leaves petiolate, membranaceous, ternate-pinnately or pinnately 
dissected, the leaflets filiform to lanceolate. Petioles sheathing. 
Inflorescence of loose compound umbels; peduncles terminal and 
axillary. Involucre of numerous entire or divided bracts. Rays 
numerous, spreading-ascending. Involucel of numerous entire bract- 
lets shorter or longer than pedicels. Pedicels spreading. Flowers 
white; petals ovate to obovate with a broad, shallowly bilobed, 
inflexed apex; calyx teeth minute; styles slender, more than twice 
as long as the depressed-conic stylopodium. Carpophore entire or 
2-parted to base, flat. Fruit oblong to ovoid, compressed laterally, 
glabrous; ribs acute; vittae solitary in the intervals, 2 on commissure; 
seed subterete in transection, the face plane. 



8 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A Mediterranean genus. Two species have been cultivated as 
ornamentals and for medicinal purposes. Both have been sporadi- 
cally introduced in the western hemisphere but only one is reported 
from Peru. 

Ammi visnaga (L.) Lam. Fl. Fr. 3: 462. 1778. Daucus visnaga 
L. Sp. PL 242. 1753. Apium visnaga Crantz, Class. Umbell. 104. 
1767. Selinum visnaga Krause in Sturm, Fl. Deutsch. ed. 2. 12: 44. 
1904. Carum visnaga K.-Pol. Bull. Soc. Nat. Mosc. II. 29: 198. 1916. 

Erect biennial, 2-8 dm. tall; leaves deltoid, the blades 5-20 cm. 
long, pinnately decompound, the ultimate divisions linear to filiform, 
setaceous to cuspidate-acute, divaricate, entire, 5-35 mm. long, 0.5-1 
mm. broad; petioles about 10 cm. long; cauline leaves ternately 
or pinnately dissected; peduncles 8-14 cm. long; involucral bracts 
equaling or exceeding rays; rays 60-100, subfiliform, unequal, 2-5 
cm. long, spreading in flower but rigidly contracted in fruit, borne 
on a discoid receptacle; involucel of numerous setiform-acute, entire 
bractlets, ascending at maturity and equaling or exceeding pedicels; 
pedicels numerous, filiform, unequal, 3-13 mm. long, spreading 
in flower but rigidly contracted in fruit, borne on a small discoid 
receptacle; carpophore entire; fruit oblong-ovoid to ovoid, 2-2.5 mm. 
long, 1.5-1.7 mm. broad. 

Arequipa: Pr. Islay: Lomas de Mollendo, 400 m., Angulo & 
Sagastegui 2589; lower edge of loma zone, foot of hills east of Mol- 
lendo, 50 m., West 8205. Cajamarca: Pr. Chota: cerca a Llama, 
2000 m., Ferreyra 8401. Pr. Cajamarca: alrededores de la ciudad, 
terreno humedo, Ridoutl 316. La Libertad: Pr. Santiago de Chuco: 
Andamarca, Paredes 1587. Pr. Huamachuco: Hda. Yanazara, en 
borde de camino, 2650 m., Ldpez & Sagastegui 2857. Lima: Pr. 
Lima: between Lima and Vitarte, edge of field, Asplund 13906; re- 
particion near Lima, Hood. Moquegua: Pr. Mariscal Nieto: Valle 
de Ilo, desembocadura del Rio Ilo, 10-20 m., Ferreyra 11629. 

Known locally as bisnaga, cicuta, and culantro de burro. Eurasia, 
sporadically introduced, Pennsylvania to Florida, Oregon to Cali- 
fornia, Oaxaca, Bermuda, temperate South America. 

APIUM L. 

Celeri Adans. Fam. PL 2: 498. 1763. Helosciadium Koch, Nova 
Acta Acad. Leop.-Carol. 12: 125. 1825. Ciclospermum Lag. Amen. 
Nat. 101. 1821. 



FLORA OF PERU 9 

Slender or stout, erect, ascending, or prostrate, herbaceous or 
somewhat woody at base, caulescent (in our species), usually branch- 
ing, glabrous annuals, biennials or perennials, from taproots or creep- 
ing rootstocks, rooting at nodes. Leaves petiolate, membranaceous, 
simply pinnate to ternate-pinnately decompound, the leaflets sub- 
orbicular to filiform. Petioles sheathing. Inflorescence of loose 
to subcompact, compound or rarely simple umbels. Peduncles lat- 
eral and terminal, or some or all of the umbels sessile. Involucre 
wanting to conspicuous. Rays usually rather few, spreading- 
ascending. Involucel wanting to conspicuous. Pedicels spreading 
or ascending. Flowers white or greenish; petals ovate to sub- 
orbicular, acute, plane, or the apex somewhat inflexed; calyx teeth 
minute or obsolete; styles short, spreading or divaricate, the stylo- 
podium short-conic to depressed. Carpophore entire or shortly 
bifid. Fruit oblong-oval to orbicular or ellipsoid, compressed later- 
ally and somewhat constricted at commissure, glabrous or rarely 
somewhat setulose; ribs filiform, subequal, obtuse, prominent; vittae 
solitary in the intervals, 2 on commissure; seed sub terete in tran- 
section, the face plane. 

A genus of some 30 species, principally of Eurasia and the 
southern hemisphere. 

Plants perennial; leaves pinnate, the leaflets ovate to suborbicular. 

A. graveolens. 
Plants annual; leaves pinnately decompound, the leaflets linear to 

filiform. 
Leaflets filiform, several times longer than broad; fertile flowers 

mostly more than ten in each umbellet A. leptophyllum. 

Leaflets linear, not several times longer than broad; fertile flowers 
mostly fewer than ten in each umbellet A. laciniatum. 

Apium graveolens L. Sp. PI. 264. 1753. Seseli graveolens Scop. 
Fl. Cam. ed. 2. 1: 215. 1772. Slum apium Roth, Fl. Germ. 1: 128. 
1788. S. graveolens Vest, Man. Bot. 517. 1805. Apium vulgare 
Bubani, Fl. Pyren. 2: 344. 1900. Selinum graveolens Krause in 
Sturm, Fl. Deutsch. ed. 2. 12: 38. 1904. Celeri graveolens Britt. in 
Britt. & Brown, 111. Fl. ed. 2. 2: 660. 1913. Carum graveolens K.-Pol. 
Bull. Soc. Nat. Mosc. II. 29: 199. 1916. 

Plants perennial, the stems erect or ascending, about 5-15 dm. 
high, from a taproot; leaves oblong to obovate, the blades 7-18 cm. 
long, 2-5 cm. broad, pinnate with few pairs of ovate to suborbicular 



10 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaflets, 2-4.5 cm. long, 2-5 cm. broad, usually 3-lobed to near mid- 
dle; petioles 3-26 cm. long; cauline leaves frequently cuneate, some- 
times laciniate; umbels sessile or short-pedunculate, the peduncles 
up to 1.2 cm. long; involucre wanting; rays 7-16, 0.7-2.5 cm. long; 
involucel wanting; pedicels 1-6 mm. long; calyx teeth minute but 
evident; carpophore shortly bifid; fruit suborbicular to ellipsoid, 
about 1.5 mm. long, 1.5-2 mm. broad. 

Illustrations: Bot. Jahrb. 28: pi 2, fig. 30. 1899 (fruit section). 

Arequipa: Pr. Arequipa: alrededores de los Banos de Jesus, cerca 
a Arequipa, 2400-2500 m., Ferreyra 12071. Lima: Pr. Lima: cer- 
canias de Lima, Isern 45 (Cuatrecasas 2466) ; Callao, mouth of Rio 
Rimac, Soukup 2136; Hacienda Balconcillo, 157 m., Ridoutt. 

The cultivated celery, locally known as apio. Europe, adventive 
throughout the western hemisphere. 

Apium laciniatum (DC.) Urb. in Mart. Fl. Bras. 11 (1) : 343. 
1879. Helosciadium laciniatum DC. Me*m. Soc. Phys. Ge"neve 4: 495. 
1829. Sison laciniatum 1'Her. ex DC. I.e. in synon. Helosciadium 
laciniatum var. humile Hook. & Arn. Bot. Misc. 3: 353. 1833. H. 
laciniatum var. elatius Hook. & Arn. I.e. H. gracile Clos ex Gay, Fl. 
Chile 3 : 124, pi. 31 , /. 2. 1847 (including varieties a and 0) . H. deserti- 
cola Phil. Fl. Atacam. 26. 1860. H. biternatum Phil. Anal. Univ. Chile 
85. 718. 1894. Apium biternatum Reiche, Anal. Univ. Chile 104: 
836. 1899; Fl. Chile 3: 115. 1902. A. laciniatum var. gracile Reiche, 
Anal. Univ. Chile 104: 835. 1899; Fl. Chile 3: 114. 1902. A. lacini- 
atum var. gracile f. deserticolum Wolff, in Engl. Pflanzenr. 90 (IV. 
228) : 56. 1927. A. laciniatum var. gracile f . elatius Wolff, I.e. A. la- 
ciniatum var. hispidulum Wolff, I.e. 

Plants slender, annual, branching above, 1.5-2 dm. high; leaves 
oblong-ovate, the blades 0.5-3 cm. long, 0.5-3 cm. broad, ternate, 
then 1-2-pinnate, the segments linear, up to 4 mm. long, about 1 mm. 
broad, entire to incised-lobed; petioles to 3.5 cm. long, the sheath 
conspicuously white-scarious-margined ; cauline leaves similar; um- 
bels axillary, sessile, 2-3-rayed, the rays unequal, to 10 mm. long; 
involucre and involucel wanting; fruiting pedicels less than 10, un- 
equal, to 4 mm. long; flowers greenish to pinkish-white; calyx teeth 
inconspicuous; carpophore entire; fruit ovoid, 1.5-2 mm. long, 1-2 
mm. broad, glabrous, to rugulose or hispidulous. F.M. Negs. 8859, 
31809. 

Illustrations: Gay, Hist. Chile, Atlas, pi. 31 J.I. 1847 (as Helo- 
sciadium gracile). 



FLORA OF PERU 11 

Ancash: Pr. Casma: lomas de Lupin entre Huacho y Casma, 
loma arcillosa, 540 m., Ferreyra 8673; Mathias 3672. Arequipa: Pr. 
Islay: Lomas de Mollendo, 500-600 m., Ferreyra 12615; near Mol- 
lendo, 250 m., Hutchison 1861. Pr. Caraveli: Lomas de Pongo 
(Acari), 750 m., Velarde 11*61; Lomas de Capace, km. 648 from Lima, 
ca. 2 km. south of Chala, 200 m., Hutchison 1290. La Libertad: 
Pr. Trujillo: Cerro Cabezon, loma arenosa, 500 m., Angulo 1192; 
aired edores de Ascope, 120 m., en borde de acequia, L6pez 188. 
Lima: Pr. Lima: Lomas de Quilmana, al sur de Lima, 480 m., Ferreyra 
3990; Miraflores, cerca a Lima, 157 m., Maisch; Lima, terrenes de 
cultivo, Cali; Lomas de Amancaes, cerca a Lima, 400 m., Ferrey- 
ra 11828; Lomas de Caracoles, entre Lima y Canete, 200-300 m., 
Ferreyra 11875; San Geronimo, about 500 ft., rocky hillside, Mac- 
bride 5911; vicinity of Lima, San Agustin, Asplund 13832. Pr. 
Chancay: 5 km. north of Barranca, Stork, Beetle & Morrison 9094,' 
Lomas de Lachay, cerca a Chancay, 300-400 m., Cerrate 859; Lomas 
de Pacar (Pativilca), 400 m., Velarde 2309; Lomas de Chancay, 
arcilloso-pedregoso, 300 m., Ferreyra 8737. Pr. Canete: hills of 
Perico near Mala, rocky places, 12-20 m., Vargas 9316. Moquegua: 
Pr. Mariscal Nieto: Lomas de Ilo, 250-350 m., Ferreyra 11604. 
Tacna: Pr. Tacna: Lomas de Morro Sama, 600-700 m., Ferreyra 
1 2646. Without locality, Dombey. 

Frequent in rocky places in loma vegetation to Chile. Occasion- 
ally a garden weed known as capuchilla. 

Apium leptophyllum (Pers.) F. Muell.; Benth. & Muell. Fl. 
Austr. 3: 372. 1866. Sison ammi L. sensu Jacq. Hort. Vindob. 2: 
95. 1773, not S. ammi L. 1753. Sium nodiflorum Walt. Fl. Car. 115. 
1788, not S. nodiflorum L. 1753. Seseli ammi Savi, Due Cent. 71. 
1804. Pimpinella leptophylla Pers. Syn. PI. 1: 324. 1805. Aethusa 
leptophylla Spreng. Umbell. Prodr. 22. 1813. Pimpinella domingensis 
Willd. ex Spreng. in Roem. & Schult. Syst. Veg. 6: 411. 1820, as synon. 
Ciclospermum ammi Lag. Amen. Nat. 2: 101. 1821. Helosciadium lep- 
tophyllum DC. Me"m. Soc. Phys. Geneve 4: 493. 1829. Apium ammi 
Urban, in Mart. Fl. Bras. 11 (1) : 341. 1879. A. leptophyllon Maza, 
Noc. Bot. Sist. 74: 1893. A. ammi var. leptophyllum Kuntze, Rev. 
Gen. 3 (2) : 111. 1898. Selinum leptophyllum Krause in Sturm, Fl. 
Deutsch. ed. 2. 12: 28. 1904. Helosciadium ammi Britt. Fl. Bermuda 
279. 1918. Cyclospermum leptophyllum Sprague, Jour. Bot. 61: 131. 
1923. Apium ammi var. genuinum Wolff, in Engl. Pflanzenr. 90 
(IV. 228) : 54. 1927. Apium depressum Jones, Contr. W. Bot. 18: 63. 
1933. 



12 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Plants annual, alternately branching above, 0.5-6 dm. high; 
leaves oblong-ovate, the blades 3.5-10 cm. long, 3.5-8 cm. broad, 
3-4-pinnately decompound, the leaflets linear to filiform, 1.5-7 mm. 
long, 0.5-1 mm. broad; petioles 2.5-11 cm. long, the sheath white- 
scarious-margined; cauline leaves ternate-pinnately decompound, the 
leaflets filiform, 4-35 mm. long; umbels compound, sessile, the rays 
to 2 cm. long; involucre and involucel wanting; pedicels 15-20, 2- 
8 mm. long; calyx teeth inconspicuous; carpophore entire or very 
shortly bifid; fruit ovoid, 1.2-3 mm. long, 1.5-2 mm. broad. 

Illustrations: Jacq. Hort. Vindob. 2: pi. 200. 1773. Mart. Fl. 
Bras. 11 (1): pi. 91. 1879. Urban, Bot. Jahrb. 28: pi 2, f. 35. 1899 
(fruit). Urban & Wolff, Ark. Bot. 22A (10): pi. 1. 1929. 

Amazonas: Pr. Chachapoyas: Hacienda Shani, Cevasco. Ancash: 
Pr. Bolognesi: Quero al este de Huasta, monte rigido, 3800 m., 
Cerrate 2518. Ayacucho: Pr. Andahuaylos: above Chincheros, km. 
203 north of Abancay, 3600 m., Hutchison 1716. Cajamarca: Pr. 
Hualgayoc: Mont. Nancho, 7700 ft., Raimondi. Cusco: Pr. Cusco: 
Laguna Huaypo, entre Cusco y Yucay, 3900 m., Ferreyra 9866; 
Saxihuaman, Mathias 3678. Pr. Calca: alrededores del Calca, 
2800 m., Vargas 4.083. Pr. Paucartambo: laderas de Paucartambo, 
arcillo-pedregoso, 3400 m., Vargas 910. Huancavelica: Pr. Huan- 
cavelica: west of and above Huando, road to Huancavelica, ca. 5 km. 
above bridge over Rio Palca, 3650 m., Hutchison 1677. Huanuco: 
Pr. Tingo Maria: Cordillera Azul below Tea Garden on road to La 
Divisoria, Mathias 3444- Junin: Pr. Tarma: cercanias de Tarma, 
Isern (Cuatrecasas 2552}. La Libertad: Pr. Trujillo: Rio Moche, 
arcillo-arenoso, 30 m., Ridoutt 173; Santa Clara (Valle Chicama), en 
borde de acequia, 55 m., L6pez 187. Lima: Pr. Huarochiri: Matu- 
cana, ditch, 2400 m., Asplund 11099; Matucana, at base of stone 
fence, 8000 ft., Macbride & Feather stone 382. Pr. Lima: Chaclacayo, 
entre Lima y Chosica, 700 m., Ferreyra 11138; Rimac Valley, 700 m., 
Rauh & Hirsch 113; Chinchin (Cheuchin), Ruiz & Pavdn; roadsides, 
waste ground, etc., 500 ft., Macbride & Featherstone 53; Chosica, 
800 m., Soukup 2048; Hacienda Villa, 18 km. sur de Lima, 40-50 m., 
Ferreyra 10466; Lima, Hipodromo San Felipe, 157 m., Ferreyra 11109; 
inmediaciones de la Hacienda Infantas cerca a Lima, Ridoutt. Pr. 
Chancay: Supe, in sandy wash near the sea, 3 m., Eyerdam 9076. 
Puno: Pr. Puno: Cerros de Puno, 3900 m., Ferreyra; Isla de Aman- 
tani, 3900 m., Aguilar; Capachica Peninsula, Lake Titicaca, 12,500 
ft., Tutin 1215. Pr. Huancane": Conima, cerro Calvario, 3900 m., 
Aguilar; Miajachi, camino a Huancane", orillas del rio, 3800-3900 
m., Aguilar 163. 



