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Full text of "Flora of Peru"

ILL 

AT URBA .AMPAI 

BIOLOGY 

JUL 1 9 1QQ7 



FIELDIANA 
Botany 

Published by Field Museum of Natural History 



New Series, No. 7 



FLORA OF PERU 

J. FRANCIS MACBRIDE 
AND COLLABORATORS 



FAMILY COMPOSITAE: PART II 

TRIBE ANTHEMIDEAE 
MICHAEL O. DILLON 



UN! 



May 19, 1981 
Publication 1319 



FLORA OF PERU 

FAMILY COMPOSITAE: PART II 

TRIBE ANTHEMIDEAE 



FIELDIANA 
Botany 

Published by Field Museum of Natural History 



New Series, No. 7 



FLORA OF PERU 

J. FRANCIS MACBRIDE 
AND COLLABORATORS 



FAMILY COMPOSITAE: PART II 

TRIBE ANTHEMIDEAE 

MICHAEL O. DILLON 

Assistant Curator 

Department of Botany 

Field Museum of Natural History 



Accepted for publication December 9, 1980 
May 19, 1981 
Publication 1319 



Library of Congress Catalog Card No.: 80-66384 

ISSN 0015-0746 
PRINTED IN THE UNITED STATES OF AMERICA 



CONTENTS 

List of Illustrations vi 

Tribe Anthemideae 1 

Key to Genera of Anthemideae 1 

I. Achilla 2 

1. Achillea millefolium 2 

II. Artemisia 3 

Key to Species of Artemisia 3 

1. Artemisia absinthium 3 

2. Artemisia annua 4 

III. Chrysanthemum 4 

1. Chrysanthemum coronarium 5 

IV. Cotula 5 

Key to Species of Cotula 5 

1 . Cotula australis 6 

2. Cotula coronopifolia 6 

3. Cotula mexicana 7 

V. Dendranthema 9 

1. Dendranthema morifolium 9 

VI. Leucanthemum 10 

1. Leucanthemum vulgare 10 

VII. Matricaria 11 

1 . Matricaria recutita 12 

VIII. Plagiocheilus 12 

Key to Species of Plagiocheilus 13 

1. Plagiocheilus bogotensis 13 

2. Plagiocheilus frigidus 13 

3. Plagiocheilus solivaeformis 14 

IX. Santolina 15 

1. Santolina chamaecyparissus 15 

X. Saliva 16 

Key to Species of Saliva 16 

1. Saliva neglecta 16 

2. Saliva stolonifera 17 

XI. Tanacetum 18 

Key to Species of Tanacetum 18 

1. Tanacetum cinerariifolium 19 

2. Tanacetum parthenium 20 

3. Tanacetum vulgare 20 

Acknowledgments 20 

Index . ..21 



LIST OF ILLUSTRATIONS 

1. Cotula mexicana 8 

2. Plagiocheilus frigidus 14 

3. Saliva neglecta 17 



vi 



Tribe ANTHEMIDEAE 1 

Anthemideae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 192. 1819. TYPE: Anthemis 
L. 

Anthemidaceae Link, Handbuch 1: 752. 1929. 

Perennial to annual herbs, or more rarely shrubs or subshrubs, often aromatic. Leaves 
mostly alternate, rarely opposite, often dissected, lobed or toothed. Inflorescences cor- 
ymbose, racemose, paniculate, or solitary. Capitula usually heterogamous, radiate or dis- 
ciform, rarely homogamous, discoid; involucres cylindrical to globose; phyllaries 1-4-se- 
riate, imbricate, wholly scarious or herbaceous with scarious margins and apices; 
receptacles flat to convex or convex-conical, naked or rarely pubescent or with persistent 
or deciduous paleae; marginal florets usually pistillate, the corollas ligulate or rarely absent, 
homochromatic white, yellow, or red (occasionally pink) to heterochromatic white or cream 
distally and yellow proximally, entire or 1-3-denticulate; disc florets hermaphroditic or 
functionally male, the corollas tubular to funnelform, 4-5-merous (rarely 3-merous), usu- 
ally homochromatic yellow to occasionally deep red, the anthers basally obtuse or short- 
tailed, the terminal appendages deltoid to ovate, acute to rounded, the style branches 
truncate, terminally penicillate with the stigmatic surfaces in two, parallel, marginal stripes. 
Achenes di- or isomorphic, 2-10-ribbed or 1-3-winged, circular or angled-terete to strongly 
dorsiventrally or laterally compressed; pappus coroniform or occasionally absent. 

The Anthemideae is a large tribe consisting of some 101 genera and about 
1,400 species (Heywood & Humphries, 1977), occurring primarily in the tem- 
perate Northern Hemisphere. In Peru, many species are introduced weeds or 
escapees from cultivation; however, various members of Cotula, Saliva, and Pla- 
giocheilus are considered native to South America. 

REFERENCE 

HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 
wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 
Academic Press, London. 



1. Receptacles conspicuously paleaceous 2 2. 

2. Capitula discoid or disciform; outer florets with corollas regularly 5-lobed, 

yellow IX. Santolina. 

2. Capitula radiate; outer florets with corollas ligulate, white, yellow or 

pinkish l.Achillea. 

I. Receptacles epaleaceous (rudimentary paleae present in Tanacetum parthenium) 3. 

Assisted by National Science Foundation Grant DEB-79-05078 (Alwyn H. Gentry, prin- 
cipal investigator). 

^ultivars of Dendranthema morifolium often possess capitula lacking disc florets and 
having a proliferation of pistillate, ray florets, each subtended by a palea similar to an 
inner phyllary. 



2 FIELDIANA: BOTANY 

3. Capitula radiate; outer florets with corollas ligulate and exceeding the disc .... 4. 

4. Leaves pinnatifid 5. 

5. Involucres ca. 5 mm wide; receptacles conical and elongating with age 

VII. Matricaria. 

5. Involucres 10-20 mm wide; receptacles convex to planar 6. 

6. Achenes isomorphic, cylindrical or clavate, 3-10-ribbed . . XI. Tanacetum. 
6. Achenes dimorphic, the outer 3-angled, the inner cylindrical or 

subcylindrical III. Chrysanthemum. 

4. Leaves entire, dentate or deeply lobed, never pinnatifid 7. 

7. Capitula usually solitary; ray florets uniseriate VI. Leucanthemum. 

7. Capitula in lax corymbs; ray florets multiseriate V. Dendranthema. 

3. Capitula discoid or disciform; outer florets with corollas tubular, variously lobed or 

bilabiate (absent in Cotula), never conspicuously radiate 8. 