FLORA OF PERU 13 

A subtropical weed. Mexico, Central America and West Indies 
to Argentina and Chile, sporadic to New York, Oregon and Califor- 
nia. Apium leptophyllum is a wide spread ruderal occurring through- 
out the world in subtropical environments. In marginal habitats or 
in areas of heavy grazing or mowing it may be modified in vegetative 
characters to such an extent that it is difficult to distinguish from 
A. laciniatum. Apium laciniatum is, as far as we know, confined to 
the loma vegetation. The following specimens are referred to A. lep- 
tophyllum on the basis of geographical location since they were not 
collected in the coastal lomas of Peru. However, in vegetative char- 
acters they suggest A. laciniatum. Ancash : Pr. Huaraz : sandy wash, 
3100-3150 m., along streams south of Rio Santa, opposite Huaraz, 
Pennell 15333. Huanuco: Pr. Huanuco: Mito, about 9000 ft., Mac- 
bride & Feather stone 1682. Lima: Pr. Canta: bare rocky slope of 
small canyon, 3000-3200 m., Pennell 14594. Pr. Huarochiri: Matu- 
cana, 8000 ft., Macbride & Featherstone 236. Puno: Pr. Melgar: 
Chuquibambilla, rocky clay soil on puno, 3900-4000 m., Pennell 
13349; 3850-3900 m., Pennell 13405. 

ARRACACIA Bancroft 

Arracacha DC. Bibl. Univ. Sci. & Arts 40: 78. 1829. Velaea DC. 
Coll. Me"m. 5: 61. 1829. Velka D. Dietr. ex Steud. Nom. Bot. ed. 2. 
2: 746. 1841. 

Reference: Constance, Bull. Torrey Club 76: 39-52. 1949. 

Stout or slender, erect, herbaceous or somewhat woody, caules- 
cent, branching or simple, glabrous to pubescent perennials, from 
taproots or tubers. Leaves petiolate, once to several times ternate, 
pinnate, or ternate-pinnate, the leaflets or ultimate divisions various. 
Petioles sheathing. Inflorescence of loose to somewhat compact 
compound umbels; peduncles terminal and lateral, or rarely only 
terminal, occasionally some umbels sessile. Involucre wanting or 
vestigial. Fertile rays few to numerous, spreading-ascending to 
divaricate and reflexed. Involucel of few short to long, narrow 
bractlets, shorter to longer than fruit or wanting. Fertile pedicels 
few, spreading-ascending to reflexed. Flowers white, greenish-yel- 
low, greenish, reddish-brown or maroon, or purple; petals oblanceo- 
late to obovate with a narrower inflexed apex; calyx teeth obsolete; 
styles short to long, erect to spreading or reflexed, the stylopodium 
conic and conspicuous to depressed and indistinct. Carpophore 2- 
parted to the base or only bifid, flat or terete. Fruit lanceolate or 
oblong to ovoid, usually narrowed at apex, compressed laterally, 



14 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

glabrous or pubescent; ribs prominent, acute to obtuse, or filiform 
and indistinct; vittae solitary to several in the intervals, 2-several on 
commissure; seed subterete in transection, often channeled under 
vittae, the face sulcate or concave. 

The genus is well represented by some 25 species in the flora of 
Mexico and Central America. The five endemic Peruvian species 
are the southernmost extension of the genus. One species, A. xan- 
thorrhiza, "arracacha," is commonly cultivated in Latin America. 

Leaflets spinulose-serrate; stylopodium conic; carpophore bifid about 

% its length; fruit tapering at apex A. elata. 

Leaflets variously serrate to incised or lobed, not spinulose-serrate; 
stylopodium depressed; carpophore 2-parted to the base (un- 
known in A. peruviana) ; fruit blunt at apex. 
Rays 4-8 cm. long; bractlets linear, herbaceous, exceeding reddish- 
brown flowers A. peruviana. 

Rays 1-4 cm. long; bractlets broad and scarious, or linear but 

shorter than purple or greenish flowers. 
Bractlets scarious or scarious-margined, lanceolate to obovate. 
Bractlets 4-8, obovate to lanceolate, scarious, 5-10 mm. long, 
exceeding the flowers; ribs of fruit very prominent and 

corky; vittae small, 2-3 in the intervals A. incisa. 

Bractlets 3-6, ovate-acuminate, narrowly scarious-margined, 
2-6 mm. long, shorter than the flowers; ribs of the fruit 
filiform; vittae large, solitary in the intervals. 

A. equator ialis. 
Bractlets herbaceous, linear. 

Plants 5-12 cm. high; fruit (immature) oblong, 10 mm. long, 
2-3 mm. broad, constricted below apex; vittae solitary 

in the intervals A. xanthorrhiza. 

Plants 3-4 dm. (to 1 m.) high; fruit ovoid, 6-7 mm. long, 
4-5 mm. broad, not constricted; vittae 2-3 in the in- 
tervals A. andina. 

Arracacia andina Britt. Bull. Torrey Club 18: 37. 1908. 

This species was described from Bolivian collections. Peruvian 
specimens tentatively referred here by Constance in 1949 (Bull. 
Torrey Club 76: 49) are apparently A. equator ialis. Better and more 
mature material is needed to define these taxa. A fragmentary col- 
lection at Chicago Natural History Museum (no. 870541) labeled in 



FLORA OP PERU 15 

handscript "Cuchero, Peruvia, 1829" (Huanuco) and with a printed 
locality "Brasilien leg. Poeppig" may be referable here. However, 
the material is too immature and too fragmentary for positive iden- 
tification. Furthermore, the mixture of data on the label makes the 
locality suspect. 

Arracacia elata Wolff, Bot. Jahrb. 40: 304. 1908. 

Stout, clambering, branching, up to 4.5 m. high, the foliage essen- 
tially glabrous; leaves ovate, 1-3 dm. long, ternate-l-2-pinnate, the 
leaflets lanceolate to ovate, acute or acuminate, cuneate to rounded 
at base, the lower distinct and short-petiolulate, the upper sessile 
and confluent, 3-6 cm. long, 1-4 cm. broad, sharply spinulose- 
serrate and often incised toward base, the lower surface paler and 
glabrous, strongly reticulate, a squamulose tuft on the upper side of 
the sulcate rachis at the base of the larger leaflets; petioles 15-45 cm. 
long, their lower one- third broadly sheathing; cauline leaves similar, 
the uppermost with petioles wholly sheathing and inflated; inflores- 
cence branching, the peduncles arising axially, 10-25 cm. long, 
squamulose at apex; involucre wanting, or occasionally of a single 
leaf sheath; fertile rays 4-15, slender, spreading-ascending, 3-6 cm. 
long, scaberulous at apex; involucel of 8-10 linear to lanceolate, en- 
tire or few-toothed unequal bractlets 3-15 mm. long, the longer 
exceeding the flowers but shorter than the fruit; fertile pedicels 1-8, 
spreading, 5-8 mm. long; flowers greenish or yellow, the petals oval; 
stylopodium conic, the styles slender, recurved; carpophore bifid 
about one-fourth its length, rigid; fruit ovoid, 5 mm. long, 3 mm. 
broad, glabrous, the ribs prominent, obtuse, with narrow, sharply 
V-shaped intervals; vittae very small and indistinct, forming a con- 
tinuous layer between seed and pericarp; seed deeply channeled under 
the intervals, the face sulcate. F.M. Neg. 3473. 

Amazonas: Pr. Chachapoyas: "Ostl. von Chachapoyas zwischen 
den Tambo Ventillas u. Piscohuanuma," 3300 m., Weberbauer 4423, 
type (entre el tambo Ventillas y Piscohuanuma, camino de Chacha- 
poyas a Moyobamba). Ayacucho: Pr. Huanta: Choimacota Valley, 
3000 m., Weberbauer 7584. Cajamarca : 3500-3800 m., Vargas 10369. 
Cusco: Pr. Urubamba: woods, Veronica, 3000 m., Rauh & Hirsch 
1049. Huanuco: Pr. Pachitea: Tambo de Vaca, 13,000 ft., Macbride 
4456. La Libertad: Pr. Otuzco: Chilte (Hda. Llague'n), en quebrada 
pedregosa, Ldpez 0615. Puno: Pr. Carabaya: entre Ayapata y 
Kahualluyoc, 3595-3800 m., Vargas 10747. 

Known only from Peru. 



16 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Arracacia equatorialis Const. Bull. Torrey Club 76: 46. 1949. 

Slender, branching, 4-8 dm. high, the foliage squamulose; leaves 
triangular-ovate, 6-9 cm. long, biternate or bipinnate, the leaflets 
ovate to lanceolate, acute to acuminate, cuneate to truncate at base, 
the lower distinct and short-petiolulate, the upper sessile and con- 
fluent, 1.5-3 cm. long, 1-2.5 cm. broad, mucronate-serrate and the 
larger incised or lobed, squamulose on sheaths, rachises, veins, and 
margins beneath with linear scales, squamulose or merely scaberu- 
lous above, the lower surface paler and reticulate, a squamulose tuft 
on the upper side of the sulcate rachis at the base of the larger leaflets; 
petioles 10-20 cm. long, sheathing only at base; cauline leaves re- 
duced upward, with linear to lanceolate, elongate divisions, the obo- 
vate petioles wholly sheathing, little inflated; inflorescence branching, 
the peduncles arising in whorls or singly, 2-12 cm. long, the terminal 
sometimes subsessile, squamulose at apex; involucre usually wanting; 
fertile rays 2-6, slender, spreading-ascending, 2-4 cm. long, scaber- 
ulous; involucel of 3-6 ovate-acuminate, entire, subequal, narrowly 
scarious-margined bractlets 2-6 mm. long, shorter than flowers and 
fruit; fertile pedicels 1-3 (-5), stout, ascending, 3-5 mm. long; flowers 
purple, the petals obovate; stylopodium depressed, the styles slender, 
spreading-ascending; carpophore 2-parted to base, lax, filiform; fruit 
ovoid-oblong, 8-9 mm. long, 3-4 mm. broad, glabrous, the ribs fili- 
form, acute; vittae large, solitary in the intervals, 2 on the com- 
missure; seed channeled under the tubes, the face deeply sulcate. 

Callacate, Jelski 354- Apurimac: Pr. Andahuaylas: quebrada 
Posoconi, 2650 m., Vargas 8795. Cusco: San Sebastian, grassy place 
on summit of bluff, 3300-3400 m., Pennell 13628. Pr. Cusco: Cerro 
Sape, frente a Sacsahuaman, cerca a la ciudad de Cusco, 3400 m., 
Ferreyra 2675. Pr. Paruro: Araypallpa, 3100 m., Vargas J^ll. 
Junin: Pr. Tarma: Huasa-huasi, Ruiz & Pav6n; entre Palca y Hua- 
capistana, 2400-2700 m., Weberbauer 1745; Carpapata, 2500 m., 
C err ate 2806; entre Palca y Carpapata, 2500 m., Cerrate 929; que- 
brada pedregosa, 2300-2500 m., L6pez 802; Huacapistana, Valle de 
Tarma, 2400 m., Velarde 722; Chanchamayo, Isern (Cuatrecasas 
2417). Pr. Huancayo: Huancayo, alrededores, Soukup 3579. 

Southern Peru to Ecuador. 

Arracacia incisa Wolff, Bot. Jahrb. 40: 305. 1908. 

Stout, caulescent, branching, 3-12 dm. high, the foliage squam- 
ulose; leaves triangular-ovate to ovate-lanceolate, 1-2.5 dm. long, 
ternate-pinnate or bipinnate, the leaflets triangular-ovate to ovate- 



FLORA OF PERU 17 

oblong, acute, cuneate or truncate at base, the lower distinct and 
short-petiolulate, the upper sessile and the larger pinnately incised, 
squamulose on margins and along veins on both surfaces, the lower 
surface paler and reticulate, a squamulose tuft on the upper side of 
the sulcate rachis at the base of the larger leaflets; petioles 8-16 cm. 
long, narrowly sheathing at base, the sheaths scaberulous on the 
veins; cauline leaves with wholly sheathing, inconspicuously inflated 
petioles; inflorescence branching, the peduncles arising axially and 
terminally, 2-12 cm. long, squamulose at apex; involucre wanting, or 
of 1 or 2 sheathing bracts; fertile rays 4-8, stout, spreading-ascend- 
ing, 1-4 cm. long, scaberulous at least at apex; involucel of 4-8, 
obovate to lanceolate, scarious, denticulate-margined, unequal bract- 
lets, 5-10 mm. long, the green central portion projecting as an 
acuminate point, exceeding flowers but shorter than fruit; fertile 
pedicels 2-6, stout, spreading, usually 2-5 mm. long, scaberulous; 
flowers dark purple or greenish, the petals obovate; stylopodium de- 
pressed, the styles slender, divaricate; carpophore 2-parted to base, 
lax; fruit ovoid, 5-8 mm. long, 3.5-6 mm. broad, the ribs very prom- 
inent and corky, acute; vittae small, 2-3 in the intervals, 3-6 on the 
commissure, frequently some accessory ones under the ribs or in 
the intervals; seed scarcely channeled under the intervals, the face 
deeply sulcate. F.M. Neg. 3472. 

Ancash: Pr. Bolognesi: Capillapunta, Cerro al S. de Chiquian, 
3560 m., Ferreyra 5712, Cerrate 155. Cusco: Pr. Paucartambo: Pau- 
cartambo Valley, Hacienda Churu, 3500 m., Herrera 1391; Ken- 
cumayo, 3300 m., Woytkowski 199. Pr. Cusco: Cusco, Rose & Rose 
19034. Huanuco: Pr. Pachitea: steep rocky open grassy slope, Hua- 
cachi, estacion near Muna, 6500 ft., Macbride 4163. Lima: Pr. 
Huarochiri: "in declivibus rupestribus prope Tambo, ad viam fer- 
ream inter oppida Lima et Oroya," 2650 m., Weberbauer 165, type; 
Viso, sandy hillside, 2800 m., Goodspeed, Stork & Norton 11540; 
valley of Rio Rimac near Lima-Oroya highway at km. 90 east of 
Lima, 3000 m., Goodspeed & Weberbauer 33059; Matucana, 8000 ft., 
Macbride & Featherstone 326, 2949; Rio Blanco, 12,000 ft., Macbride 
& Featherstone 730. Pr. Canta: cerca a Culluay entre Canta y La 
Viuda, 3600-3700 m., Ferreyra 12964 Pasco: Pr. D. Carrion: in 
shrub on southwestern canyon slope, Yanahuanca, 10,000 ft., Mac- 
bride & Featherstone 1244- 

Known only from Peru. 

This species and A. peruviana have been generally confused, 
largely because of the inadequacy of the original description. The 



18 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

involucels of the two species are entirely distinct. Arracacia incisa, 
with its conspicuous scarious involucels, deep purple flowers, and 
blunt, prominently ribbed fruit, is one of the most distinctive species 
of the genus. The taproot is fleshy and has a fragrance of anise. 

Arracacia peruviana (Wolff) Const. Bull. Torrey Club 76: 45. 
1949. Velaea peruviana Wolff, Bot. Jahrb. 40: 303. 1908. 

Slender, branching, 6-9 dm. high, squamulose to scaberulous 
throughout, the stem base clothed with dry sheaths, from a branched 
taproot; leaves ovate-lanceolate, 2-3 dm. long, bipinnate, the leaflets 
ovate to lanceolate, acute, cuneate at base, the lower distinct and 
short-petiolulate, the terminal sessile and confluent, 2-5 cm. long, 
1-4 cm. broad, coarsely sinuately lobed and mucronulate-serrate, 
squamulose on veins and margins, the lower surface paler and retic- 
ulate; petioles 1-3 dm. long, sheathing below; cauline leaves pinnate, 
the uppermost with short, wholly sheathing petioles; inflorescence of 
alternate axillary peduncles, 7-15 cm. long, squamulose at apex; in- 
volucre wanting, or of a single leaf sheath; fertile rays 5-10, slender, 
spreading, 4-8 mm. long, squamulose especially at apex; involucel 
of 6-10 entire linear bractlets 5-9 mm. long, exceeding flowers but 
shorter than fruit; fertile pedicels 2-6, spreading, 5-6 mm. long, 
squamulose or scaberulous above; flowers reddish-brown, the petals 
obovate; stylopodium depressed, the styles slender, spreading-erect; 
carpophore unknown; fruit ovoid, 4-6 mm. long, 3-4 mm. broad, 
glabrous, the ribs filiform; vittae large, solitary in the intervals, 2 on 
the commissure; seed face deeply and narrowly sulcate. F.M. Neg. 
28335. 

Ancash: Pr. Cajatambo: infra Ocros, 3000-3200 m., Weberbauer 
2748, type. Ayacucho: Pr. Huanta: mountains northeast of Huanta, 
Weberbauer 7513. Lima: Pr. Yauyos: Cuchapaya-pampa, cerca (ar- 
riba) a Tupe, 2830 m., Cerrate 1027. Moquegua: Pr. Mariscal Nieto: 
Carumas, Weberbauer 7269. 

"Rainy-green formation," moist shrub areas, Peru, 2800 to 3200 
meters. 

Arracacia xanthorrhiza Bancroft, Trans. Agr. Hort. Soc. Ja- 
maica 1825 : 5. 1825. Conium arracacha Hook. Exot. Fl. pi. 152. 1825. 
Arracacha esculenta DC. Bibl. Univ. Sci. & Arts 40: 78. 1829. Ban- 
croftia xanthorrhiza Billb. Linn. Samf. Handl. 1: 40. 1833. 

Stout, caulescent, branching, 5-12 dm. high, glaucous, the foliage 
squamulose and scaberulous; leaves broadly ovate, 10-35 cm. long 



FLORA OF PERU 19 

and broad, biternate or bipinnate, the leaflets ovate-lanceolate to 
triangular-ovate, acuminate, cuneate to rounded at base, the lower 
distinct and often short-petiolulate, the upper sessile and confluent, 
4-12 cm. long, 1.5-6.5 cm. broad, coarsely simply or doubly mucro- 
nate-serrate and incised or lobed, squamulose or scaberulous on ra- 
chises, veins, and margins with flattened oblong or linear scales, the 
lower surface pale and reticulate, a squamulose tuft on the upper 
side of the sulcate rachis at the base of the larger leaflets; petioles 
0.8-4.5 dm. long, sheathing only at base; cauline leaves reduced up- 
ward, mostly ternate or 3-parted, with lanceolate, acuminate divi- 
sions, the lower alternate and petiolate, the upper often opposite and 
wholly sheathing with narrow, scarcely inflated sheaths; inflores- 
cence branching, the peduncles arising in whorls or singly, 3-10 cm. 
long, squamulose or scaberulous at apex; involucre wanting; fertile 
rays 5-12, slender, spreading-ascending, 1.5^4 cm. long, scaberulous; 
involucel of 5-8 linear, entire, unequal, herbaceous bractlets 2-5 mm. 
long, shorter than flowers and fruit; fertile pedicels 3-8, slender, 
spreading-ascending, 2-4 mm. long; flowers purple or greenish, the 
petals oval; stylopodium depressed, the styles slender, ascending; 
carpophore 2-parted to base; immature fruit oblong, 10 mm. long, 
2-3 mm. broad, constricted below the apex, glabrous, the ribs prom- 
inent, acute; vittae rather large, solitary in the intervals, 2 on the 
commissure; seed channeled under the vittae, the face deeply sulcate. 

Illustrations: Bot. Mag. 58: pi. 3092. 1831. DC. Mem. Soc. Phys. 
G&ieve 6 : pi 1. 1833. 

Cusco: Pr. Cusco: Colimas del Saxaihuaman, 3600 m., Herrera 
858. Lima: Pr. Huarochiri: Altura de Surco, 3000-3200 m., Fer- 
reyra 679. 

This is the commonly cultivated arracacha of northern South 
America. Little is known of its domestication and its origin in the 
wild. An account of the cultivation of the species is given by W. H. 
Hodge, The Edible Arracacha, a little-known root crop of the Andes, 
Economic Botany 8: 195-221. 1954. 

AZORELLA Lam. 

Fragosa Ruiz & Pav. Prodr. Fl. Peruv. 43: pi. 34. 1793. Chamitis 
Banks ex Gaertn. Fruct. 1: 94, pi. 22, f. 4. 1788. 

Low and spreading, usually cushion-forming glabrous or pubes- 
cent perennial herbs. Leaves petiolate, membranaceous to fleshy or 
coriaceous, simple and entire to 5-7-lobed. Petioles sheathing and 



20 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

persistent on the branches. Inflorescence of simple umbels; pedun- 
cles terminal, or some or all umbels sessile. Involucre present, usually 
hidden by the terminal leaves. Pedicels spreading-ascending or erect, 
often elongate. Flowers greenish-white to yellow; petals oval, with- 
out a narrower inflexed apex; calyx teeth prominent to obsolete; 
styles short, persistent or not, the stylopodium conical to depressed. 
Carpophore lacking. Fruit ovoid to orbicular, terete to somewhat 
compressed dorsally, glabrous; ribs prominent to obscure; vittae 
minute, solitary under the ribs, a woody endocarp surrounding the 
seed cavity; seed sub terete to flattened dorsally in transection, the 
face plane. 

A characteristic genus of cushion plants of the high Andes and 
lower latitudes of South America, commonly known as llareta, or 
yareta, and used as fuel. 

Plants forming hard, woody cushions 1 m. or more high; umbels 

1-5-flowered. 

Leaves entire; fruit orbicular, about 4.5 mm. long. . .A. compacta. 
Leaves lobed; fruit ovate, about 2.5 mm. long. 

Leaves shortly 3-lobed, the lobes subequal A. corymbosa. 

Leaves shortly 5-lobed, the median lobe elongate. .A. pulvinata. 

Plants forming low mats or cushions to about 10 cm. high; umbels 
mostly more than 5-flowered. 

Leaf blades linear, mostly entire A. diapensioides. 

Leaf blades neither linear nor entire, crenate, toothed, or deeply 
lobed. 

Petioles narrowed above the sheath, leaf blades with mostly 
obtuse lobes or crenations. 

Leaf blades cuneate and decurrent, the margins usually shal- 
lowly bilobed at apex A. biloba. 

Leaf blades truncate to cordate, the margins crenate to cre- 
nately lobed A. crenata. 

Petioles pulvinate, leaf blades deeply 3-7-incised, the lobes 
spinulose A. muUifida. 

Azorella biloba (Schlecht.) Wedd. Chlor. And. 2: 195, pi 66 B. 
1860. Fragosa biloba Schlecht. Linnaea 28: 478. 1856. Azorella bi- 
loba ft strigosa Wedd. Chlor. And. 2: 195. 1860. ?A. biloba a normalis 
f . subglabra Kuntze, Rev. Gen. 3 (2) : 111. 1898. 



FLORA OF PERU 21 

Plants cespitose, 1-5 cm. high, from a long, stout taproot, acau- 
lescent, the leaves clustered at the apex of the rootstock; leaves 
petiolate, the petioles to 2.5 cm. long, sparsely strigose-setose, fibril- 
lose and membranous, dilated below; leaf blades oblong, cuneate and 
decurrent, 0.5-25 mm. long, 3-10 mm. broad, palmately nerved, 
shallowly to deeply 2-lobed at the apex, rarely entire or inconspic- 
uously 3-lobed, the lobes obtuse, cuneate, glabrous to densely stri- 
gose; inflorescence sessile or short-pedunculate, mostly shorter than 
the leaves, the peduncles more or less strigose-pilose, the hairs brown- 
ish-white; involucral bracts several, shorter than the flowers, linear- 
lanceolate, obtuse, ciliate-strigose especially on the margins; umbels 
about 10-flowered, the flowers greenish-white or yellowish; pedicels 
to 10 mm. long, glabrous; fruit ovate, about 2.5 mm. long, about 
2 mm. broad, glabrous, the 3 dorsal ribs conspicuous; calyx teeth 
prominent, deltoid; stylopodium depressed-conical, the styles per- 
sistent. 

Illustrations: Wedd. Chlor. And. 2: pi. 66 B. 1860. Bull. Inter- 
nat. Ceska Akad. FT. Jos. 13: pi. 3, figs. 4, 5. 1909. 

Cusco: Pr. Calca: paso de Amparacs, en rocas, 4500 m., Vargas 
4044- Pr. Quispicanchis: Cordillera Ausangate, 4500 m., Rauh & 
Hirsch 1189. Pr. Canchis: Uyumiri near Sicuani, 14,000 ft., Stafford 
965. Pr. Cusco: Cord, de Cusco, Gay. Junin: Acopalca-nevado, 
plus 4000 m., Soukup 3704. Puno: Pr. Carabaya: bajando abra de 
Antapampa, laderas, 4300 m., Vargas 6838; 4150 m., Vargas 6848. 
Carabaya, Lechler, type. Pr. Huancane": in pampa, Moho, 3125 m., 
Shepard 95. 

High Andes to Ecuador, Bolivia, and Argentina. 

Azorella compacta Phil. Anal. Mus. Nac. Chile 8: 28. 1891. 
Laretia compacta Reiche, Anal. Univ. Chile 104: 784. 1899; Reiche, 
Fl. Chile 3: 63. 1902. Azorella prismatoclada Domin, Repert. Sp. 
Nov. 4: 297. 1907. A. columnaris Wolff, Bot. Jahrb. 40: 288. 1908. 
A. yareta Hauman, Physis 4: 485, /. 3d, 4c, 5, 6. 1919. 

Plants forming hard, dense cushions 1 m. or more high with 
branching, woody, angular stems, clothed with old leaves; leaves 
rosulate, sessile, thick, carinate, ovate to oblong, 3-6 mm. long, 
1-4 mm. broad, expanded toward base, mucronate or obtuse, gla- 
brous, the leaves on peripheral branches larger and sometimes hori- 
zontally oriented; umbels sessile to short-pedunculate, 4-5-flowered; 
pedicels 1-2 mm. long; fruit orbicular, about 4.5 mm. long and 
broad, glabrous; calyx teeth persistent. F.M. Neg. 28319. 



22 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Illustrations: Bull. Internat. Ceska Akad. FT. Jos. 13: pi 3,f. 13; 
14: pi. 4, f. 4, 5. 1909. Drude in Engl. & Prantl, Pflanzenf. 3 (8): 
131, /. 52. 1897. Reiche, Bot. Jahrb. 28: pi. 1, f. 16. 1899 (fruit 
section). 

Arequipa: Pr. Arequipa: Solitario, 4400 m., en rocas, Vargas 
13113; cimbral i Salinas 4350 m., arenoso-pedrigosos, Vargas 12695. 
Mount Chacani, near Arequipa, 17,000 ft., Bailey; near Arequipa, 
16,400 ft., Douglass; crevices of rocks and rocky slopes, Nevado de 
Chacani, 4300-4600 m., Pennell 13317; southern slope of Chacani, 
near Arequipa, 15,000-16,000 ft., Bailey. Pr. Cailloma: Vincocaya, 
Rose & Rose 18946; near Crucero Alto, in dense clumps, 2 ft. across, 
15,000 ft., R. S. Williams 2504. Moquegua: Pr. Mariscal Nieto: 
Cordillera above Torata, 4300 m., Weberbauer 7477. Puno: Pr. Chu- 
cuito: entre Chilliwa i Mazocruz, Livini, 4100-4600 m., Vargas 13012. 

To Argentina, Bolivia and Chile. 

This species is a source of firewood in the high Andes where it is 
commonly known as yareta. We believe this taxon to be quite dis- 
tinct from the genus Laretia to which it has been referred. In habit 
as well as in other morphological characters A. compacta is closely 
related to A. corymbosa. A discussion of this plant has been pub- 
lished by W. H. Hodge, Yareta, Fuel Umbellifer of the Andean 
puna, Econ. Bot. 14: 113-118. 1960. 

Azorella corymbosa (R. & P.) Pers. Syn. 1: 303. 1805. Fragosa 
corymbosa R. & P. Fl. Peruv. 3: 27, pi. 250, f. a. 1802. 

Plants cespitose forming large hard cushions, branching from 
a woody caudex, the branches clothed with old leaves; leaves rosulate, 
the blades pulvinate, truncate, 3-4 mm. long, about 1.5 mm. broad, 
deeply 3-lobed, the lobes subequal, glabrous on the dorsal surface, 
sparsely fibrillose ventrally, palmately veined, the veins departing 
from the apex of the dilated petiole; petioles 7-8 mm. long, wholly 
pulvinate, fibrillose especially on the margins; umbel sessile, few- 
flowered, the pedicels stout, swollen above, 7-10 mm. long, glabrous; 
fruit ovate, 2-2.5 mm. long, about 2 mm. broad, the ribs incon- 
spicuous; calyx teeth inconspicuous; stylopodium depressed, the 
styles not persistent. F.M. Neg. 18267. 

Huanuco: Ruiz & Pavdn, type. Pr. Pachitea: in mounds on 
rocky outcrops, Tambo de Vaca, 13,000 ft., Macbride 4899. 

Andes of Ecuador and Peru. 



FLORA OF PERU 23 

Azorella crenata (R. & P.) Pers. Syn. 1: 303. 1805. Fragosa 
crenata R. & P. Fl. Peruv. 3: 27, pi 249, f. c. 1802. F. reniformis 
R. & P. I.e. 26, pi. 249, f. b. F. cladorrhiza R. & P. I.e. 27, pi 250, 
f. b. Azorella "cladorhisa" Pers. Syn. 1: 303. 1805. A. reniformis 
Pers. I.e. Bolax dadorrhizus Spreng. in Roem. & Schult. Syst. Veg. 6: 
362. 1820. B. crenatus Spreng. I.e. ? Fragosa glabra Lag. Amen. Nat. 
2: 93. 1821. Azorella crenata ft compacta Wedd. Chlor. And. 2: 195. 
1860. A. crenata var. mutisiana Hieron. Bot. Jahrb. 20 : Beibl. 49 : 71. 
1895. A. mutisiana Hieron. ex Wolff, Repert. Sp. Nov. 17: 266. 1921. 

Plants cespitose, 2-12 cm. high, from a long slender branching 
root, acaulescent and cushion-forming; leaves clustered at apex of 
rootstock, petiolate, the petioles 0.3-10 cm. long, densely strigose- 
pilose, fibrillose below; leaf blades obovate to orbicular, 0.4-18 mm. 
long, 4-20 mm. broad, obtuse, the margins entire below, crenate to 
crenately lobed above, the lobes mostly obtuse, subglabrous to hirsute- 
pilose above, more or less strigose beneath with grayish-white hairs, 
the hairs to 4 mm. long, the primary nerves 5-9, palmate; peduncles 
1-3 cm. long, densely strigose-pilose with grayish-white hairs, shorter 
than the leaves; involucral bracts 5-7, equaling or exceeding the 
pedicels, linear-lanceolate, acute, conspicuously pilose especially on 
the margins; umbels 5-12-flowered, the flowers greenish-white to 
yellowish; pedicels 2-7 mm. long, glabrous to pilose; fruit sub- 
globose, 2-3 mm. long, about 1.5 mm. broad, glabrous, the dorsal 
ribs conspicuous; calyx teeth prominent; stylopodium conical, the 
styles persistent. F.M. Negs. 18266, 3430, 37074. 

Illustrations: Bull. Internat. Ceska Akad. Fr. Jos. 13: pi 3, f. 2. 
1909. Weberbauer, Vegetat. Erde 12: 209, /. 44- 1911; El Mundo 
Veg. And. Per. 371, /. 26. 1945. 

Ancash: Conchucos, 4500 m., Weberbauer 7240. Cusco: puna, 
Rio Marcapata, 3800 m., Rauh & Hirsch 1266. Pr. Anta: Cordillera 
Salcantay, 4500 m., Rauh & Hirsch 1447. Huancavelica: arriba 
de Laria, Dist. Conico, 3800-3950 m., Tovar 2494. Huanuco: Pr. 
Huamalies: Monzon, Weberbauer 3319; Huamalies, Ruiz & Pav6n, 
type of Fragosa cladorrhiza. Pr. Huanuco: puna de Panao, Asplund 
13716; Mito, 9000 ft., Macbride & Feather stone 1787. Junin: Pr. 
Junin : tundra vegetation from Lago Capillacocha to Lago Aguacocha, 
north of Carhuamayo, 4200-4400 m., Constance & Tovar 3596; 
Pampa de Junin, 26 km. southeast of Junin, open grassy steppe, 
4100 m., Constance & Tovar 3589. Pr. Tarma: punas de Tarma, 
3900 m., Cerrate 981; 3800^000 m., L6pez 833; Tarma, Ruiz & 
Pavdn, type of Fragosa crenata. Pr. Yauli: Mar Tunel, 4000 m., 



24 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Velarde 1960. Pr. Huancayo: quebradas east of Huancayo, 3400 m., 
Stork & Horton 10213. Pasco: Cord. Raura, puna, 4300 m., Rauh 
& Hirsch 1837. Pr. Oxapampa: cumbre entre Capillacocha y Pau- 
cartambo, estepa de gramineas, 4300-4500 m., Ferreyra 5187. 
Piura: Huancabamba (Piura Divide), 10,000 ft., Sandeman 4312. 
Without locality: Dombey; Ruiz; Mutis ex Herb. Humboldt; Ruiz 
& Pavdn. 

Paramos and grass steppes at 3000 to 4500 meters altitude in 
the Andes, from Venezuela to Peru. 