8. Outer or marginal florets with corollas bilabiate VIII. Plagiocheilus. 

8. Outer or marginal florets with corollas tubular or filiform, regularly 3-5-lobed 

(absent in Cotula) 9. 

9. Corollas of the disc florets regularly 5-dentate 10. 

10. Inflorescences compact spikes or panicles; capitula pendent II. Artemisia. 

10. Inflorescences corymbose; capitula erect XI. Tanacetum (T. vulgare). 

9. Corollas of the disc florets 3-4-lobed 11. 

11. Capitula sessile in leaf clusters X. Soliva. 

11. Capitula pedunculate IV. Cotula. 

I. ACHILLEA 
Achillea L., Sp. PL 899. 1753. TYPE: A. millefolium L. 

Perennial herbs. Leaves alternate, 3-4-pinnatifid to entire. Inflorescences corymbose or 
corymbiform-paniculate, rarely solitary, terminal. Capitula radiate, heterogamous, sub- 
sessile or pedunculate; involucres oblong-cylindrical, ovoid, hemispherical, or depressed; 
phyllaries 3 4-seriate, graduate, the margins scarious; receptacles conic or convex; paleae 
lanceolate or oblong, membranaceous; ray florets 2-15, pistillate, fertile, the ligules 3- 
dentate, white, yellow, or sometimes pinkish; disc florets 8-100, hermaphroditic, fertile, 
the corollas tubular, subcompressed, the bases pouched and enveloping tops of achenes, 
5-lobed distally, white, yellow, or pink, the anthers basally obtuse, the terminal appendages 
ovate, the style branches truncate, penicillate. Achenes oblong or obovate, compressed, 
glabrous, smooth; pappus absent. 

A genus of some 85 species from southeastern Europe and southwestern and 
central Asia. It is represented throughout Central and South America by the 
following cultivated and frequently escaping species. 

1. Achillea millefolium L., Sp. PI. 899. 1753. TYPE: Europe, exact locality and 
collector unknown (LINN, holotype, not seen, IDC Microfiche 117. 609: 
II. 7). 

Aromatic, perennial herbs from creeping rootstocks, to 0.5 m tall; stems erect, simple 
or branched, woolly-pilose. Leaves oblanceolate to oblong in outline, 2-3-pinnatifid, 4-10 
cm long, 0.5-1.5 cm wide, the ultimate segments linear to linear-lanceolate, 1-2 mm long, 
ca. 1 mm wide, spinulose-tipped, loosely pilose. Inflorescences corymbiform-paniculate, 
50-100-flowered; peduncles 1-5 mm long. Capitula 4-5 mm high, 2-3 mm wide (excluding 
rays); involucres oblong to ovoid; phyllaries 4-seriate, the outer ovate, 1.5-2.0 mm long, 
ca. 1 mm wide, sparsely woolly-pilose to subglabrous, the inner oblong to lanceolate, 
4.0-4.5 mm long, ca. 1 mm wide, glabrescent, all keeled, the margins scarious, brown, 
erose; ray florets usually 5, the ligules orbicular, 2.5-3.0 mm long, ca. 3 mm wide, white 
or pinkish white to ochroleucous; disc florets 10-25, the corollas 2.5-3.0 mm long, white 
to ochroleucous. Achenes obovate, 1.5-2.0 mm long, compressed, glabrous; pappus ab- 
sent. Chromosome number: n = 9 (x = 9). 3 

Chromosome numbers have been extracted from the following chromosome indexes: 
Fedorov, 1969; Moore, 1973, 1974, 1977. 



MACBRIDE: FLORA OF PERU 3 

CUZCO. Anta: Chaccan Chico, 3,490 m, Brunei 477 (F, MO); Santa Ana, El 
Chaccan, 3,492 m, Brunei 562 (MO), 636 (F, MO). 



II. ARTEMISIA 
Artemisia L., Sp. PI. 845. 1753. TYPE: A. vulgaris L. 

Annual, biennial, or perennial herbs or shrubs, aromatic, frequently tomentose. Leaves 
alternate, entire or pinnatifid, often pedate. Inflorescences racemose or paniculate, rarely 
solitary. Capitula homogamous or heterogamous, numerous, pendulous, or rarely erect; 
involucres oblong to depressed-globose, narrowly campanulate or hemispherical; phyl- 
laries imbricate, few-seriate, the margins scarious; receptacles convex or hemispherical, 
naked or conspicuously pilose; florets iso- or dimorphic, the outer pistillate, the corollas 
filiform, apically bi- or trifid; disc florets hermaphrodite or masculine, the corollas tubular, 
the limb campanulate, 5-dentate, the anthers obtuse or minutely pointed basally, the 
terminal appendage deltoid, sometimes subulate-acuminate, the style branches of female 
florets linear, truncate-penicillate, those of the hermaphrodite florets linear or linear-spat- 
ulate, usually penicillate, undivided in functionally male florets. Achenes subglobose to 
fusiform, terete or somewhat compressed, smooth, 2-ribbed or multistriate, incurved ba- 
sally; pappus absent. 

Approximately 400 species, primarily north temperate, including centers of 
diversity in Asia, Europe, and western North America. Its members tend to 
dominate the steppe communities of Asia, "sagebrush" communities of the New 
World, and the Karoo scrub of South Africa (Hey wood & Humphries, 1977). The 
genus is represented by two naturalized weeds in Peru. 

REFERENCE 

HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 
wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 
Academic Press, London. 

KEY TO SPECIES OF Artemisia 

1. Perennial; cauline leaves sericeous, the lobes oblong; receptacles pilose 

1. A. absinthium. 

1. Annual; cauline leaves glabrous, the lobes linear-lanceolate; receptacles naked 

2. A. annua. 

1. Artemisia absinthium L., Sp. PI. 848. 1753. TYPE: Europe (western edge of 
Asia, ?Russia), exact locality and collector unknown (LINN, holotype, not 
seen, IDC Microfiche 117. 567: I. 5). 

Suffrutescent, perennial herbs to 1 m tall, aromatic; stems erect, sericeous at least above. 
Leaves ovate in outline, 2-3-pinnatifid, 1.5-3.0 cm long, 1.5-2.5 cm wide, the ultimate 
segments oblong, 1.0-1.5 mm wide, subacute, grayish or whitish, sericeous on both sur- 
faces; petioles to 2 cm long. Inflorescences narrow to broad panicles, bracteate. Capitula 
disciform, heterogamous, ca. 3 mm high, 3-5(-6) mm wide, pedunculate, often nodding; 
involucres depressed-globose; phyllaries 2-3-seriate, graduate, the outer oblong, herba- 
ceous, 1-2 mm long, ca. 0.5 mm wide, sericeous, the inner obovate, mostly scarious, ca. 
2 mm long, ca. 1.5 mm wide, glandular; receptacles conspicuously pilose; florets 20-50, 
the outer pistillate, fertile, the corollas filiform, to 1 mm long, the inner florets herma- 
phroditic, fertile, the corollas cylindric-campanulate, 1.0-1.5 mm long, all yellowish, gla- 
brous. Achenes fusiform, glabrous, ca. 1 mm long. Chromosome number: n = 9. 

This species is native to Eurasia and North Africa and has been introduced 
into the New World. It is commonly cultivated for ornamental or medicinal 



4 FIELDIANA: BOTANY 

purposes. In Peru, it is known as "ajenjo/' a name also used for the genus 
Ambrosia. 

JUNIN. Jauja: Jauja, Cerrate 3810 (MO). 

2. Artemisia annua L., Sp. PI. 847. 1753. TYPE: Siberia, exact locality and col- 
lector unknown (LINN, holotype, not seen, IDC Microfiche 117. 566: II. 4). 

Erect annuals to 2 m tall; stems puberulous to glabrous, reddish. Leaves ovate in outline, 
2-3-pinnatifid, sessile, the basal segments 3-10 mm long, deeply toothed, remote from 
next distal pair, median segments 3-4 cm long, regularly and deeply toothed, the ultimate 
lobes linear-lanceolate, 1-5 mm long, 0.5-1.0 mm wide, glabrous. Inflorescences pani- 
culate. Capitula disciform, heterogamous, 1.5-3.0 mm high, 2.0-3.5 mm wide, pedun- 
culate, often nodding; involucres globose; phyllaries 2-3-seriate, graduate, the outer ob- 
long, herbaceous, ca. 0.6 mm long, the inner ovate-oblong, mostly scarious, ca. 2 mm 
long, 1.5-2.0 mm wide; receptacles naked; florets 20-25, the outer pistillate, fertile, the 
corollas filiform, to 1 mm long, the inner florets hermaphroditic, fertile, the corollas cy- 
lindric-campanulate, 0.6-1.0 mm long, all yellowish, glabrous. Achenes narrowly turbi- 
nate, ca. 0.8 mm long, obscurely striate. Chromosome number: n = 9. 

This species is a native of Asia and is widely naturalized in central and southern 
Europe and throughout the New World. In Peru, it is represented by cultivated 
material only. 

JUNIN. Jauja: Jauja, Ridoutt s.n. (MO). 

III. CHRYSANTHEMUM 
Chrysanthemum L., s.l., Sp. PI. 887. 1753. TYPE: C. coronarium L. 

Annual herbs, glabrous or pubescent, often strong scented. Leaves alternate, the mar- 
gins entire, toothed, incised or variously dissected, pubescent or glabrate. Inflorescences 
solitary or 2-5 on branch tips, often long pedunculate. Capitula radiate, heterogamous; 
involucres hemispherical or campanulate; phyllaries 3-4-seriate, imbricate, the margins 
scarious, the costa darkened; receptacles convex, epaleate; ray florets 13-21(-34), the ligules 
entire or dentate, pistillate, usually fertile, white or yellow, and rarely with reddish bases; 
disc florets 50-200, hermaphroditic, fertile, the corollas yellow, the tube laterally expanded 
and 2-winged, the anthers basally obtuse or truncate, the terminal appendage ovate, acute, 
the style branches narrowly oblong, truncate, penicillate. Achenes without vallecular se- 
cretory canals or epicarpic mucilaginous cells, those of the ray florets 3-angled, with the 
ribs often winged, those of the disc florets cylindrical to cylindrical-triquetrous, ribbed, 
the adaxial rib sometimes winged; pappus absent. 

The genus Chrysanthemum, when interpreted in a broad sense, includes about 
200 species, mainly natives of Europe, western Asia, and northern Africa, with 
species naturalized on nearly every continent. Considerable biosystematic evi- 
dence indicates that this genus has been quite heterogeneous, and recent work- 
ers (Heywood & Humphries, 1977) have recircumscribed the generic limits, thus 
limiting the genus to a group of three species: C. carinatum, C. coronarium, and 
C. segetum, all of northern Africa and Europe. These changes have been reflected 
in several recent floras (e.g., Flora of Turkey, 1975; Flora Europaea, 1976). In the 
present treatment, several taxa traditionally treated under Chrysanthemum s.l. 
are treated under other genera (cf. Tanacetum, Dendranthema, and Leucanthemum). 

REFERENCES 

GRIERSON, A. J. C. 1975. Chrysanthemum. In Davis, P. H. (ed.), Flora of Turkey. 5: 253-255. 
HEYWOOD, V. H. 1976. Chrysanthemum. In Tutin, T. G., et al. (eds.), Flora Europaea. 4: 
168-169. 



MACBRIDE: FLORA OF PERU 5 

HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 
wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 
Academic Press, London. 

1. Chrysanthemum coronarium L., Sp. PI. 890. 1753. TYPE: Europe, exact lo- 
cality and collector unknown (LINN, holotype, not seen, IDC Microfiche 117. 
603: II. 3). 

Annual herbs to 0.75 m tall; stems erect, glabrous. Leaves oblong to obovate in outline, 
2-3-pinnatifid, to 5 cm long, 2-3 cm wide, sessile, the ultimate segments 1-2 mm wide, 
acute. Inflorescences solitary; peduncles 5-15 cm long, ebractate. Capitula ca. 1 cm high, 
1.0-1.5 cm wide (excluding rays); involucres hemispherical; phyllaries 3-seriate, ovate- 
oblong, 5-10 mm long, 3-5 mm wide, the margins scarious, brownish; ray florets ca. 21(- 
34), the ligules 1.0-1.5 cm long, ca. 5 mm wide, yellow, cream, or white; disc florets 
50-100, the corollas 4-5 mm long, yellow. Achenes dimorphic, the outer triquetrous, the 
angles produced into wings to 1.5 mm wide, the inner laterally compressed with adaxial 
wings, prominent ribs on abaxial face and rounded ribs on lateral faces, sometimes those 
of the center lacking wings, all covered with sessile, non-mucilaginous glands. Chro- 
mosome number: n = 9. 

This species is native to the Mediterranean region and is now a widespread 
weed in many temperate areas. In Peru, it is cultivated for ornament and often 
escapes to roadsides and waste places. 

JUNIN. Jauja: Jauja, Ridoutt s.n. (MO). CUZCO. Anta: Chaccan Chico, El 
Chaccan, 3,490 m, Brunei 484 (MO). 

IV. COTULA 

Cotula L., Sp. PI. 891. 1753. TYPE: C. coronopifolia L. 