Azorella diapensioides A. Gray, Bot. Wilkes Exped. 1: 702. 
1854. A. diapensioides glabra Wedd. Chlor. And. 2: 190. 1860. 
?A. diapensioides a denticulata Wedd. I.e. A. glabra Wedd. I.e. 
pi. 67 A. 1860. ? A. lehmanii Hieron. Bot. Jahrb. 20: Beibl. 49: 
71. 1895. 

Plants cespitose, forming hard, dense, somewhat convex bright 
green to blue-green cushions to 10 cm. high and 50 cm. across, 
branching from a woody caudex, the branches clothed with per- 
sistent leaf sheaths; leaves rosulate, the leaf blades linear, 4-10 
mm. long, 1.5-2 mm. broad, obtuse, apiculate to spinulose-tipped, 
decurrent, entire or the outermost rarely 1-3-lobed, the lateral lobes 
short, glabrous to sparingly strigose; petioles to 10 mm. long, pul- 
vinate below, narrowed to the blade, sparingly fibrillose on the 
margins; umbels sessile, several-flowered, the flowers pale yellow 
to greenish; pedicels stout, to 4 mm. long, glabrous; fruit 2.5 mm. 
long, about 2 mm. broad, the ribs inconspicuous; calyx teeth in- 
conspicuous; stylopodium depressed, the styles not persistent. 

Illustrations: Engl. & Prantl, Pflanzenf. 3 (8): 127, /. 50. 1897. 
Schimper, Pflanzen-Geogr. 782, f. 441 (1 & 2}. 1898. Jour. N. Y. 
Bot. Card. 33: 55, /. 1, 2. 1932. 

Apurimac: Soroccha Pass on trail Chincheros-Andahuaylas, 
open puna and among rocks, 4300 m., West 3740. Arequipa: Pr. 
Arequipa: hacia el Solitario, 4400 m., pedregoso, Vargas 13114- 
Cusco: Vargas 5592. Pr. Espinar: alrededores de Yauri, 3900 m., 
Vargas 12077. Huancavelica: Pr. Huancavelica: puna, Huancavel- 
ica, 3700 m., Rauh & Hirsch 360a. Junin: Pr. Junin: tundra vege- 
tation from Lago Capillacocha to Lago Aguacocha, north of Car- 
huamayo, 4200-4400 m., Constance & Tovar 3593, 3595; Hondores, 
southeast of Laguna Junin, between Junin and San Bias, open 
grass steppe, 4200 m., Constance & Tovar 3592; Pampa de Junin, 
26 km. southeast of Junin, open grass steppe, 4100 m., Constance 



FLORA OF PERU 25 

& Tovar 3590. Pr. Yauli: 25 km. northeast of La Oroya toward 
Tarma, bare open grass steppe, 4000 m., Constance & Tovar 3585; 
Ticlio, stony slope, 4900 m., Asplund 11656; Paso de Anticona, 
4500 m., Cerrate 992; Mar Tunel, 4000 m., Velarde 1952. Lima: 
high Andes, Casa Caucha, Wilkes Expedition. Casapalca, 15,500 
ft. in mats on slopes of alpine basin, Macbride & Featherstone 847. 
Pasco: Cord. Raura, puna, 4300 m., Rauh & Hirsch 1845. Pr. 
Pasco: Cerro de Pasco, 4400 m., Vargas 6596; Huaron, naked earth, 
4200 m., Asplund 11804. Puno: Pr. Chucuita: hacia Chilliwa, 4100 
m., Vargas 13010. Pr. Puno: near Puno, 4000 m., Soukup 210, 214. 
Pr. Lampa: Santa Lucia, Stordy; crece terrenes mui humedos i 
resiste hasta 5000 m. altura, Stordy 32. Tacna: Pr. Tacna: Tacora, 
Weddell (type of A. diapensioides a denticulata). Without definite 
locality: Andes, McLean. Alpamarca, Andes, Crooke. 

Common in grass steppes or puna and also in crevices of rocks, 
forming dense cushions, or hard, thin, somewhat convex mats. Lo- 
cally used as fuel and known as cuncuma, estrella chica, yareta and 
llareta. High Andes to Bolivia. 

Azorella multifida (R. & P.) Pers. Syn. 1: 303. 1805. Fragosa 
multifida R. & P. Fl. Peruv. 3 : 27, pi. 249, f. a. 1802. Bolax multifidus 
Spreng. ex Roem. & Schult. Syst. Veg. 6: 362. 1820. Azorella laxa 
Wolff, Bot. Jahrb. 40: 288. 1908. A. weberbaueri Wolff, I.e. 287. 1908. 

Plants forming small convex, bluish-green, gray to bright green 
mats to 75 cm. across, loosely branching near periphery, the branches 
clothed above with persistent leaf sheaths; leaves rosulate, the 
leaf blades thin, ovate to oblong, 3-13 mm. long, to 10 mm. broad, 
decurrent, 3-7-incised-lobed, the lobes spinulose-tipped, the margins 
entire or serrulate, the median lobe elongate, glabrous dorsally, 
more or less fibrillose above, pinnately veined; petioles pulvinate 
below, to 20 mm. long, fibrillose especially on margins; umbels 
sessile, several-flowered, the flowers greenish-white to yellowish; 
pedicels slender, to 8 mm. long, only slightly enlarged at apex; 
calyx teeth not persistent; fruit ovate, 1.5-2 mm. long, 1-1.5 mm. 
broad; stylopodium low-conical, the styles very short, erect. F.M. 
Neg. 3429. 

Illustrations: Rich. Ann. Ge"n. Sci. Phys. 4: pi. 51, no. 4- 1820. 
Wedd. Chlor. And. 2: pi. 66 C. 1860. 

Ancash: Pr. Bolognesi: Chiquian, Pampa de Lampas, 4100 m., 
Cerrate 1526; Mahuay, 4700 m., Cerrate 2223. Apurimac: Pr. Anda- 



26 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

huaylas: km. 165 carretera entre Andahuaylas y Chincheros, puna, 
3500 m., Ferreyra 2808. Pr. Abancay: entre Urahuasi y Abancay 
(km. 24), 3400 m., Ferreyra 2786; km. 45 east of Abancay, at Say- 
huite, 3600 m., Hutchison 1744; Ampay, 3200-3800 m., Vargas 8406. 
Ayacucho: Pr. Huananga: 75 mi. south of Ayacucho on road to 
Abancay, beginning descent to Ocros, 4000 m., Hutchison 1707. 
Cajamarca: Pr. Hualgayoc: montanas de Nancho, 10,800 ft., Rai- 
mondi 1750, 1749. Pr. Cutervo: Cutervo, Jelski 1114. Cusco: Pr. 
Cusco: between Cusco y Chinchero, 3800 m., West 7167. Pr. Quis- 
picanchis: Cord. Ausangate, 4800 m., Rauh & Hirsch 1257. Pr. 
Anta: Cordillera Salcantay, 4500 m., Rauh & Hirsch 1415. Pr. 
Urubamba: Cordillera Veronica, puna, 4100 m., Rauh & Hirsch 
998; Pacechac, Urubamba Valley, Hill 186, 185, Mathews 516. 
Huancavelica: arriba de Laria, Dist. Conico, 3000-3950 m., Tovar 
2495. Pr. Tayacaja: Hda. Huari, 3700 m., Velarde 2013. Huanuco: 
Pr. Dos de Mayo: Cordillera Huayhuash, 4300 m., Rauh & Hirsch 
1843; Huayhuash, rocks, 4100 m., Rauh & Hirsch 1888. Pr. Huama- 
lies: in montibus a Monzon, 3400-3500 m., Weberbauer 3341, type 
of Azorella laxa. Junin: Pr. Junin: tundra vegetation from Lago 
Capillacocha to Lago Aguacocha, north of Carhuamayo, 4200- 
4400 m., Constance & Tovar 3594, 3597; encima de la laguna de 
Capillacocha, cerca a Carhuamayo, 4300 m., Ferreyra 3926; tundra 
vegetation above Lago Aguacocha, north of Carhuamayo, 4400 m. 
Constance & Tovar 3598; Hondores, southeast of Laguna Junin be- 
tween Junin and San Bias, 4200 m., Constance & Tovar 3591. Pr. 
Tarma: entre Oroya & Tarma, 4000 m., Velarde 606. Pr. Yauli: 
alrededores de Yauli, 3500-3600 m., Tovar 3024. San Jose*, 13,000 
ft., Macbride & Featherstone 1103. La Libertad: Pr. Bolivar: La- 
guna de los Ichus, Bolivar, 3600 m., L6pez & Sagastegui 3244- 
Lima: Pr. Huarochiri: supra Lima, 4600 m., Weberbauer 5180, type 
of Azorella weberbaueri. Pasco: Pr. Pasco: Cerro de Pasco, calcare- 
ous rock, 4300 m., Asplund 11767; "in frigidissimus Andium inter 
Cerro de Pasco de fodinas Gallquijirca," Poeppig (?); puna, Cord. 
Raura, 4400 m., Rauh & Hirsch 1817. Piura: Pr. Huancabamba: 
above Huancabamba, road to Conchaque, 3100 m., Hutchison 1625. 
Puno: Pr. Carabaya: abra Antapampa-juro-juro, 4300 m., Vargas 
6841. Without definite locality: Dombey, type; Ruiz & Pavon. 

Forming large branched prostrate mats at high elevations in 
the Andes to Colombia and Bolivia. Locally known as pasapamaquin 
and yareta. Frequent in moist locations on grass steppes or puna 
or among rocks. 



FLORA OF PERU 27 

Azorella pulvinata Wedd. Chlor. And. 2: 194, pi. 66 A. 1860. 

Plants forming great cushions, several meters broad, branching 
from a woody caudex, the branches clothed with persistent leaf 
sheaths; leaves rosulate, the leaf blades about 3 mm. long, shortly 
5-lobed, the median lobe longer than the lateral, glabrous dorsally, 
the ventral surface densely clothed with long white hairs giving 
the upper surface of the cushion a frosted appearance, palmately 
veined, the veins departing from the apex of the dilated petiole; 
petioles wholly pulvinate, about 7 mm. long, fibrillose on margins; 
umbels sessile, ?l-flowered; pedicels stout, to 6 mm. long; fruit 
oblong, about 2.5 mm. long, about 1.5 mm. broad, the ribs in- 
conspicuous; calyx teeth inconspicuous; stylopodium depressed- 
conical, the styles short. 

Illustrations: Bull. Internat. Ceska Akad. Fr. Jos. 13: pi. 3, 
f. 11. 1909. Weberbauer, Vegetat. Erde 12: 200, /. 30. 1911. El 
Mundo Veg. And. Per. 378, /. 55. 1945. Math. & Const. Bull. 
Torrey Club84:/. 1. 1957. 

Ancash: Pr. Pallasca: Cordillera of Palagatos, 4450 m., Weber- 
bauer 7242. Cusco: Pr. Anta: Salcantay, 4700 m., Rank & Hirsch 
1483. Huanuco: Pr. Dos de Mayo: Huayhuash, 4900 m., rocks, 
Rauh & Hirsch 1887. 

High Andes to Bolivia, at the upper limits of vegetation. 

BOWLESIA R. & P. 

Erect, prostrate or vining, slender, herbaceous, caulescent, 
branching, stellate-pubescent and/or stellate-glochidiate to glabrate, 
annuals or perennials, from slender taproots. Leaves petiolate to 
subsessile above, stipulate, alternate or opposite, membranaceous, 
simple, palmately veined and palmately lobed or divided. Petioles 
not sheathing, the stipules scarious, lacerate. Inflorescence of simple, 
few-flowered, subcompact umbels. Peduncles slender, axillary and 
terminal. Involucre of a few subulate, lacerate bracts or obsolete. 
Pedicels very short or obsolete. Flowers white, greenish-yellow, 
pinkish, or purplish; petals oblong to ovate, obtuse; calyx teeth 
prominent, ciliate, or obsolete; styles short, equaling to slightly 
exceeding the low-conic stylopodium. Carpophore lacking. Fruit 
broadly ovoid to narrowly oblong, constricted at the commissure 
and flattened dorsally or terete, the carpels plane or concave dorsally, 
the fruit thus appearing 2- or 4-winged, glabrate to stellate-pubescent 



28 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

or glochidiate; ribs obsolete; vittae obsolete; seed flattened dorsally 
in transection, the face plane or convex. 

A genus of about 12-15 species, widely distributed throughout 
temperate South America, extending to Mexico and the south- 
western United States. 

Calyx and involucre evident; ovaries, fruits, and stems stellate- 
pubescent, soft to the touch. 
Upper umbels, or all of them, subsessile; fruit globose, inflated 

and becoming indented dorsally, unwinged B. incana. 

Umbels usually all pedunculate; fruit ovoid or ovate, strongly 
flattened and either plane or concave dorsally, thus appearing 
2- or 4-winged. 

Ovaries and lower leaf surfaces densely stellate-pannose or 
-tomentose at least when young; lobes of basal and lower 
cauline leaves usually subequal, broadest near middle; petals 
usually conspicuously stellate-pubescent dorsally. 

B. tropaeolifolia. 

Ovaries and lower leaf surfaces stellate-hirsutulous (or the former 
rarely glabrous) ; central leaf lobe usually markedly longer 
than the others, the lobes all broadest at base; petals 

usually glabrous B. lobata. 

Calyx and involucre obsolete or very inconspicuous; ovaries and 
fruits armed with glochids; stems beset with hard stellate 
processes, harsh to the touch. 

Fruit glochids very slender, "asteroid," with 4-6 very short sub- 
rotate rays B. flabilis. 

Fruit glochids slender, "coprinoid," with 8-12 slender, closely 

reflexed rays. 
Leaves divided to base, the leaflets pinnately parted. 

B. palmata. 

Leaves usually not lobed more than halfway to base, the leaf- 
lets few-toothed to entire. 

Stems scabrous with mostly 8-rayed processes; fruit dark 
brown, oblong-ovoid, 3-4 mm. long, 1.5-2 mm. broad. 

B. setigera. 

Stems scabrous with mostly 4-rayed processes, to glabrous; 

fruit light brown, ovoid, 2-3 mm. long, 1-2 mm. broad. 

Stems rather stout, armed with usually 4-rayed processes; 

both carpels equally armed with glochids 0.5-1 mm. 

long B. sodiroana. 



FLORA OF PERU 29 

Stems filiform, subglabrate; one of each pair of carpels 
subglabrate, the other with glochids less than 0.5 mm. 
long B. tenella. 

Bowlesia flabilis Macbr. Field Mus. Bot. 11: 32. 1931. Drusa 
acutangula Drude in Engl. & Prantl, Pflanzenf. 3 (8): 126. 1897. 
Bowlesia palmata var. acutangula Kuntze, Rev. Gen. 3 (2) : 112. 1898, 
not B. acutangula Benth. 1856. 

Annual, prostrate to ascending and scandent, 1.5-10 dm. long, 
dichotomously branched from a slender taproot, the stems sparsely 
to densely scabrous with sessile or short-stalked, 4-rayed stellate 
processes, the young foliage and inflorescence stellate-hirsute; lower 
leaves alternate, the upper alternate or opposite, petiolate, thinly 
membranaceous or occasionally somewhat thicker, the basal and 
lower cauline orbicular-reniform, 1-6 cm. long, 1.2-7.5 cm. broad, 
cordate to truncate at base, palmately 3-, 5-, or 7-lobed about to 
middle, with narrow to broad sinuses, the lobes lanceolate to ovate, 
acute, subequal or the terminal larger, entire or the larger with 1 
or 2 lateral teeth or lobes, strigose or strigulose above at least 
on veins, with simple hairs, hirsutulous beneath with sessile or 
short-stalked, 4-rayed (cruciate) stellate hairs, bristly on veins and 
at summit of petiole, the petioles slender, 1.5-7.5 cm. long, somewhat 
bristly, dilated at base into an oval, scarious-margined sheath bearing 
3-6 pairs of linear to linear-lanceolate, hyaline, acuminate, stipular 
processes 1-3 mm. long; upper cauline leaves reduced, narrower, 
truncate at base, usually 3-lobed or entire; peduncles obsolete, the 
umbels usually 1-4-flowered, subsessile in leaf axils of main stem 
or on slender axillary branches; involucre apparently lacking, but 
stipules and pedicel hairs often forming a concealing semi-involucre; 
flowers greenish-white; calyx obsolete; petals oval to ovate, 0.4-0.6 
mm. long, plane, glabrous, subacute; ovaries glochidiate-roughened; 
pedicels up to 1 mm. long, densely white-hirsute, spreading; fruit 
broadly ovate to ovate-oblong, 2.5-4 mm. long, 1.8-2.5 mm. broad, 
4-winged, narrowed at apex, a little cordate at base, the carpels 
strongly flattened dorsally, concave dorsally, the commissural face 
convex, usually sparsely stellate with sessile or subsessile, mostly 
4-rayed hairs on the dorsal face and with or without 2 rows of hairs 
on the commissural face, the margins and dorsal ribs conspicuously 
armed with a border of prominently stalked, very shortly 4-6-rayed 
"asteroid" glochids, the rays spreading-reflexed, the stalks unequal, 
up to 2 mm. long. 



30 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Illustration: PeYez-Moreau, Physis 12: 86, /. 1. 1936 (as Drusa 
acutangula). 