Annual or perennial herbs, often diminutive, prostrate to decumbent, often rhizomatous 
or stoloniferous^ glabrous to pilose, often with pellucid glands. Leaves alternate, 2-3- 
pinnatifid, rarely simple, toothed to entire, periolate or not, the bases amplexicaul or only 
partly so. Inflorescences solitary, terminal and axillary; peduncles sometimes swollen 
under the capitula. Capitula monoecious, dioecious, or gynomonoecious, disciform, het- 
erogamous; involucre hemispherical to campanulate; phyllaries 2-many-seriate, subequal, 
herbaceous, scarious at least marginally; receptacles conical, flat or convex, rarely hemi- 
spherical; epaleate; marginal florets pistillate, 1-many-seriate, the corollas filiform, some- 
times 2-toothed or with a minute ligula (sometimes corollas lacking); disc florets herma- 
phroditic, fertile, or functionally male, the corollas cylindric-campanulate, funnelform, or 
tubular, sometimes with the bases sheathing and extending over the ovary, 4-lobed (rarely 
3-lobed), the anthers 4 (rarely 3), basally obtuse or minutely tailed, the terminal appendage 
ovate or lanceolate, the style branches of the marginal florets linear-lanceolate, of the disc 
florets oblong, truncate, penicillate. Achenes generally stipitate, terete, or dorsally com- 
pressed, winged or not, dorsally convex; pappus absent. 

About 90 species, almost cosmopolitan, but mainly South African and in New 
Zealand, some in North America, Asia, Australia, New Guinea, South America, 
and the Falkland Islands. 

REFERENCE 

CARD, J. A. 1961. Las especies de Cotula (Compositae) del centro de la Republica Argentina. 
Kurtziana 1: 289-298. 

KEY TO SPECIES OF Cotula 
1. Receptacle pilose; disc corollas 3-lobed, stamens 3; achenes isomorphic 



3. C. mexicana. 



6 FIELDIANA: BOTANY 

1. Receptacle glabrous; disc corollas 4-lobed, stamens 4; achenes dimorphic 2. 

2. Plants annual, pubescent; marginal florets in 3 series; disc corollas white 

1. C. australis. 

2. Plants perennial, glabrous; marginal florets in 1 series; disc corollas yellow 

2. C. coronopifolia. 

1. Cotula australis (Sieb. ex Spreng.) Hook, f., Fl. Nov. Zel. 128. 1853. 

Anacyclus australis Sieb. ex Spreng., Syst. Veg. 3: 497. 1826. TYPE: New Zealand ("Nov. 
Holl."), exact locality and collector unknown (?P [Herb. Cosson], holotype, not seen). 

Annual herbs to 20 cm tall, basally branched, thinly villous. Leaves 1-2-pinnatifid, 
oblong in outline, to 6 cm long, to 2 cm wide, the lobes linear-lanceolate, acute, often 
glabrous above, villous below, sessile, the bases dilated, subamplexicaul. Inflorescences 
solitary, terminating branches; peduncles to 8 cm long, villous with appressed, ascending 
trichomes. Capitula monoecious, disciform, heterogamous, ca. 2 mm high, 4-5 mm wide; 
involucres hemispherical; phyllaries 2-seriate, equal, oblong, 1.5-2.0 mm long, 0.6-1.0 mm 
wide, apically obtuse or rounded, the margins scarious; marginal florets stipitate, pistillate, 
the corollas absent; disc florets hermaphroditic, the corollas tubular, 0.6-0.8 mm long, 4- 
lobed, white. Achenes dimorphic, the outer oblong, ca. 1 mm long, dorsally compressed, 
laterally winged, glandular, the inner ones obconical, 0.5-0.8 mm long, flattened on the 
inner face, convex on the outer face, the margins thickened. Chromosome number: n = 
10. 

This species was originally described from New Zealand and is naturalized 
in many regions of the world. It is widespread in Peru, occupying a wide range 
of habitats. 

AMAZONAS. Chachapoyas: ca. 43 km NE of Balsas on road to Chachapoyas, 
ca. 2,900 m, Dillon & Turner 1720 (F, USM). AREQUIPA. Caraveli: Lomas de 
Ocopa, entre Uauca y Atiquipa, 500-600 m, Ferreyra 11478 (MO); Islay: Mollendo, 
1,500-2,000 m, Ferreyra 18658 (MO), Lomas de Mollendo, 560 m, Herrera-Rivera 
40 (MO), 7 km de Matarani, 800 m, Lopez 6 (MO). CAJAMARCA. Cajamarca: 
Sunchubaba, 2,410 m, Cabanillas et al. 7 (MO); Celendin: ca 23 km SW of Celendin, 
3,100 m, Dillon & Turner 1645 (F, MO, TEX, USM). JUNIN. Tarma: Paucartambo, 
2,800 m, Woytkowski 6694 (F, MO). LA LIBERTAD. Trujillo: Barraza, 50 m, Lopez 
& Sagastegui 7990 (MO). LIMA. Canta: Rucuma, entre Canta y Obrajillo, 2,700 
m, Mexa 159 (MO), San Buenaventura, 2,700-2,800 m, Penned 14570 (F), Puru- 
marca, cerca a Canta, Sanchez 52 (MO); Chancay: Lomas de Lachay, km 88 
carretera Panamericana Norte, 400 m, Cerrate 5738 (MO), Naupay, 2,700 m, 
Cerrate et al. 6315 (MO), Lomas de Chancay, 300 m, Ferreyra 8744 (MO), Lachay, 
entre Chancay y Huacho, 380-400 m, Ferreyra 8765 (MO); Huarochiri: Viso, 2,740 
m, Macbride & Featherstone 637 (F); Lima: Lomas de Lurin, 350-400 m, Ferreyra 
11920 (MO), Loma de Amancae, 120-410 m, Gentry 16479 (F, MO). PIURA. 
Huancabamba: 40-43 km E of Olmos on road to Pucara, 1,600-1,850 m, Gentry 
et al. 22664 (F, MO, USM). 

2. Cotula coronopifolia L., Sp. PI. 892. 1753. TYPE: Ethiopia, exact locality and 
collector unknown (LINN, holotype, not seen, IDC Microfiche 117. 605: I. 4). 