Cusco: Pr. Calca: Pisac, ca. 3200 m., Velarde 1343. Pr. Uru- 
bamba: Ollantaytambo, ca. 3000 m., Cook & Gilbert 367a, 872, 1201. 
Huancavelica: Pr. Huancavelica: Peccsuca, 5 km. east of Conaica, 
Tovar 277. Huanuco: Pr. Huanuco: wood-clearing, Mito, Macbride 
& Feather stone 1718, type. Junin: Pr. Huancayo: Huancayo, Soukup 
1987. 

Rocky places in the Andes, from Ecuador to Bolivia and northern 
Argentina, at elevations of 2200 to 3480 meters; also in Mexico. 
Locally known as opuisoro and uphuisuru and considered a styptic 
to stop flow of blood; also used for the lungs. 

Bowlesia incana R. & P. Fl. Peruv. 3: 28, pi. 268, f. a. 1802. 
B. tenera Spreng. Syst. 1: 880. 1825. B. geraniifolia Cham. & 
Schlecht. Linnaea 1: 382. 1826. B. nodiflora Presl ex DC. Prodr. 4: 
75. 1830. B. rotundifolia Phil. Linnaea 28: 651. 1856. B. incana f. 
tenera Urban, Mart. Fl. Bras. 11 (1) : 292, /. 7811. 1879. B. incana 
f. crassifolia Urban, I.e. B. brevipes Phil, ex Reiche, Fl. Chile 3: 59. 
1902, nomen in synon. B. septentrionalis Coult. & Rose, Contr. U. S. 
Nat. Herb. 7: 31. 1900. B. rubra Larranga, Escritos Larranga 2: 112. 
1923. B. asiatica Nasir, Madrono 12: 217, figs. 1-11. 1954. 

Illustrations: Spreng. Sp. Umbell. pi. 5, f. 10. 1818. Engl. & 
Prantl, Pflanzenf . 3 (8) : 125, /. 49A-F. 1897. Cabrera, Man. Fl. 
Buenos Aires 347, /. 128E-G. 1958. 

This species was described by Ruiz and Pavon as "habitat in 
Peruviae collibus ad Huanuci, Rondos et Pillao tractus." The only 
collection examined which purports to be from Peru is the ostensible 
isotype: Dombey, "In Peruvia," at Chicago Natural History Mu- 
seum, and also F.M. Neg. 3410 of Berlin isotype. To our knowledge 
the species has not been collected in Peru since, and it is presumed 
that the original collections were actually obtained in Chile. Bow- 
lesia incana is weedy and has become widely distributed as a weed 
but, curiously enough, not in Peru. 

Bowlesia lobata R. & P. Fl. Peruv. 3: 28, pi. 251, f. b. 1802. 
?B. diversifolia Meyen, Reise, 1 : 447. 1834, nomen. B. palmata var. 
/3 A. Gray, Bot. Wilkes Exped. 1: 696. 1854, in part. B. acutangula 
Benth. PI. Hartw. 186. 1856. B. palmata /3 acutangula Kuntze, Rev. 
Gen. 3 (2) : 112. 1898. Drusa acutangula Drude in Engl. & Prantl, 
Pflanzenf. 3 (8): 126. 1897, as to name only. B. acutiloba Wolff, 
Bot. Jahrb. 40: 284. 1908. 



FLORA OP PERU 31 

Perennial, vining, 3-6 dm. high or long, dichotomously branched 
from a multicipital caudex terminating in a slender taproot, the 
stems sparsely stellate with 4-5 (-8) -rayed sessile or stalked stellate 
hairs, or glabrate, the young foliage and inflorescence stellate- 
hirsutulous; leaves mainly opposite, weakly clustered toward base, 
petiolate, thinly membranaceous, the basal and lower cauline orbic- 
ular-reniform to broadly ovate, 1-7 cm. long, 1-7.5 cm. broad, 
strongly cordate and usually with a narrow sinus, palmately (3-) 
5-7 (-9) -lobed about to the middle, the lobes oval or ovate-triangular 
to lanceolate, obtuse to acute or acuminate, broadest at base, the 
terminal usually longer than the others, all entire to shallowly trilo- 
bulate, weakly bristly-mucronulate, strigose or strigulose above with 
mostly simple or 2-3-forked hairs, thinly hirsutulous beneath with 
short-stalked or sessile 4-8-rayed stellate hairs, the petiole slender, 
2-12 cm. long, dilated into an oval sheath, its margin bearing 1-5 
pairs of linear to lanceolate, hyaline, acute or acuminate, fimbriate 
stipular processes 1-3 mm. long; upper cauline leaves, or all but the 
lowermost, reduced and shorter-petiolate upward, entire or 3-lobed 
and truncate or rounded; inflorescence of solitary or paired axillary 
peduncles bearing simple umbels, the umbels 1-3 (-5) -flowered; 
peduncles weak, short to very slender, 0.5-6 cm. long; involucre 
of several stipule-like hyaline linear or lanceolate bracts; flowers 
white or rose- or purple-tinged; calyx lobes linear to linear-lanceolate, 
hyaline, 0.5-1 mm. long, somewhat persistent, reflexed, shorter than 
petals; petals oblong-oval to oval, 1-2 mm. long, plane, glabrous 
or sparsely stellate-pubescent dorsally, a little narrowed at either 
end; ovaries usually stellate-pubescent, less commonly glabrous; 
pedicels 0.5-2.5 mm. long, or abortive, spreading or recurved; fruit 
ovoid or ovoid-triangular, 1.5-3 mm. long, 1.5-2 mm. broad, 2- 
or 4-winged, narrowed at apex, a little cordate, the carpels strongly 
flattened dorsally, plane or concave dorsally, the commissural face 
lightly convex, sparsely stellate-pubescent on both surfaces with 
sessile, 4-8-rayed hairs, or glabrous. F.M. Negs. 3412, 31815. 

"In peruvia ad Chancay, etc." Ruiz; "in peruviae Andium al- 
pibus versus Pillao vicum, in locis apricis et fruticetis," Ruiz, type. 
Cusco: Pr. Paucartambo: Llulluchayocc, 3750 m., Vargas 1+298. 
Cusco-Puno: Pucara, Weberbauer 453. Lima: Pr. Huarochiri: be- 
tween Rio Blanco and Chicla, ca. 3600 m., Asplund 11306; Rio 
Blanco, 3000-3500 m., Kittip & Smith 21605. Pr. Yauyos: Capia, 
abajo de Tupe, 2800 m., Cerrate & Tovar 1085. Pr. Canta: Obrajillo, 
Wilkes Expedition. 



32 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Andes of Ecuador, central and southern Peru, Bolivia and north- 
western Argentina, at elevations of 1600 to 4600 meters. 

Bowlesia palmata R. & P. Fl. Peruv. 3: 28, pi. 251, f. a. 1802. 
B. palmata a lacerata Kuntze, Rev. Gen. 3 (2) : 112. 1898. 

Annual, erect, prostrate or climbing, 3-12 dm. long, dichoto- 
mously branched from a slender taproot, the stems densely scabrous 
with short-stalked or sessile, 4-rayed stellate processes, the young 
foliage and inflorescence densely hirsutulous with stellate and sim- 
ple hairs; lower leaves alternate, the others opposite, petiolate, thinly 
membranaceous, the basal and cauline ovate to orbicular-reniform, 
1.5-5 cm. long, 2-7 cm. broad, cordate to truncate, palmately 3- or 
5-parted nearly to base, with rather narrow sinuses, the lobes lan- 
ceolate to ovate, subequal, entire to laciniately dissected, acute or 
acuminate, bristly mucronate, strigose above with mostly simple 
large trichomes, or a few smaller 2-4-forked hairs intermixed, hir- 
sutulous beneath with sessile or subsessile, 4-6-rayed stellate hairs, 
the petioles slender, slightly dilated at base into a margin bearing 
3-8 pairs of linear-filiform, hyaline stipular processes 1-^4 mm. long; 
the uppermost cauline leaves reduced, narrower and shorter-petiolate, 
usually 3-lobed; peduncles usually abortive, or some rarely elongate 
(up to 3.5 cm. long!), the 1-4-flowered umbels sessile or subsessile in 
leaf axils; involucre apparently lacking; flowers greenish-yellow or 
yellowish-white; calyx lobes abortive, lanceolate, up to 0.2 mm. long; 
petals oval, 0.5-0.7 mm. long, plane, glabrous, narrowed at apex; 
ovaries sparsely covered with sessile or subsessile stellate hairs, the 
margins and dorsal midrib armed with stalked glochids; pedicels 
abortive, up to 1 mm. long; fruit triangular-ovoid, 2.5-4 mm. long, 
2-3 mm. broad, narrowed at apex, strongly flattened dorsally, 
strongly concave dorsally with inrolled margins, 4-winged, the com- 
missural surface convex, sparsely stellate on both surfaces with 
usually 8-rayed hairs, the margins and dorsal midribs armed with 
very slender, stalked, "coprinoid," 8-12-rayed glochids up to 2 mm. 
long, their rays sharply reflexed. 

Illustrations: Rich. Ann. Ge"n. Sci. Phys. 4: pi. 51, f. 3. 1820. 
Weberbauer, Vegetat. Erde 12: 142, /. 12. 1911; El Mundo Veg. 
And. Per. 231, /. 18. 1945. 

"Peru," Dombey, type. Ancash: Pr. Bolognesi: Conay, 6 km. 
abajo de Chiquian, 2600 m., Ferreyra 7.40-4, Cerrate 600. Pr. Casma: 
Lomas de Lupin, Mathias 3671. Arequipa: Pr. Islay: Mollendo, 
Hitchcock 22390; 10 km. east of Islay, 300 m., Worth & Morrison 



FLORA OF PERU 33 

15718, 15714. Pr. Caraveli: Lomas de Capace, km. 651, several km. 
east of gas station "Call" and up canyon, 250 m., Hutchison 1308. 
La Libertad: Pr. Trujillo: Cerro Campana, 500 m., Ldpez 091 4, que- 
brada bottom, 750 m., Hutchison 1359. Lima: Pr. Lima: Lima, 
Isern 2516; San Geronimo, ca. 500 ft., Macbride 5897; Lomas de 
Amancaes, 420 m., Ferreyra 3962, 400 m., Ferreyra 11825, 250 m., 
Soukup 3121, 200-600 m., Pennell 15927; Atacongo Lomas, 525 m., 
Stork, Horton & Vargas 9285, 300 m., Ferreyra 183; Cerros al norte 
de Chosica, 1800-1900 m., Weberbauer 886; Hacienda Pando, Soukup 
2561; Santa Clara, Rose & Rose 18618, Pr. Chancay: Lomas de 
Pacar (Pativilca), 300 m., Velarde 2322; Lomas de Pativilca, 5 km. 
north of Barranca, 300 m., Stork, Horton & Vargas 9224; Cerro north 
of Barranca, 80 m., Beetle 9088; Lomas de Lachay cerca a Chancay, 
400-560 m., Ferreyra 3873, 8727, 500 m., Velarde 2277, Mathias 3674, 
300-400 m., Cerrate 841, 400 m., Ridoutt. 

Moist slopes of coastal lomas and higher hills, from southern 
Ecuador and throughout western, northern and central Peru (to 
northern Bolivia?), at elevations of 80 to 2600 meters. 

Bowlesia setigera Wolff, Bot. Jahrb. 40: 385. 1908. B. lobata 
var. chiclensis Ball, Jour. Linn. Soc. Bot. 22: 40. 1885, mistakenly 
published as "chilensis"! 

Rather stout annual, decumbent at base and ascending, 1-5 dm. 
long, more or less dichotomously branched from a slender taproot, 
the stems stout, densely to sparsely scabrous with sessile or short- 
stalked, mostly 8-rayed stellate hairs or platelets, the young foliage 
and inflorescence stellate-hirsutulous; lower leaves alternate, the 
upper opposite, petiolate, thinly membranaceous, the basal and lower 
cauline orbicular-reniform, 1-4 cm. long, l;,5Ht cm. broad, narrowly 
cordate, palmately 3-7-lobed to below middle with rather narrow 
sinuses, the lobes narrowly lanceolate to ovate-lanceolate, acute, en- 
tire or with 1 or 2 lateral teeth, strongly bristly mucronate, strigose 
above with forked or few-branched bristles, hirsutulous beneath with 
sessile or stalked 5-8-rayed stellate hairs, often bristly on veins and 
margins, the petioles slender, 1-4 cm. long, stellate-hirsute and 
bristly, dilated into an oval sheath, its margins bearing 3-8 pairs of 
linear, hyaline, acute, entire stipular processes 2-5 mm. long; upper 
cauline leaves like the basal, only the uppermost sometimes reduced, 
subsessile, and trilobed; peduncles very slender (to 5 cm.) to abortive, 
usually paired in middle and upper leaf axils, the umbels 2-5-flowered; 
involucre of an inconspicuous tuft of linear stipule-like bracts, or 



34 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

usually lacking; flowers greenish-white; calyx lobes obsolete; petals 
oval, about 0.8 mm. long, plane, glabrous, a little narrowed at either 
end; ovaries rather densely covered with sessile, mostly 8-rayed stel- 
late hairs and stalked glochids; pedicels up to 1 mm. long, or abortive; 
fruit dark brown, oblong-ovoid, 3-^4 mm. long, 1.5-2 mm. broad, 
more or less 4-angled, a little cordate, the carpels compressed and 
concave dorsally, their margins inrolled, the commissural face con- 
vex, stellate-pubescent on both surfaces with mostly 8-rayed hairs, 
the margins and dorsal midribs densely armed with prominently 
stalked, usually shortly 8-12-rayed "coprinoid" glochids, the stalks 
up to 1 or 1.5 mm. long. F.M. Neg. 3417. 

"Canruru, Andes Peruvie"nne," Savatier. "Peru," Wilkes Expe- 
dition. Ancash: Pr. Cajatambo: prope Ocros, 3200-2400 m., Weber- 
bauer 2659, type. Lima: Pr. Huarochiri: Rio Blanco, 3000-3500 m., 
Killip & Smith 216^1; "ex regione media Andium Peruviae, in con- 
valle fluminis Rimac, 6-11000' s. m.," Ball; "ex saxosis Andium 
Peruviae juxta pagum Chicla, 12-13000' s. m.," Ball, type of B. lo- 
bata var. chiclensis; valley of Rio Rimac, near Lima-Oroya highway 
at km. 70 east of Lima, alt. 1850 m., Goodspeed 33126; Infiernillo, 
entre San Mateo i Rio Blanco, carretera Lima-Huancayo, 3300- 
3350 m., Ferreyra 7006, in part. 

Southern Ecuador to central Peru, at elevations of 1850 to 3500 
meters. 

Bowlesia sodiroana Wolff, Bot. Jahrb. 40: Beih. 91: 49. 1907. 
Bowlesia sp. Muiioz, Sinop. Fl. Chil. pi. 191. 1959. 

Annual, prostrate or ascending or subscandent, 0.5-6 dm. long, 
dichotomously branched from a slender taproot, the stems slender, 
scabrous with short-stalked or sessile, mostly 4- (occasionally 5-, 6-, 
or 8-) rayed stellate scales, the young foliage and inflorescence white- 
hirsute with simple and stellate hairs; lower leaves alternate, the 
upper opposite, petiolate, thinly membranaceous, the basal and lower 
cauline ovate to orbicular-reniform, 1-4 cm. long, 1.5-5.5 cm. broad, 
cordate, palmately 3-, 5-, or 7-lobed very shallowly or to well below 
middle, the lobes ovate to lanceolate, acute to acuminate, subequal 
or the terminal elongate, coarsely toothed or shallowly lobed to en- 
tire, bristly mucronate, strigose above with large, simple or distally 
forked barbellate trichomes and sometimes a few smaller 2-4-forked 
hairs, hirsute or hirsutulous beneath with short-stalked, usually 4- 
(sometimes 6- or 8-) rayed stellate hairs, with some giant trichomes 
on veins, the petioles slender, 1-7 cm. long, bristly at summit, dilated 



FLORA OF PERU 35 

into an oval sheath, its margins bearing 3-8 pairs of linear-lanceolate 
to linear-filiform, hyaline, acuminate, fimbriate, stipular processes 
1-4 mm. long; uppermost cauline leaves narrower, shorter petiolate, 
and often 3-lobed; peduncles weak, usually paired in axils, 1-5 cm. 
long, the umbels usually 1-5-flowered, usually one umbel solitary 
and often subsessile in axils; involucre apparently lacking; flowers 
greenish-white or purplish; calyx lobes obsolete; petals ovate, 0.5- 
0.6 mm. long, plane, glabrous, a little narrowed at either end; ovaries 
sparsely stellate-pubescent and glochidiate; pedicels to 0.5 mm. long, 
hirsute at apex, spreading; fruit ovoid, 2.5 mm. long, 1.5-2 mm. 
broad, 4-winged, rounded at apex, a little cordate, the carpels flat- 
tened dorsally, concave dorsally, the margins a little inrolled, the 
commissural face convex, stellate and papillate with sessile 8-rayed 
hairs on both surfaces, the ribs armed with very slender, stalked, 
"coprinoid," 8-12-rayed glochids up to 1 mm. long, their rays sharply 
reflexed. 