Perennial herbs to 20 cm tall; stems stoloniferous, glabrous, usually much-branched, 
ascending. Leaves linear to lanceolate, entire to deeply laciniate, or 1-2-pinnatifid, 2-5 cm 
long, 0.5-1.5 cm wide, glabrous, the bases sessile and sheathing. Inflorescences solitary, 
terminal; peduncles to 6 cm long, glabrous. Capitula monoecious, disciform, heteroga- 
mous, 2-3 mm high, 9-10 mm wide; involucres hemispherical; phyllaries 3-seriate, equal, 
oblong-lanceolate, 3-5 mm long, 1-2 mm wide, apically obtuse, glabrous, striate, the 



MACBRIDE: FLORA OF PERU 

margins scarious; marginal florets uniseriate, stipitate, pistillate, the corollas absent; disc 
florets hermaphroditic, the corollas tubular, 1.2-1.5 mm long, 4-lobed, yellow. Achenes 
dimorphic, the outer oblong, 1.3-1.8 mm long, dorsally compressed, laterally winged, 
glandular, the inner ones obconical, 1.0-1.3 mm long, flattened on the inner face, convex 
on the outer face, the margins thickened. Chromosome number: n = 10. 

This species was originally described from Africa and is adventive throughout 
the world. It frequently inhabits aquatic or inundated areas in fresh- and salt- 
water. 

AREQUIPA. Arequipa: alrededores de los Barios de Jesus, cerca de Arequipa, 
2,400-2,600 m, Ferreyra 12047 (MO), Yura, ca. 2,700 m, Solomon 2826 (MO), 
Chilean Valley, Stafford 566 (F), no exact locality, Vargas 2030 (MO); Islay: cerca 
a Mollendo, 300 m, Ferreyra 12098 (MO). PUNO. Puno: near Puno, 4,000 m, 
Soukup 410 (F). 

3. Cotula mexicana (DC.) Cabrera, Bol. Soc. Argent. Bot. 8: 207. 1960. 

Hippia minuta L. f., Suppl. 389. 1781. TYPE: LINN, not seen. 

Saliva pedicellata Ruiz & Pavon, Syst. Veg. 215. 1798. TYPE: Peru, exact locality unknown, 

Ruiz & Pavon s.n. (MA, not seen). 
Saliva pygmaea H.B.K., Nov. Gen. Sp. PI. 4: 238. ed. fol. 1818. TYPE: Colombia, Tolima, 

"Andium Quindeunsim," Humboldt & Bonpland s.n. (P, holotype, not seen, IDC Mi- 
crofiche 6209. 113: II. 4). 
Gymnostyles minuta (L. f.) Spreng., Syst. Veg. 3: 500. 1826. TYPE: based upon Hippia 

minuta L. f. 
Gymnostyles peruviana Spreng., Syst. Veg. 3: 500. 1826. nom. superfl. (Based upon Saliva 

pedicellata Ruiz & Pavon.) 

Saliva minuta (L. f.) Sweet, Hort. Brit. ed. 2. 217. 1830. 
Saliva mexicana DC., Prodr. 6: 143. 1837. TYPE: Mexico, no exact locality, Sesse & Mocino 

3959 (G, holotype illustration; F, isotype specimen). 

Soliva pedunculata Ruiz & Pavon ex Steudel, Nom. Bot. ed. 2, 2: 609. 1841. orth. mut. 
Cotula pygmaea (H. B. K.) Benth. & Hook, ex Hemsl., Biol. Centr. Amer. Bot. 2: 230. 

1881 (non C. pygmea Poir., Encyc. Suppl. 2: 371, 1810). 
Cotula minuta (L. f.) Schinz, Mem. Soc. Sci. Nat. Neuchatel 5: 429. 1913 (non C. minuta 

Forster, Prodr. 57. 1786). 

Lancisia minuta (L. f.) Rydb., N. Amer. Flora 43: 287. 1916. 
Cotula pedicellata (Ruiz & Pavon) Cabrera, Notas Mus. La Plata, Bot., 14: 139. 1949 (non 

C. pedicellata Compton, J. S. African Bot. 7: 189. 1941). 
Cotula cabrera Caro, Kurtziana 1: 295. 1961. nom. superfl. (Based upon Soliva pedicellata 

Ruiz & Pavon.) 

Annual, diminutive herbs to 3 cm tall; stems ascending to procumbent, pubescent. 
Leaves pinnatifid, l-2(-3) cm long (including petiole), 5-15 mm wide, oblanceolate in 
outline, the segments oblanceolate to elliptic, 4-paired, 2-5 mm long, 0.6-0.9 mm wide, 
apically obtuse to subacute, mucronate, entire, glabrous to sparsely villous, villous below; 
petioles 5-10 mm long, villous, basally dilated, subamplexicaul. Inflorescences solitary; 
peduncles axillary, to 1 cm long, villous. Capitula monoecious, disciform, heterogamous, 
1.5-2.0 mm high, ca. 2 mm wide; involucres campanulate; phyllaries 2-seriate, equal, 
oblong, 1.0-1.2 mm long, 0.4-0.5 mm wide, sparsely villous, apically obtuse to acute, 
roseate, the margins scarious; receptacles flat, pilose; marginal florets 2-seriate, stipitate, 
pistillate, fertile, ca. 10, the corollas absent, the stipe ca. 0.2 mm long, the ovary obovate 
to oblong, compressed; disc florets ca. 3, hermaphroditic, functionally male, the corollas 
tubular, ca. 1 mm long, 3-lobed, white. Achenes isomorphic, obovate, 1.3-1.5 mm long, 
0.8-1.0 mm wide, dorsally compressed, laterally 2-winged, smooth, glabrous. Chromo- 
some number: n = ca. 56. 

This species occupies humid, montane habitats from Mexico to Costa Rica and 
from Colombia to Argentina. In Peru, it is only known from a few collections 
from Depto. Amazonas in the north and Depto. Puno in the south. 




u 



MACBRIDE: FLORA OF PERU 9 

The earliest legitimate specific epithet for this taxon is mexicana; the epithets 
minuta, pygmaea, and pedicellata are all illegitimate due to homonymy. 

AMAZONAS. Chachapoyas: middle eastern Calla-Calla slopes, near km 
411^16 of Leimebamba-Balsas road, 3,100-3,250 m, Wurdack 1331 (F). PUNO. 
Carabaya: Crucero Alto, 4,540 m, Stafford 654 (F); Azangaro: Chuquibambilla, 
3,850-3,900 m, Pennell 13391 (F). 

V. DENDRANTHEMA 

Dendranthema (DC.) Des Moul., Actes Soc. Linn. Bordeaux 20: 561. 1860. TYPE: 
D. indicum (L.) Des Moul. = Chrysanthemum indicum L. 

Perennial herbs, or rarely suffruticose. Leaves alternate, oblanceolate to ovate, lobed to 
2-pinnatifid (rarely entire), petiolate. Inflorescences solitary to corymbose. Capitula ra- 
diate, heterogamous; involucres hemispherical; phyllaries ca. 3-seriate, imbricate, mar- 
ginally scarious, brown, erose; receptacles convex to conical, epaleate (paleae present in 
multi-seriate rayed cultivars); ray florets pistillate, fertile or sterile, the ligules white to 
purple or yellow (variously colored in cultivars); disc florets hermaphroditic (absent in 
some cultivars), the corollas tubular-obconical, usually yellow, the tube unwinged, apically 
5-lobed, the anthers basally obtuse, the terminal appendage lanceolate to ovate, the style 
branches oblong, truncate, penicillate. Achenes isomorphic, cylindrical-obconical, 5-8- 
ribbed, without vallecular secretory canals, with or without epicarpic mucilaginous cells; 
pappus absent. Chromosome number: n = 9 (x 9). 

The genus consists of some 50 species of perennial herbs and subshrubs 
distributed throughout the Far East, including China, Japan, Korea, Mongolia, 
USSR, and Europe. The genus is of considerable horticultural interest, with 
some 7,000 cultivars of autumn-flowering chrysanthemums derived from Den- 
dranthema morifolium, the hybrid product of various native species (Heywood 
& Humphries, 1977). 

REFERENCES 

BAILEY, L. H. 1976. Hortus Third: A concise dictionary of plants cultivated in the United 
States & Canada. Initially compiled by L. H. Bailey and E. Z. Bailey; revised and ex- 
panded by the staff of Liberty Hyde Bailey Hortorium. Macmillan Publishing Co., Inc., 
New York, N.Y. 

GOSLING, S. G. (ED.). 1970. The Chrysanthemum Manual, National Chrysanthemum So- 
ciety. Essex Telegraph Press Ltd., Colchester, England. 375 pp. 

HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 
wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 
Academic Press, London. 

1. Dendranthema morifolium (Ramat.) Tzvel. Flora U.R.S.S. 26: 373. 1961. 

Chrysanthemum morifolium Ramat., J. Hist. Nat. 2: 240. 1792. TYPE: presumably P, not 
seen. 

Suffrutescent perennial herbs to 1.5 m tall, aromatic; stems erect or spreading, leafy. 
Leaves lanceolate to ovate, 4-9(-12) cm long, 4-6 cm wide, lobed, the segments entire to 
coarsely toothed, the upper surface glabrous, the lower pilose with 2-armed trichomes, 
glandular; petioles to 4 cm long, 2 auriculate segments basally. Inflorescences laxly cor- 
ymbose; peduncles bracteate. Capitula radiate; involucres hemispherical; phyllaries her- 
baceous, the outer lanceolate to oblong, 5-8 mm long, 1-2 mm wide, the inner ovate, 8-10 
mm long, 2-3 mm wide, the margins broadly scarious; receptacles strongly convex, paleate; 
ray florets multi-seriate, numerous, pistillate, the ligules 1-8 cm long, variously colored 
(commonly purple or yellow); disc florets absent in ours. Achenes 1.0-1.5 mm long, sterile. 

Autumn-flowering perennial chrysanthemums were successfully introduced 



10 FIELDIANA: BOTANY 

from China to France in 1789, and in 1792 the French botanist Ramatuelle pro- 
posed the name Chrysanthemum morifolium ( = Dendranthema morifolium) for one 
of these cultivars called "Old Purple" (Gosling, 1970). Although its origin is 
obscure, this cultivar is thought to be of hybrid origin, derived from several 
native Far Eastern species, including D. indicum, D. japonense, D. makinai, and 
D. ornatum (Bailey, 1976). It is not known when it was first introduced into the 
New World. 

The specimens examined thus far from Peru lack disc florets, with the ray 
florets produced in multiple series. Each ray floret is subtended by a palea, 
closely resembling an inner phyllary. These cultivars are sterile and are prop- 
agated through cuttings. In Peru, they are called "crisanthema." 

CUZCO. Anta: Chaccan Chico, El Chaccan, 3,493 m, Brunei 734 (MO). LO- 
RETO. Alto Amazonas: Yurimaguas, lower Rio Huallaga, 155-210 m, Williams 
4332 (F), 4517 (MO); Maynas: Iquitos and vicinity, 120 m, Williams 3607 (F), 8212 
(MO). SAN MARTIN. San Martin: Tarapoto, 750 m, Williams 5931 (F), Alto Rio 
Huallaga, 360-900 m, Williams 6607 (F). 

VI. LEUCANTHEMUM 
Leucanthemum Mill., Card. Diet. Abridg. ed. 4. 1754. TYPE: L. vulgare Lam. 

Perennial, rarely annual herbs. Leaves alternate, oblong, spatulate or obovate, the mar- 
gins crenate, serrate or pinnatifid. Inflorescences solitary or rarely lax corymbs. Capitula 
radiate, heterogamous, rarely disciform or discoid; involucres hemispherical; phyllaries 
3-4-seriate, imbricate, lanceolate to oblong, the margins scarious; receptacles usually con- 
vex, epaleate; marginal florets usually ligulate, pistillate, fertile, the corollas white or 
pinkish, rarely yellow, rarely tubular-campanulate, hermaphroditic or pistillate, yellow; 
disc florets hermaphroditic, fertile, the corollas tubular, 5-lobed, yellow, the anthers basally 
obtuse, the terminal appendage ovate, the style branches truncate, penicillate. Achenes 
isomorphic, oblong-cylindrical, 10-ribbed, the ribs bearing myxogenic cells, vallecular se- 
cretory canals present; pappus a corona or auricle, sometimes rudimentary or only on 
marginal achenes, or absent. 

A genus of approximately 25 species native to Europe, North Africa, and 
southwest Asia. In Peru, the genus is represented by the cultivated taxon, Leu- 
canthemum vulgare (= Chrysanthemum leucanthemum L.), which occasionally es- 
capes and appears naturalized. 

REFERENCES 

BOCHER, T. W., AND K. LARSEN. 1957. Cytotaxonomical studies in the Chrysanthemum 

leucanthemum complex. Watsonia 4: 11-16. 
HEYWOOD, V. H. 1976. Leucanthemum. In Turin, T. G., et al. (eds.), Flora Europaea. 4: 

174-177. 
HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 

wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 

Academic Press, London. 

1. Leucanthemum vulgare Lam., Fl. Francoise 2: 137. 1778. TYPE: based upon 
Chrysanthemum leucanthemum L. 

Chrysanthemum leucanthemum L., Sp. PI. 888. 1753. LECTOTYPE (designated by Bocher 
& Larsen, 1957): Europe, exact locality and collector unknown, Herb. Cliff. (BM, not 
seen). 