Ancash: Pr. Bolognesi: Cerro al SE de Chiquian, 3500 m., Cer- 
rate 521, 528, 3500-3600 m., Ferreyra 7323. Arequipa: Pr. Arequipa: 
quebrada de San Lazaro, near Arequipa, 9100 ft., Munz 15506; Are- 
quipa, 8-10,000 ft., Stafford 36, Pennell 13209; Volcan de Misti, 
3300-3400 m., Pennell 13224; Chiquata, 3250-3400 m., Vargas 9441a; 
Chacani, 11,000 ft., Stafford 617 A; Pr. Islay, 10 km. east of Islay, 
300 m., Worth & Morrison 1571 4; cerca a la pampa la Joya, arriba 
de Mollendo, 500 m., Ferreyra 6426. Pr. Caraveli: Lomas de Capas 
(Chala), ca. 200 m., Velarde 181. Ayacucho: Pampalca, between 
Huanta and Rio Apurimac, 3200 m., Kittip & Smith 22260. Cusco: 
Pr. Cusco: Cusco, 3000-3600 m., Herrera Lima: Pr. Canta: Obra- 
jillo, Wilkes Expedition. Pr. Huarochiri : Chicre cerro al N. de Hua- 
rochiri, 3700 m., Cerrate & Tovar 1817; Infiernillo, entre San Mateo 
i Rio Blanco, carretera Lima-Huancayo, 3300-3350 m., Ferreyra 7006 
in part; Chicla, 12,000-13,000 ft., Ball; Huamantanga, Mathews 491; 
Matucana, ca. 8000 ft., Macbride & Featherstone 407, 121. 

Ecuador to southern Peru, 2000 to 4000 meters, and in coastal 
Arequipa at 200 to 500 meters; also in Chile and in Mexico. 

Bowlesia tenella Meyen, Reise 1: 466. 1834. B. lobata var. 
humifusa Ball, Jour. Linn. Soc. Bot. 22: 40. 1885; B. mandoni Rusby, 
Mem. Torrey Club 3: 41. 1893. 

Annual, prostrate to ascending, 0.5-3.5 dm. long, dichotomously 
branched from a slender taproot, the stems filiform, sparsely pubes- 
cent with sessile or subsessile, very slender and mostly 4-rayed stel- 



36 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

late hairs, or quite glabrate, the young foliage and inflorescence 
stellate-hirsutulous; leaves mostly alternate, petiolate, thinly mem- 
branaceous, the basal and lower cauline orbicular-reniform, 0.8- 
2.2 cm. long, 1-3 cm. broad, cordate, palmately 5- or 7-lobed about 
to the middle, with narrow sinuses, the lobes ovate, acute or obtuse, 
subequal, mostly entire (or with 1 or 2 lateral teeth), weakly mucro- 
nate, strigulose above with simple or 2-3-forked hairs, hirsutulous 
beneath with sessile or subsessile, 3-5-rayed stellate hairs, bristly on 
veins, the petioles very slender, 1.5-5 cm. long, stellate-hirsute, 
dilated into an oval sheath, its margins bearing 2-4 pairs of linear 
to linear-lanceolate, hyaline, acute or acuminate, fimbriate, stipular 
processes, 1-2 mm. long; upper cauline leaves reduced, but like the 
basal; peduncles obsolete, the umbels usually 1-3-flowered, short- 
pedicellate to subsessile in leaf axils of main stem or on filiform axil- 
lary shoots (which appear like peduncles) ; involucre lacking, but the 
stipular processes and pedicel hairs often forming a concealing semi- 
involucre; flowers yellowish-white; calyx obsolete; petals oval, 0.2- 
0.4 mm. long, plane, glabrous, a little narrowed at either end; ovaries 
glochidiate-roughened; pedicels up to 3 mm. long, hirsute, spreading; 
fruit ovoid, 2-3 mm. long, 1-1.5 mm. broad, obtuse at apex, a little 
cordate, 4-winged, the carpels (always?) dimorphic, strongly flat- 
tened dorsally, plane to strongly concave dorsally, the commissural 
face convex, either sparsely to densely glochidiate-stellate on both 
surfaces and between ribs, or one carpel quite naked, the other armed 
with sessile or stalked, mostly 8-rayed "coprinoid" glochids, the 
stalks unequal, less than 0.5 mm. long. F.M. Neg. 3418. 

"Andes peruvie"nnes," Savatier. Junin: Pr. Junin: Hondores 
cerca a la laguna de Junin, 4200-4300 m., Ferreyra 5278. Lima: 
Pr. Huarochiri: "ex saxosis Andium Peruviae juxta pagum Chicla, 
12-13000' s. m.," Ball, type of B. lobata var. humifusa. Puno: Cor- 
dillera de Puno, Lechler 1701. St. Gavan, Lechler 2665. Pr. Huan- 
cane" : Conima, 3900 m., Aguilar. Pr. Puno: Isla Amantani, 3900 m., 
Aguilar. Tacna: Cordillera de Tacna, 3^000 ft., Meyen; "Laguna 
de Titicaca," Meyen. 

Rocky places at altitudes of 3400 to 4300 meters, in the Andes 
of Peru and Bolivia. 

Bowlesia tropaeolifolia Gillies & Hook. Bot. Misc. 1: 325. 1830. 
B. flexilis Meyen, Reise 1: 348. 1834. B. pulchella Wedd. Chlor. 
And. 2: 188, pi. 67B. 1860. B. cirrosa Phil. Anal. Univ. Chil. 85: 
515. 1894. B. tropaeolifolia var. cirrosa Reiche, Anal. Univ. Chil. 



FLORA OF PERU 37 

104: 783. 1899. B. tropaeolifolia var. heterophylla Speg. Anal. Mus. 
Nac. Buenos Aires 7: 294. 1902. B. tropaeolifolia var. patagonica 
Speg. I.e. B. tropaeolifolia var. gayana Domin, Repert. Sp. Nov. 4: 
299. 1907. B. rupestris Wolff, Bot. Jahrb. 40: 286. 1908. 

Perennial, 0.5-12 dm. high or long, prostrate or half-climbing to 
low and "bushy," dichotomously branched from a multicipital cau- 
dex terminating in a stout, woody, vertical taproot, the stems densely 
to sparsely pubescent with sessile or short-stalked 4-8-rayed stellate 
hairs, or glabrate, the young foliage and inflorescence white- or gray- 
ish-pannose, -tomentose, or merely -hirsutulose, with stellate hairs; 
leaves loosely clustered toward base, alternate below, opposite above, 
membranaceous but thickish, orbicular-reniform, 0.5-3.5 cm. long, 
0.7^4.5 cm. broad, cordate, with a broad to narrow sinus, palmately 
5-, 7-, or 9-lobed, usually about to the middle, the lobes ovate to 
linear-lanceolate, subequal (or the terminal rarely a little elongate), 
generally broadest near the middle, all usually entire, weakly bristly 
mucronulate, green but strigose to hirsute above with appressed or 
ascending, mostly 2-5-rayed, stalked hairs, or glabrate, white-pan- 
nose or -tomentose beneath at least when young, with stalked 5-8- 
rayed hairs; petioles slender, 1-16 cm. long, dilated into an oval 
sheath, its margin bearing 2-5 pairs of linear to ovate-lanceolate, 
hyaline, acute, fimbriate to lacerate stipular processes 0.5-3 mm. 
long; upper cauline leaves somewhat reduced and shorter-petiolate, 
often 3-lobed or even entire; peduncles weak to stout, short to very 
slender, 0.5-5 (-10) cm. long; involucre of several stipule-like linear 
to lanceolate hyaline bracts, these often somewhat obscured by the 
stellate indumentum; flowers yellowish- or greenish-white (rarely 
rose-tinged or purplish) ; calyx lobes linear-lanceolate, scarious, fim- 
briate, 0.5-0.8 mm. long, somewhat persistent, reflexed, shorter than 
petals; petals ovate to oblong, 1-1.5 mm. long, plane, usually stellate- 
pubescent on back, a little narrowed at either end; ovaries white, 
stellate-pannose or -tomentose; pedicels 0.5-2 mm. long, to abortive, 
spreading; fruit ovoid, 1-3 mm. long, 1.5-2.5 mm. broad, more or 
less 2- or 4-winged, narrowed at apex, a little cordate, the carpels 
strongly flattened dorsally, plane to strongly concave dorsally, the 
commissural face convex, densely to sparsely stellate-pubescent on 
both surfaces with 8- or 6-rayed sessile hairs. F.M. Negs. 3407, 
3416, 37062. 

Illustration: Reiche, Bot. Jahrb. 28: pi 1J.4. 1889 (fruit section). 

Arequipa: Pr. Arequipa: Chacani, 11000 ft., Stafford 617; Salinas, 
above Arequipa, 4500 m., Rauh 619; Cordillera between Cotahuasi 



38 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and Cailloma, 4500-4600 m., Weberbauer 6884- Cusco: Pr. Quispi- 
canchis: Ccateca, Kauri, 3900 m., Vargas 918. Huancavelica: Pr. 
Castro- Virreyna: high Andes between Pisco and Ayacucho, Sta. Ine"s, 
4600-4700 m., Weberbauer 5430. Junin: Pr. Yauli: Tunel Mar 
(Jauli), 4000 m., Velarde 1957. Pr. Junin: Juampacucha, 4500 m., 
Aguilar. Lima: Pr. Canta: Huamantanga, Mathews 503. Pr. Hua- 
rochiri: "An d. Lima-Oroya Bahn, Chicla, 3720 m.," Weberbauer 231, 
type of B. rupestris Wolff. Moquegua: Saylapa, near Carumas, 
3600-3700 m., Weberbauer 7349. Pasco: Pr. Pasco: Cerro de Pasco, 
ca. 4300 m., Asplund 11842. Puno: Pr. Puno: hills back of Puno, 
3152 m., Shepard 142; alrededores de Puno, 3900 m., Aguilar 88; 
south-facing, under rocks in stony slides, km. 4 north of Puno on 
road to Juliaca, overlooking Puno, 3800 m., Hutchison 1820; Isla 
Estebes, Vargas 118; Salcedo, Soukup 957; Tiquillaca, 13,000 ft., 
Stafford 1213; Sta. Lucia, Sharpe 59. Tacna: Chulunquaicum, Cor- 
dillera de Tacora, Weddell, type of B. pulchella Wedd. 

High Andes of Peru, southward through Bolivia, Chile, and cor- 
dilleran Argentina to Fuegia and Patagonia, at altitudes of 2000 to 
4700 meters at the north to less than 100 meters at the extreme south. 

CONIUM L. 

Slender, erect, herbaceous, caulescent, branching, glabrous bien- 
nials, the stems spotted, from stout taproots. Leaves petiolate, 
membranaceous, pinnately decompound, the leaflets pinnately in- 
cised. Petioles sheathing. Inflorescence a compound dichasium of 
loose compound umbels; peduncles terminal and axillary. Involucre 
of numerous, inconspicuous, lanceolate bracts. Rays numerous, 
spreading-ascending. Involucel of numerous bractlets like the 
bracts, shorter than pedicels. Pedicels spreading. Flowers white; 
petals obovate or cuneate with a narrower inflexed apex; calyx teeth 
obsolete; styles reflexed, the stylopodium depressed-conic. Carpo- 
phore entire. Fruit broadly ovoid, compressed laterally, glabrous; 
ribs prominent, obtuse, undulate, crenate; vittae very small and 
numerous, irregular; seed terete in transection, the face deeply 
and narrowly sulcate. 

A genus of two species, one native of Eurasia, the other South 
African. 

Conium macula turn L. Sp. PI. 243. 1753. Cicuta officinalis 
Crantz, Class. Umbell. 98. 1767. Coriandrum cicuta Crantz, Stirp. 



FLORA OF PERU 39 

Austr. 3: 100. 1767. Conium cicuta Neck. Delic. 142. 1768. Cicuta 
major Lam. Fl. Fr. 3: 456. 1778. Coriandrum maculatum Roth, Fl. 
Germ. 1: 130. 1788. Conium maculosum Pall. Reise Siidl. Statth. 1: 
478. 1799. Sium conium Vest, Man. Bot. 513. 1806. Selinum conium 
Krause in Sturm, Fl. Deutsch. ed. 2. 12: 79. 1904. 

Plants 5-30 dm. high; leaf blades broadly ovate, 1.5-3 dm. long, 
5-30 cm. broad, pinnately decompound into pinnately incised seg- 
ments; petioles dilated; involucre of ovate-acuminate, short bracts; 
rays 15-25 mm. long, subequal; involucel of bractlets like the bracts, 
with a conspicuous midrib; pedicels 4-6 mm. long; fruit 2-2.5 mm. 
long, about 2 mm. broad, the ribs very prominent in the dry fruit. 

Illustrations: Engl. & Prantl, Pflanzenf. 3 (8): 168. 1898 (fruit). 
Reichenb. Ic. Fl. Germ. 21: pi. 2032. 1867. Sturm, Fl. Deutsch. 
ed. 2. 12: pi. 38. 1904. 

Ancash: Pr. Bolognesi: alrededores de Aquia, 3400 m., Ferreyra 
12161. Arequipa: Pr. Arequipa: cerca del Rio Chile, borde de ace- 
quia, 2200 m., Angulo 1786. Cusco: Pr. Cusco: cerro Sape, cerca a 
la ciudad de Cusco, 3400 m., Ferreyra 2669; Cusco, 3200 m., Herrera 
308, Soukup 55; Cerro Picchu cerca al Cusco, 3500-3700 m., Ferreyra 
9844- Huancavelica: Pr. Huancavelica: roadside weed, canyon of 
the Rio Mantaro north of Mejorada, km. 372 from Lima on Abancay- 
Huancayo Road, 2950 m., Hutchison 1670. Huanuco: Pr. Dos de 
Mayo: on previously cultivated land, 2700 m., Woytkowski 82; Cha- 
vinilla, alrededores del pueblo, 3400 m., Woytkowski 1013. Junin: 
Pr. Jauja: Concepcion, valle del Mantaro, borde de chacra, 3200- 
3300 m., Ferreyra 12911; alrededores de Jauja, 3300 m., Ridoutt 891; 
Valle de Concepcion, Hood. Pr. Tarma: alrededores de Tarma, 
3200 m., Esposto, 3150 m., Ldpez 789; Acobamba, aba jo de Tarma, 
2900 m., Aguilar 528; cerca a Cochas, entre Tarma y La Oroya, 
3500-3600 m., Ferreyra 11427; entre Tarma y Oroya, subiendo de 
Tarma a la cumbre, 3300-3500 m., Ferreyra 3795; Tarma, 3000- 
3200 m., Killip & Smith 21882. Pr. Huancayo: cerca a Huancayo, 
Chavez; alrededores de Huancayo, 3317 m., Soukup 2012; Huancayo, 
3317 m., Soukup 3551. Lima: Pr. Huarochiri: Llucanchi abajo de 
Huarochiri, 3350 m., Cerrate & Tovar 1759. Puno: Pr. Huancane": 
Conima, 3900 m., Aguilar. 

Locally known as culantrillo, cicuta, and cicuta-asna-acha; and 
common along roadsides in many parts of Peru. Introduced from 
Europe, widespread as a weed in moist places in North and South 
America. Poisonous, the poison hemlock of the Greeks. 



40 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

CORIANDRUM L. 

Slender, erect, herbaceous, caulescent, glabrous annuals, from 
slender taproots. Leaves petiolate, membranaceous, pinnately dis- 
sected. Petioles sheathing. Inflorescence of loose compound um- 
bels. Peduncles terminal and lateral. Involucre usually wanting. 
Rays few, spreading-ascending. Involucel of a few small, narrow 
bractlets. Pedicels spreading. Flowers white or rose; petals oblong 
with a narrower inflexed apex, the outer usually radiant; calyx teeth 
prominent, acute, often unequal; styles slender, spreading, the stylo- 
podium conic. Carpophore 2-parted to base. Fruit orbicular, terete, 
with a hard pericarp, glabrous, the carpels boat-shaped, not separat- 
ing readily at maturity; primary ribs filiform, the secondary filiform 
or obscure; vittae absent; seed flattened dorsally in transection, the 
face concave. 

A genus of two species of the warm temperate and subtropical 
regions of the Old World. 

Coriandrum sativum L. Sp. PI. 256. 1753. Selinum corian- 
drum Krause in Sturm, Fl. Deutsch. ed. 2. 12: 163. 1904. 

Plants 2-7 dm. high; basal leaves ovate, the blades 3-15 cm. long, 
2-10 cm. broad, simple and ternately or pinnately lobed, or pinnate, 
the leaflets flabelliform, cuneate at base, 1-2 cm. long, 0.5-1 cm. 
broad, variously toothed or incised; petioles 2-15 cm. long; cauline 
leaves pinnately dissected, the leaflets linear to filiform, 2-15 mm. 
long, 0.5-1.5 mm. broad, obtuse, entire; peduncles 3-10 cm. long, 
occasionally abortive; involucre usually wanting; rays 2-8, 1-2.5 cm. 
long; involucel dimidiate, of a few linear bractlets, 2-4 mm. long; 
pedicels 2-5 mm. long; calyx teeth ovate-lanceolate, unequal, the 
outer up to 1 mm. long; petals white or rose, the outer radiant; fruit 
1.5-5 mm. in diameter. 

Illustrations: Reichenb. Ic. Fl. Germ. 21: pi. 2043. 1867. Engl. 
& Prantl, Pflanzenf . 3 (8) : 159. 1898, fl ., frt. Hegi, 111. Fl. Mitt.-Eur. 
5 (2):pl.l94,f.2. 1926. 

lea: Pr. lea: lea, terrenos de cultivo, 300 m., Herb. San Marcos. 
Junin: Pr. Tarma: Coyllarbamba, cerca de Acobamba, 2200 m., 
Ochoa 184; Tarma, 3000-3200 m., Killip & Smith 21915. Pr. Huan- 
cayo: cultivado, 3200 m., Chavez. Lima: Pr. Lima: Otero, Rimac, 
Lima, terrenos cultivados, 157 m., Ridoutt 887. Loreto: Pr. May- 
nas: La Victoria on the Amazon River, Williams 2790; Caballo- 
Cocha on the Amazon River, Williams 2394; Paraiso, Alto Rio Itaya, 



FLORA OF PERU 41 

145 m., Williams 3279. San Martin: Pr. Lamas: Roque, banana 
plantation, Melin 293. 

The coriander of commerce, native of the Mediterranean region 
and widely adventive in the western hemisphere. Locally known as 
culandra and culandro. 

DAUGUS L. 

Carota Rupr. Fl. Ingr. 466. 1860. 

Low or tall, erect, herbaceous, caulescent, branching, pubescent 
annuals or biennials, from taproots. Leaves petiolate, membrana- 
ceous, pinnately decompound, the leaflets small and narrow. Petioles 
sheathing. Inflorescence of loose compound umbels, or subcompact 
by incurving of the rays after anthesis; peduncles terminal and axil- 
lary. Involucre of numerous dissected or entire bracts, or wanting. 
Rays few to numerous, spreading or the outer often connivent over 
the short inner ones. Involucel of numerous toothed or entire bract- 
lets, or wanting. Pedicels spreading, unequal. Flowers white, or 
the central flower of each umbellet purple, or rarely all reddish, pink- 
ish, or yellow; petals obcordate, unequally cleft, with a narrower in- 
flexed apex, the outer often radiant; calyx teeth obsolete to evident; 
styles short, the stylopodium conic. Carpophore entire or bifid at 
apex. Fruit oblong to ovoid, compressed dorsally; primary ribs fili- 
form, bristly; secondary ribs winged, the wings divided into a single 
row of barbed or glochidiate prickles; vittae solitary under the sec- 
ondary ribs, 2 on the commissure; seed flattened dorsally in tran- 
section, the face shallowly concave to nearly plane. 

A genus of about 25 species of wide distribution. 

Fruiting umbels many-rayed, compact in fruit by incurving of the 
outer rays D. carota. 

Fruiting umbels few-rayed, open in fruit, the rays spreading-ascend- 
ing D. montanus. 

Daucus carota L. Sp. PI. 242. 1753. Caucalis carota Crantz, 
Class. Umbell. 113. 1767. C. daucus Crantz, Stirp. Austr. 3: 125. 
1767. Carota saliva Rupr. Fl. Ingr. 468. 1860. 

Plants biennial, 1.5-12 dm. high, the stems solitary, glabrous to 
retrorsely hispid; leaf blades oblong, 5-15 cm. long, 2-7 cm. broad, 
the leaflets linear to lanceolate, 2-12 mm. long, 0.5-2 mm. broad, 
acute, mucronate, entire or few-cleft, glabrous to hispid especially 



42 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

on veins and margins; petioles 3-10 cm. long; divisions of cauline 
leaves often elongate; peduncles 2.5-6 dm. long, retrorsely hispid; 
involucre of filiform, elongate, pinnately divided or rarely entire, 
scarious-margined bracts, 3-30 mm. long, usually reflexed; rays 
numerous, unequal, 3-7.5 cm. long, compact in fruit; involucel of 
linear, acuminate, entire or rarely pinnate, more or less scarious, 
ciliate bractlets, equaling or exceeding the flowers; pedicels unequal, 
3-10 mm. long; flowers white, yellow or pinkish, the central flower 
of each umbellet usually purple or pinkish; fruit ovoid, 3-4 mm. long, 
about 2 mm. broad, broadest at middle. 

Illustrations: Reichenb. Ic. Fl. Germ. 21: pi. 2000. 1867. Bailey, 
St. Cycl. Hort. 674, 675. 1922. 

Lima: Pr. Lima: cercanias de Lima, Isern (Cuatrecasas 2465). 

The cultivated carrot, native of Europe, adventive throughout 
the western hemisphere. 

Daucus montanus Humb. & Bonpl. ex Spreng. in R. & S. Syst. 
Veg. 6: 482. 1820. D. toriloides DC. Prodr. 4: 214. 1830. Torilis 
peruviana Presl in DC. Prodr. 4: 214. 1830, in synon. Daucus aus- 
tralis Poepp. in DC. I.e. 

Plants annual (or biennial?), 1.2-10 dm. high, the stems usually 
solitary, erect, simple or few-branched, papillate-hispid with mostly 
retrorse hairs to glabrate; leaf blades oblong, 3.5-11 cm. long, 2.5- 
5 cm. broad, the leaflets linear, 2-5 mm. long, about 1 mm. broad, 
acute, more or less hispid; petioles 3-12 cm. long; peduncles 4.5- 
25 cm. long, retrorsely papillate-hispid; involucre of foliaceous, pin- 
nately decompound bracts, the divisions short, linear; rays 4-20, 
unequal, spreading, 1.5-12 cm. long; involucel of a few linear, acute 
bractlets shorter than pedicels; pedicels unequal, 3-20 mm. long; 
flowers whitish, greenish, yellowish, pink, or red-purple; fruit oblong, 
3-6 mm. long, 2-3 mm. broad, the commissural surface with 2 rows 
of hispidulous hairs. 

Illustrations: Saunders, Refug. Bot. 5: pi. 299. 1873. 

Ancash: Pr. Bolognesi: Mahuay, 4300 m., Cerrate 2221; Distr. 
de Huasta, 3800 m., Cerrate 2521. Pr. Huaraz: gravelly soil between 
rocks, Cerro Shaurema, southeast of Huaraz, 3200-3300 m., Pennell 
15290. Apurimac: Pr. Abancay: Ampuy, 3200 m., Stork, Norton & 
Vargas 10610; Quera, Lambrana, 3050 m., Vargas 1958; Abancay, 
Banco de Fomento Agropecuario 103. Arequipa: Pr. Islay: near 
Mollendo, 250 m., Hutchison 1861a. Cusco: Pr. Urubamba: Machu 



FLORA OF PERU 43 

Picchu, 1800 m., Velarde 1113; Machu Picchu Station, Hutchison 
1763; ruins of Machu Picchu, Hutchison 1764, Mathias 3676. Pr. 
Cusco: ruinas de Sacsayhuaman, cerca a la ciudad de Cusco, 3400- 
3500 m., Asplund 2637, Ferreyra 2637, 3600 m., Vargas 1179; Cusco, 
Rauh & Hirsch 718, Hen era 3158, 2401. Pr. Paucartambo : laderas 
de Paucartambo, arcillo-pedregoso, 3400 m., Vargas 911. Pr. Calca: 
Calca, Vargas 3888. Pr. Anta: Cordillera Salcantay, Pampa Soray, 
4000 m., Rauh 1495. Huancavelica: Pr. Tayacaja: entre Colca- 
bamba y Paucarbamba, 2800 m., Tovar 2059. Pr. Huancavelica: 
abajo de Huando, 3300 m., Tovar 1222; west of and above Huando, 
road to Huancavelica, ca. 5 km. before (above) bridge over Rio 
Palca, 3650 m., Hutchison 1678. Huanuco: Pr. Huanuco: open 
grassy banks, 15 mi. southeast of Huanuco, 10,500 ft., Macbride & 
Feather stone 2119; Mito, 9000 ft., Macbride & Feather stone 1737; 
Pillao, 2700 m., Woytkowski 105, 34105, 35105; roadside, about 1 km. 
above Chinchao, Mathias 5163; Carpish, cumbre entre Huanuco y 
Tingo Maria, 2700-2800 m., Ferreyra 10017; Carpish, slope towards 
Huanuco, 2750 m., Asplund 13097; Mitotambo arriba de Mito, 3200- 
3300 m., Ferreyra 10358; Hda. Paty, abajo de Carpish, 1700-1800 m., 
Ferreyra 9388; Cuchero, Poeppig 1292; Peruvia subandina, locis 
cultis ad Cuchero, Poeppig 1292. Pr. Dos de Mayo: Chavinillo, 
3600 m., Woytkowski 1042, 3400 m., Woytkowski 837. Junin: Pr. 
Tarma: entre Palca y Huacapistana, 2800 m., Ferreyra 11009; Hua- 
capistana, entre Tarma y San Ramon, margen izq. del Rio Tarma, 
1800-1900 m., Ferreyra 479a; Tarma, debajo de Huacapistana, cerca 
al puente 1^ km., 1700-1800 m., Ferreyra 11206; entre Tarma y 
Oroya, 3600 m., Constance & Tovar 2360; San Juan, cerca a Hua- 
capistana, entre Tarma y San Ramon, 2200-2300 m., Ferreyra 11317; 
1800-1900 m., Ferreyra 11248; Huacapistana, 1800-2400 m., Killip & 
Smith 24134; Valley of the Huasahuasi, Hutchison 1110; mouth of 
gorge, lower end of Huasahuasi Valley, Hutchison 1140; Chancha- 
mayo Valley, above La Merced at Cumbre Yacunay near summit, 
2500 m., Hutchison 1182. Pr. Huancayo: agua de las Virgenes, cerca 
a Huancayo, 3270-3300 m., Tovar 329. Pr. Yauli : Oroya near Lima, 
10-13,000 ft., Kalenborn 62; alrededores de Yauli-Hca., 3500-3600 m., 
Tovar 3021. La Libertad: Pr. Bolivar: arriba de Longotea, 3000 m., 
L6pez & Sagastegui 3179. Pr. Trujillo: lomas del Cerro Campana, 
600 m., Angulo & Ldpez 2029. Lambayeque: Pr. Lambayeque: 
border depts. Lambayeque & Piura, abra Porculla Pass on Olmos- 
Maranon Highway, 1650 m., Hutchison 1380. Lima: Pr. Huaro- 
chiri: Chicla, 3700 m., Asplund 11325; on Lima-Oroya Hwy. above 



44 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Rio Blanco, 3600 m., Hutchison 1659; Rio Blanco, about 12,000 ft., 
Macbride & Feather stone 671. Pr. Lima: Lomas de Lurin, al sur de 
Lima, 350^400 m., Ferreyra 11912; Lomas de Atocongo, 200-300 m., 
Ferreyra 11993. Pr. Canta: Antaicocha, Cerro Colorado, east of 
Canta, 3500-3600 m., Pennell 14628. Piura: Pr. Huancabamba: 
arriba de Palambla, 1500-1600 m., Ferreyra, Cerrate & Tovar 10841. 
San Martin: Pr. Mariscal Caceres: on road to Divisoria, 59 km. 
from Tingo Maria on highway to Pucallpa, 1540 m., Allard 21307; 
La Divisoria, Mathias 3511. Pr. Lamas: San Roque, 1350-1500 m., 
Williams 7129. Without locality: Haenke. 

A weedy native, with odor of parsley, widespread in western 
South America from Chile to Colombia, extending north to Sonora. 

DOMEYKOA Phil. 

Reference: Math. & Const. A revision of Asteriscium and some 
related hydrocotyloid Umbelliferae, Univ. Calif. Publ. Bot. 33: 99- 
184. 1962. 

Prostrate or spreading, fleshy annual or woody-based perennial 
herbs, caulescent, dichotomously branched with flexuous, terete 
branches, glabrous and glaucous. Leaves petiolate, membranaceous, 
simple, lobed, or 1-3-ternate or -pinnate, palmately, ternately, or 
pinnately veined; cauline leaves opposite, alternate, or weakly fas- 
cicled, petiolate to sessile, or sessile and conspicuously clasping to 
perfoliate. Petioles sheathing at base. Inflorescence of simple, few- 
to many-flowered globose umbels. Peduncles solitary in or opposite 
upper leaf axils, or umbellately clustered in terminal groups of 2-7, 
sometimes obsolete. Involucre of several linear-lanceolate or orbic- 
ular, free or partially connate bracts, usually equaling to longer than 
pedicels. Flowers white, yellowish, or purple; calyx lobes evident, 
ovate to obovate, entire or trifid; petals broadly ovate to obovate, 
acutish, plane or a little incurved, or very shortly inflexed at apex, 
stipitate to subsessile, the dorsal gland evident; styles short to mod- 
erately long, the stylopodium depressed to low-conic. Carpophore 
wanting. Pedicels short. Fruit orbicular to oval, subtetragonal, 
truncate at apex, more or less constricted at the commissure and 
compressed laterally, glabrous, the mature carpels cuneate-rectangu- 
lar in transection, about as broad as long, with plane sides, plane or 
very weakly concave dorsally, the margins prominent but unwinged, 
the commissural face intrusively sulcate, the ribs filiform; vittae 
small, solitary under the ribs; a thin woody endocarp surrounding 
the seed cavity. 



FLORA OF PERU 45 

A genus of about four species, endemic to Peru and northern 
Chile, confined to the coastal slope. 

Basal leaves repandly spinulose-dentate; flowers yellowish-white; 
calyx lobes about 0.5 mm. long, often conspicuously 3-fid or 
3-lobed; styles 0.8-1 mm. long D. amplexicaulis. 

Basal leaves deeply 3- or 5-lobed or parted; flowers usually purplish 
or rose-colored; calyx lobes obscure, up to 0.3 mm. long, entire; 
styles about 0.5 mm. long D. saniculifolia. 

Domeykoa amplexicaulis (Wolff) Math. & Const. Univ. Calif. 
Publ. Bot. 33: 175, /. 84. 1962. Astericium amplexicaulis Wolff, Bot. 
Jahrb. 40: 289. 1908. 

Slender, prostrate or spreading annual herbs, the branches 1- 
10 dm. long, striate; leaves somewhat rosulate at base, orbicular to 
obovate, 0.9-3.5 cm. long, 1.2-5.0 cm. broad, a little cuneate, re- 
pandly spinulose-dentate; petiole 1-4 cm. long, weakly dilated and 
narrowly scarious-margined; cauline leaves prominent, alternate, 
sessile, conspicuously cordate-clasping or perfoliate, repandly spin- 
ulose-dentate; peduncles mostly umbellately clustered in terminal 
groups of 2-4, a few solitary in upper axils, spreading-ascending, 
0.5-4 cm. long, each group or solitary peduncle subtended by a disc- 
shaped perfoliate leaf or bract; umbels compact, 3-8 mm. in diam- 
eter, with usually 15-40 flowers; bracts 5-8, obovate to narrowly 
oblanceolate, 1-4 mm. long, 1-3 mm. broad, often imbricate, some- 
what connate, conspicuously lacerate at apex; flowers yellowish- 
white; calyx lobes oval to obovate, 0.5 mm. long, deeply 3-fid or 
-lobed, or rarely entire, conspicuously shorter than petals; petals 
spatulate-obovate, 0.7-1 mm. long, abruptly acuminate, the short 
apex slightly incurved to inflexed, conspicuously stipitate; stylo- 
podium low-conic, shorter than the slender, spreading styles; ped- 
icels stout, 1-1.5 mm. long, spreading-ascending; fruit orbicular or 
oval, ca. 1.5 mm. long, 1.25-1.5 mm. broad, sharply angled, the 
carpels slightly concave dorsally. F.M. Neg. 3465. 

Arequipa: Pr. Caraveli: Atico, entre Chala y Camana, 100- 
200 m., Ferreyra 12489, 100 m., Ferreyra 11540; Atico, entre Chala 
y Ocofia, 50-100 m., Ferreyra 11944, 50-100 m., Ferreyra 12010; 
ca. 5 km. south of Atico, km. 730 south of Lima on Pan American 
Highway, 200 m., Hutchison 1869; Lomas de Atiquipa, 400 m., Ve- 
larde 4549a. Pr. Islay: Mollendo, dry sandy slope just above sea 



46 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

level, Stafford 805; Tambo prope Mollendo, 300-600 m., Weberbauer 
1560, type. 

Lomas, Arequipa at elevations of 50 to 600 meters. 

Domeykoa saniculifolia Math. & Const. Univ. Calif. Publ. 
Bot. 33:179,/. 35. 1962. 