Perennial herbs to 1 m tall, aromatic; stems simple or sparsely branched, glabrous. Basal 
leaves obovate-spatulate to oblong-obovate, to 15 cm long, incised to crenate, petiolate; 



MACBRIDE: FLORA OF PERU 11 

cauline leaves oblong to oblong-lanceolate, to 10 cm long, remotely denticulate or entire, 
sessile and somewhat clasping at the base, glabrous. Inflorescences solitary. Capitula 
radiate, heterogamous, ca. 1 cm high, 1.5-2.0 cm wide (excluding rays); phyllaries lan- 
ceolate to oblong, the outer 6-9 mm long, 2.0-2.5 mm wide, the inner ca. 10 mm long, 
2.5-3.0 mm wide, the margins scarious, brownish, erose; receptacles convex; ray florets 
21-34, the ligules oblong, 2.5-3.0 cm long, ca. 5 mm wide, apically denticulate, the tube 
1-2 mm long, white; disc florets 200-300, the corollas tubular, 3-4 mm long, yellow. 
Achenes 2.0-2.5 mm long, blackish with prominent, pale-colored ribs, glabrous; pappus 
a corona or short unilateral auricle, or absent. Chromosome number: n = 9 (18, 27, 36, 
45, 54). 

This species, or species-complex, is native to temperate Eurasia from western 
Europe to China and naturalized throughout North and South America. It is 
extremely variable within its range in the Old World and has been divided into 
a large number of questionable species (Heywood, 1976). Individuals occurring 
in Peru have been previously referred to Leucanthemum maximum (Ramond) DC., 
which is a commonly cultivated, cytological variant (n = 45 or 54). Although 
the plants within our range do possess larger cauline and basal leaves and larger 
capitula, current taxonomic treatments view it simply as a variant of L. vulgare 
(Heywood, 1976). 

Its common name in Peru is "marguarita." 

CUZCO. Anta: El Chaccan, 3,550 m, Brunei 268 (F, MO). 



VII. MATRICARIA 

Matricaria L., Sp. PI. 890. 1753. LECTOTYPE (as designated by Pobedimova, 

1961): Matricaria recutita L. 

Annual herbs, strongly aromatic or odorless. Leaves alternate, variously pinnatifid, the 
ultimate segments linear or filiform. Inflorescences solitary or sometimes corymbose. Ca- 
pitula radiate, heterogamous, rarely discoid and homogamous; involucres campanulate 
to hemispherical; phyllaries 2-3-seriate, equal to subequal, the margins membranous or 
scarious; receptacles conical, elongating in fruit, epaleate; ray florets (when present) pis- 
tillate, fertile or sterile, the ligules white, tridentate; disc florets hermaphroditic, the corollas 
tubular to cylindric-campanulate, 5-lobed, the anthers basally obtuse, the terminal ap- 
pendage deltoid, acute, the style branches oblong, truncate, penicillate. Achenes iso- 
morphic, oblong to obovate, often incurved, strongly 3-5-ribbed, glabrous; pappus absent 
or minutely coroniform. 

This genus, when interpreted in the narrowest sense, consists of about five 
species of northern hemispheric annuals and represents a Mediterranean di- 
vergence from the closely related genus Anthemis L. (Heywood & Humphries, 
1977). Matricaria also shares relationships with Tripleurospermum Sch. Bip., a 
distinct genus of some 30 species of Europe, Asia, and North Africa. 

REFERENCES 

GRIERSON, A. J. C. 1974. Matricaria. In Materials for a Flora of Turkey. XXX. Compositae. 
Notes Roy. Bot. Card. Edinburgh 33: 252-254. 

HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 
wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 
Academic Press, London. 

JEFFREY, C. 1979. Note on the lectotypification of the names Cacalia L., Matricaria L. and 
Gnaphalium L. Taxon 28: 349-351. 

POBEDIMOVA, E. G. 1961. Matricaria. In Shishkin, B. K., and Bobtov, E. G., Flora U.R.S.S. 
26: 147-152, Moscow-Leningrad 



12 FIELDIANA: BOTANY 

1. Matricaria recutita 4 L., Sp. PI. 891. 1753. TYPE: Europe, exact locality and 
collector unknown (LINN or BM, not seen). 

Matricaria courrantiana DC, Prodr. 6: 52. 1837. TYPE: ex Teneriffa, Courrant s.n. (G-DC, 
holotype, not seen, IDC Microfiche 800. 1002: HI. 2). 

Annual herbs to 45 cm tall, aromatic; stems erect, branched, striate, glabrous. Leaves 
oblong in outline, 5-7 cm long, 2-3-pinnatifid, the primary segments 10-12-paired, nar- 
rowly linear. Inflorescences solitary or rarely laxly corymbose; peduncles 1-9 cm long. 
Capitula radiate, heterogamous, 5-6 mm high, 5-6 mm wide (excluding rays); involucres 
hemispherical; phyllaries 2-seriate, equal, lanceolate to oblanceolate, 2.5-3.0 mm long, 
0.3-0.8 mm wide, apically acute to obtuse, the margins membranous, erose, brown; re- 
ceptacles conical, hollow, epaleate; ray florets 12-15, pistillate, fertile, the ligules oblong, 
5-8 mm long, 2-3 mm wide, tridentate, white; disc florets 200-400, the corollas tubular, 
1.2-1.5 mm long, yellow. Achenes obovate, 0.6-0.9 mm long, smooth dorsally, the inner 
surface 3-5-ribbed, glabrous, apically truncate; pappus absent, or a membranous corona, 
fimbriate, white, 0.3-0.9 mm long (disc), 1.3-1.9 mm long (rays). Chromosome number: 
n = 9. 

This taxon has its origin in Eurasia and is adventive in the New World. It is 
commonly found in markets of Central and South America, where it is sold for 
preparation of tea used for stomach troubles. In Peru, it is commonly cultivated 
and frequently escapes, becoming naturalized in fields and other disturbed areas. 
It is called "manzanilla" throughout Central and South America. 

CUZCO. Anta: Cillapuya, El Chaccan, 3,613 m, Brunei 302 (F, MO); Cuzco: 
2 km S of San Jeranimo, 10 km SE of Cuzco, Solomon 3009 (MO). LIMA. Huar- 
ochiri: Matucana, Macbride & Featherstone 458 (F). 



VIII. PLAGIOCHEILUS 

Plagiocheilus Arnott ex DC., Prodr. 6: 142. 1837. TYPE: P. tanacetoides Haenke 
ex DC. 