Rather slender, prostrate or ascending, annual (?) or weakly peren- 
nial herb, the branches 1-3 dm. long, faintly ribbed; leaves somewhat 
rosulate at base, orbicular to obovate, 0.6-2 cm. long, 0.8-2.2 cm. 
broad, cuneate, deeply 3- or 5-lobed or -parted, the lobes oblanceolate 
to obovate, coarsely spinulose-dentate; petiole 2-5 cm. long, weakly 
dilated; cauline leaves prominent, alternate, sessile, oval, conspicu- 
ously cordate-amplexicaul, deeply 3-7-lobed or coarsely toothed; 
peduncles mostly umbellately clustered in terminal groups of 2 or 3, 
or some solitary in upper axils, spreading-ascending, 0.5-1 cm. long, 
each group or solitary peduncle subtended by a disc-shaped perf oliate 
leaf or bract; umbels compact, 5-7 mm. in diameter, with usually 
8-15 flowers; bracts 5 or 6, narrowly oblanceolate to orbicular, 2- 
4 mm. long, 0.5-3 mm. broad, more or less connate below, acute and 
entire to lacerate at apex; flowers purplish-violet, rose-colored, or 
rarely greenish-white; calyx lobes broadly triangular, entire, up to 
0.3 mm. long, but usually obscure or obsolete; petals ovate, 0.8- 
1 mm. long, acutish, plane or a little incurved but without a definite 
inflexed apex, shortly stipitate; stylopodium low-conic, about as long 
as the short, recurved styles; pedicels stout, 1-1.5 mm. long, angled, 
spreading-ascending; fruit orbicular, 1-1.5 mm. long and broad, sub- 
tetragonal, sharply angled, the carpels slightly concave dorsally. 

Arequipa: Pr. Caraveli: Los Cerillos entre Nazca y Chala, 700 m., 
Ferreyra 18451; Los Cerillos, 52 km. al sur de Nazca, 700 m., Rahn. 
lea: Pr. Nazca: Lomas de San Nicolas, al sur de Nazca, 800-900 m., 
Ferreyra 13395. Moquegua: Pr. Mariscal Nieto: Lomas de Ilo, 600- 
650 m., Ferreyra 12571+; 4-5 km. north of Ilo on the pampa, 500 m., 
Hutchison 483. Tacna: Pr. Tacna: Lomas cerca al Morro Sama, 
500-600 m., Ferreyra 12544, type. 

Lomas, lea to Tacna, at elevations of 500-900 meters. 

EREMOCHARIS Phil. 

Reference: Math. & Const. A revision of Asteriscium and some 
related hydrocotyloid Umbelliferae, Univ. Calif. Publ. Bot. 33: 99- 
184. 1962. 



FLORA OF PERU 47 

Asteriscium sect. Bustillosia Benth. & Hook. Gen. Plant. 1: 877. 
1867, pro parte; Asteriscium subgen. Bustillosia Drude in Engl. & 
Prantl, Pflanzenf . 3 (8) : 134. 1897, pro parte. Asteriscium subgen. 
Gymnophytum Wolff, Bot. Jahrb. 40: 292-293. 1908, pro parte. 

Erect or decumbent, suffrutescent or woody-based perennial 
herbs, caulescent, dichotomously branched with flexuous or spines- 
cent branches, glabrous and glaucous. Leaves alternate or fascicled, 
petiolate, membranaceous to fleshy, 1-3 pinnately or ternately di- 
vided or lobed, rarely entire, cuspidate, pinnately veined. Petioles 
obscurely sheathing. Inflorescence of simple, few- to many-flowered 
globose umbels. Peduncles mostly umbellately clustered in terminal 
groups of 2-7, some solitary in upper axils. Involucre of several 
small, lanceolate to ovate, entire bracts shorter than pedicels. 
Flowers yellow or greenish-white to deep maroon or reddish-violet; 
calyx lobes evident, ovate-lanceolate to triangular-ovate, usually 
conspicuously shorter than the petals; petals oval to obovate with 
a narrower inflexed apex */ to about as long as the blade, stipitate 
to subsessile, the dorsal gland conspicuous to evident; styles short 
to slender, the stylopodium low-conic. Carpophore wanting. Ped- 
icels slender. Fruit oblong-oval to orbicular, subtetragonal, truncate 
at apex, more or less constricted at the commissure and compressed 
laterally, glabrous, the mature carpels cuneate-rectangular in tran- 
section, about as broad as long with plane sides, plane or very weakly 
concave dorsally, the commissural face intrusively sulcate, the ribs 
filiform; vittae small, solitary under the ribs; a thin woody endocarp 
surrounding the seed cavity. 

A genus of nine species, endemic to Peru and northern Chile, and 
largely confined to the coastal slope. 

Leaves all entire, white-margined E. integrifolia. 

Leaves deeply trilobed to ternately dissected, not evidently margined. 
Branches spinescent, the older with exfoliating bark. 
Branches ribbed; umbels 10-15 mm. in diameter, 8-20-flowered; 

filaments 4-5 mm. long, long-exserted E. hutchisonii. 

Branches striate; umbels 3-10 mm. in diameter, 3-12-flowered; 

filaments 1.5-2 mm. long, shortly exserted. 
Erect, strongly spinescent shrub; leaves prominently fasci- 
cled; petals falsely emarginate at point of inflexion; 

Cusco E. triradiata. 

Decumbent and weakly spinescent; leaves scarcely fascicled; 
petals obtuse at point of inflexion; Tacna. . . .E. confinis. 



48 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branches slender and flexuous, not spinescent, without promi- 
nently exfoliating bark. 

Petal apex much shorter than blade; styles little longer than 
stylopodium E. ferreyrae. 

Petal apex equaling to surpassing blade; styles markedly longer 

than stylopodium. 
Basal leaves 2-3-ternate, the ultimate divisions filiform. 

E. piscoensis. 

Basal leaves 3-lobed or 3-parted, the ultimate divisions linear- 
lanceolate to orbicular. 
Cauline petioles prominently winged; flowers violet. 

E. tripartite,. 
Cauline petioles unwinged; flowers yellow. . . E. longiramea. 

Eremocharis confinis Johnston, Contr. Gray Herb. 85: 173. 1929. 

Low, decumbent shrub, 3-4 dm. tall, intricately branched, the 
branches spinescent, 3-15 cm. long, terete and finely grooved, the 
older with exfoliating bark; leaves numerous, alternate or weakly 
fascicled below, fleshy, 2-2.5 cm. long, 0.5-1.2 cm. broad, deltoid, 
biternate, the primary lobes narrowly spatulate, 3-8 mm. long, 3- 
5 mm. broad, divaricate, subequal, the ultimate divisions up to 
3 mm. long, often mucronate-dentate; petiole linear, 1-2 cm. long, 
narrowly dilated toward base; cauline leaves little reduced upward, 
the uppermost trilobed; peduncles in groups of 2-4, spreading- 
ascending, 0.5-2.5 cm. long, a little attenuate toward apex, each 
subtended by a lanceolate to oblong leaf 1-2 mm. long; umbels com- 
pact, 3-7 mm. in diameter, with usually 3-12 flowers, the bracts 4-7, 
oblong or lanceolate, mostly obtuse, 1-2 mm. long; flowers greenish- 
white or deep purple; calyx lobes triangular-acuminate, 0.3-0.5 mm. 
long, conspicuously shorter than petals; petals orbicular to obovate, 
about 1.5 mm. long, with a cuneate, inflexed apex about 2/3 length 
of blade; stylopodium much shorter than the slender, spreading styles; 
pedicels stout, 1-2 mm. long, spreading; fruit oval, 2 mm. long, 
1.5 mm. broad, angled, the carpels slightly concave dorsally, the 
margins prominent but unwinged. 

Illustrations: Math. & Const. Univ. Calif. Publ. Bot. 33: 159, 
/. 25. 1962. 

Tacna: Pr. Tarata: Candarave, 2900 m., Weberbauer 7369, type; 
along Tarata-Tacna road midway between the two towns, west side 
of the pass, 1950 m., Hutchison 1834. 



FLORA OF PERU 49 

Known only from the area of the type collection in "open forma- 
tion, herbs, rainy green shrubs and Cereus" 

Eremocharis ferreyrae Math. & Const. Univ. Calif. Publ. Bot. 
33:163,/. 29. 1962. 

Slender, woody-based perennial herbs, 3-7 dm. tall, the branches 
slender, ascending, terete, striate, but neither definitely grooved nor 
ribbed; leaves rather numerous, solitary or very weakly fascicled 
below, 1.5-3 cm. long and broad, ovate-deltoid, deeply trilobed, the 
lobes ovate, 5-20 mm. long, 3-15 mm. broad, the central a little 
larger than the two lateral, all spinose-dentate or -lobed; petiole 
linear, 3-5 cm. long, only the short oblong sheath slightly dilated; 
cauline leaves reduced upward, becoming trifid or even entire; pe- 
duncles mostly in groups of 3-5, spreading-ascending, 1^4 cm. long, 
scarcely attenuate at apex, each subtended by a lanceolate to ovate 
leaf 2-5 mm. long; umbels compact, 4-7 mm. in diameter, with usu- 
ally 20-30 flowers, the bracts lanceolate, acute, 1-3 mm. long; flowers 
bright yellow to greenish-white; calyx lobes broadly ovate, about 
0.2 mm. long, conspicuously shorter than the petals; petals nearly 
orbicular, 0.8 mm. long, with a subulate inflexed apex about Y^ length 
of blade; stylopodium a little shorter than the recurved styles; ped- 
icels stout, about 1 mm. long, spreading; fruit orbicular, 1.5 mm. long 
and broad, sharply angled, the carpels slightly concave dorsally, the 
margins very narrowly winged. 

Arequipa: Pr. Caravel! : Atico, lomas de Atiquipa, 10-20 m., Fer- 
reyra 6463, type; Atico, entre Chala y Camana, 100-200 m., Ferreyra 
12^91; Lomas entre Chala y Atico (La Costanera), 5 m., Angulo 2574. 

Lomas, Arequipa. Known only from the region of the type local- 
ity, at elevations of 5 to 200 meters. 

Eremocharis hutchisonii Math. & Const. Univ. Calif. Publ. 
Bot. 33: 161, /. 26. 1962. 

Low shrub, 3-10 dm. tall, intricately branched, the branches 
short, stiff, and somewhat spinescent, angled and prominently ribbed, 
the older with exfoliating bark; leaves sparse, alternate, weakly fas- 
cicled below, 2-3 cm. long, 1.5-2 cm. broad, ovate, 1-2-ternate, the 
primary lobes oblong-spatulate, 0.5-1.5 cm. long, 0.5 cm. broad, 
divaricate, subequal, the ultimate divisions 1-3 mm. long, ovate- 
lanceolate, mucronate; petiole broadly linear, 1-3 cm. long, grad- 
ually dilated toward base; cauline leaves little reduced upward, the 
uppermost trilobed; peduncles in groups of 3-5, many solitary, 



50 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

spreading, 0.5-2 cm. long, a little attenuate toward the apex, each 
subtended by an ovate leaf 0.5-2 mm. long; umbels lax, 10-15 mm. 
in diameter, with usually 8-20 flowers, the bracts about 5, oblong, 
1-2 mm. long; flowers purplish-brown; calyx lobes ovate, 0.3-0.5 mm. 
long, conspicuously shorter than the petals; petals oval, about 1.5 
mm. long, with a narrower, inflexed apex of about equal length; 
stylopodium much shorter than the very slender, spreading styles; 
pedicels slender, 3-6 mm. long, spreading; fruit oval, 2 mm. long, 
1.5 mm. broad, sharply angled, the carpels strongly concave dorsally, 
the margins prominent. 

Arequipa: Pr. Caraveli: Incuyo-Chala, 3300 m., Hutchison 1277, 
type. 

Known only from the type collection. 

Eremocharis integrifolia Math. & Const. Univ. Calif. Publ. 
Bot. 33:171,/. 32. 1962. 

Low shrub, 3-6 dm. tall, much-branched, the branches slender, 
ascending, somewhat spinescent, terete but with filiform grooves; 
leaves fairly numerous, alternate and often fascicled, especially below, 
the basal and lower cauline 3-4 cm. long, 3-6 mm. broad, narrowly 
lanceolate, entire, narrowly white-margined, tapering insensibly at 
base and with no evidently distinct petiole or sheath; cauline leaves 
reduced upward, but otherwise similar to the basal; peduncles in 
groups of 2-4, ascending, 1-3 cm. long, conspicuously attenuate at 
the apex, each subtended by a linear-lanceolate to lanceolate, usually 
acute leaf 3-12 mm. long, 1-3 mm. broad, often inserted at 
different levels; umbels lax, 5-8 mm. in diameter, with 3-10 flowers, 
the bracts lanceolate to oval, obtuse, 0.5-1.5 mm. long; flowers green- 
ish-purple; calyx lobes ovate, acute, about 0.8 mm. long, almost as 
long as the petals in bud; petals orbicular, about 0.8 mm. long, taper- 
ing into a linear-lanceolate inflexed apex about 2/3 as long as the 
blade; stylopodium much shorter than the slender styles; pedicels 
2-3 mm. long, ascending; fruit orbicular, 1.5-2 mm. long and broad, 
sharply angled, the carpels truncate to slightly concave dorsally, the 
margins acute. 

"Andes Peruvie"nnes," 1876-1879, Savatier. Lima: Pr. Huaro- 
chiri: Matucana, 2400 m., Asplund 10950, type; Matucana, about 
8000 ft., Macbride and Feather stone 555. 

Known only from the vicinity of the type locality, at an eleva- 
tion of 2400 to 2600 meters. 



FLORA OF PERU 51 

Eremocharis longiramea (Wolff) Johnston, Contr. Gray Herb. 
85: 174. 1929. Asteriscium longirameum Wolff, Bot. Jahrb. 40: 292. 
1908. Eremocharis dissecta Johnston, I.e. 

Slender subshrub or woody-based perennial herb, 0.5-1.5 m. tall, 
much-branched above, the branches slender, strictly erect or ascend- 
ing, terete but with definite filiform ribs; leaves sparse, alternate, 
the basal 5-7 cm. long, 3-6 cm. broad, ovate-cuneate, biternate, the 
leaflets cuneate-oblanceolate, subequal, irregularly trilobed, the lobes 
cuneate to linear-lanceolate, or dentate, 3-5 cm. long, 1-2 cm. broad; 
petiole linear, 2-3 cm. long, only the very short sheath dilated; cau- 
line leaves reduced upward, becoming deltoid, ternate, and subsessile; 
peduncles in groups of 2-5, very slender to rather stout, ascending or 
spreading, mostly 0.5-5.5 cm. long, scarcely attenuate at apex, each 
subtended by a lanceolate to linear leaf 0.5-7 mm. long; umbels lax, 
8-15 mm. in diameter, with usually 6-25 flowers, the bracts lanceo- 
late, 0.5-2 mm. long; flowers yellow to greenish-white; calyx lobes 
triangular, acuminate, 0.3-0.6 mm. long, conspicuously shorter than 
the petals; petals orbicular, 0.8-1 mm. long, tapering into a linear- 
lanceolate inflexed apex of approximately equal length; stylopodium 
much shorter than the slender, ascending or spreading styles; ped- 
icels slender, 3-5 mm. long, spreading; fruit orbicular, 1.5-2 mm. in 
diameter, sharply angled, the carpels slightly concave dorsally, the 
margins acute or narrowly winged. F.M. Neg. 3457. 

Illustrations: Math. & Const. Univ. Calif. Publ. Bot. 33: 166, 
/. 30. 1962. 

Ancash: Pr. Huaylas: Caraz, 2200-2500 m., Weberbauer 2993, 
type. Pr. Casma: Lomas de Lupin, entre Barranca y Huarmey, 
350-400 m., Ferreyra 13537, Mathias 3673. Pr. Huaraz: Casma 
Valley, 2000-3000 m., Rank & Hirsch 2113. Pr. Yungay: entre 
Carhuaz y Caraz, 2600-2200 m., Vargas 10299. Cajamarca: Pr. 
Jae"n: 900-2400 m., Vargas 10454; between Acapulco and Pucard, 
1150 m., Hutchison 1398; Puente del Rio Huancabamba, entre Pu- 
cara y Abra de Porculla, falda rocono con elementos subxerofilos, 
1200-1300 m., Ferreyra 13727; entre Abra de Porculla y Jae"n, 1300- 
1400 m., Ferreyra 13634. La Libertad: Pr. Otuzco: Laderas del 
Cerro Chologday, 2640 m., Angulo 0914; Plazapampa, 2200 m., Ce- 
vasco "D." Pr. Trujillo: Cerro Cabezon, 670 m., Angulo & Ldpez 
1480; El Portachuelo, Ascope-San Benito, 900 m., Ldpez 1277 
Lima: Pr. Canta: along Rio Chillon near Viscas, 1800-2000 m., 
Pennell 14481. Pr. Huarochiri: valley of Rio Rimac, near Lima- 
Oroya Highway at km. 60 east of Lima, 1500 m., Goodspeed & Met- 



UNIVERSITY Qf 
ILLINOIS LIBRARY 



52 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

calf 30227; between Matucana & San Bartolome", Rose & Rose 18755. 
Piura: Cerro Prieto, above 1200 ft., Haught F 155. Pr. Huanca- 
bamba: Carretera a Jae"n, 1200 m., Angulo 2103; Abra Porculla Pass, 
east side, 1350 m., Hutchison 1392. Pr. Paita: Cerro Prieto, La Brea, 
Amotape Hills, Talara, Haught 29; M