Hippia H.B.K., Nov. Gen. Sp. PI. 4: 301. 1820, not Hippia L., Mant. PI. Alt.: 158, 291. 
1771. TYPE: H. peduncularis H.B.K. = P. peduncularis (H.B.K.) Wedd. 

Annual or perennial herbs. Leaves alternate, 1-2-pinnatifid. Inflorescences solitary or 
corymbose. Capitula disciform, heterogamous; involucres hemispherical; phyllaries 2-3- 
seriate; receptacles convex to conical; marginal florets multiseriate, pistillate, the ligules 
bilabiate, the outer entire or bidentate, the inner short, entire or bidentate, white; disc 
florets hermaphroditic, functionally male, the corollas funnelform, yellow, (4-)5-lobed, the 
anthers basally obtuse, the terminal appendages ovate, the style branches of marginal 
florets long, filiform, of the disc florets short, truncate. Achenes obconical, subcompressed 
laterally, glandular, puberulent or glabrous; pappus absent. 

A genus of perhaps five species from Colombia to Argentina. In Peru, three 
species are known from various high-elevation, moist habitats. 

The tribal status of this genus is open to question. Grau (1977) and Robinson 
& Brettell (1973) have suggested that it belongs in the Astereae. However, it is 
here accepted in the Anthemideae, a position supported by its general mor- 
phology, breeding system, and floral biology (Heywood & Humphries, 1977; 
Turner, pers. comm.). Its closest affinities appear to be with other southern 
hemispheric genera, including Cotula, Centipeda, Soliva, and Abrotanella. 

4 Other synonyms are listed by Grierson (1974). Only the names listed have been used 
for Peruvian material. 



MACBRIDE: FLORA OF PERU 13 

REFERENCES 

CUATRECASAS, J. 1954. Notas a la Flora de Colombia. XIII. Revista Acad. Colomb. Ci. Exact. 
9: 233-249. 

GRAU, J. 1977. Astereae Systematic review. In Heywood, V. H., et al. (eds.), The Biology 
and Chemistry of the Compositae, pp. 539-565. Academic Press, London. 

HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 
wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 
Academic Press, London. 

ROBINSON, H., AND R. D. BRETTELL. 1973. Tribal revisions in the Asteraceae. X. The 
relationship of Plagiocheilus. Phytologia 26: 159-162. 

KEY TO SPECIES OF Plagiocheilus 

1. Inflorescences corymbose 1. P. bogotensis. 

1. Inflorescences solitary 2. 

2. Leaves 2-pinnatifid, pilose with flaccid, uniseriate trichomes ... 3. P. solivaeformis. 

2. Leaves 1-pinnatifid, arachnoid-tomentose 2. P. frigidus. 

1. Plagiocheilus bogotensis (H.B.K.) Wedd., Chlor. And. 1: 62. 1865. 

Hippia bogotensis H.B.K., Nov. Gen. Sp. PI. 4: 237. ed. fol. 1818. TYPE: Colombia, Cun- 
dinamarca, "prope Santa Fe de Bogota," Humboldt & Bonpland s.n. (P, holotype, not 
seen, IDC Microfiche 6209. 113: II. 2). 

Leptinella bogotensis (H.B.K.) DC., Prodr. 6: 142. 1837. 

Plagiocheilus prostratus Benth., PI. Hartw. 136. 1844. TYPE: Ecuador, Loja, "in montibus 
Loxa," Hartweg s.n. (K, holotype, not seen; P, isorype, not seen; F, photo). 

Annual or perennial herbs to 0.3 m tall; stems decumbent to ascending, arachnoid to 
glabrate. Basal leaves obovate-spatulate to oblong-obovate in outline, to 14 cm long (in- 
cluding petiole), 2-pinnatifid, the ultimate segments lanceolate, sparsely pilose; petioles 
to 5 cm long, basally dilated, clasping; cauline leaves ovate to obovate in outline, 2.0-3.5 
cm long, 1.5-2.0 cm wide, 2-pinnatifid, the segments lanceolate, pilose, sessile, the bases 
dilated, incised, clasping. Inflorescences corymbose. Capitula 2-3 mm high, 4-5 mm wide; 
phyllaries 2-seriate, equal, broadly lanceolate to ovate, ca. 1.5 mm long, ca. 1 mm wide, 
apically obtuse, the margins scarious, nerved, persistent; marginal florets multiseriate, 
pistillate, the ligules white, bilabiate, the outer narrowly spatulate, ca. 0.5 mm long, the 
inner minute; disc florets 30-50, the corollas funnelform, ca. 1.5 mm long, 5-lobed, yellow. 
Achenes obconical, subcompressed laterally, ca. 3 mm long, puberulent, glandular, whit- 
ish; pappus absent. 

Plagiocheilus bogotensis occupies moist, montane habitats from Colombia to 
northern Peru (2,100-3,100 m). It most closely resembles P. tanacetoides Haenke, 
a lowland, annual species of northeastern Argentina and is distinguished from 
that species by its more deeply incised and generally less pubescent leaf seg- 
ments. 

Bentham's Plagiocheilus prostrata, described from Ecuador, appears to be a 
variation in habit and is here reduced to synonymy. 

AMAZONAS. Luya: Leimebamba, 2,100 m, Woytkowski 7802 (F, MO). CA- 
JAMARCA. Cajamarca: Pampa de la Culebra, around la Encanada, 2,900 m, 
Sagastegui et al. 8076 (F, MO). LA LIBERTAD. Huamachuco: mountain slopes 
above Yamobamba, 3,000-3,100 m, Conrad 2726 (F, MO); Santiago de Chuco: 
above Cachicadan, 2,800 m, Stork & Morton 9977 (F, MO). 

2. Plagiocheilus frigidus Poepp. & Endlich., Nov. Gen. Sp. PI. 3: 48, t. 248B. 
1843. TYPE: Peru, Pasco, "circum Cerro de Pasco," Poeppig s.n. (W, holotype, 
not seen). 



14 



FIELDIANA: BOTANY 




FIG. 2. Plagiocheilus frigidus. (From Gentry et al. 19226, MO.) 



Perennial, suffruticose herbs, caespitose; stems stoloniform. Leaves oblong-spatulate in 
outline, 8-25 mm long (including petiole), ca. 5 mm wide, pinnatifid, the segments entire 
or 1-lobed, lanceolate, 1.5-2.5 mm long, 0.6-1.0 mm wide, apiculate, glabrous to sparsely 
arachnoid; petioles to 1 cm long, margined, basally arachnoid. Inflorescences solitary; 
peduncles to ca. 2 cm long, 1-2-bracteolate, the bracteoles linear to oblanceolate, entire 
to laciniate. Capitula 2.5-3.0 mm high, 4-5 mm wide; phyllaries ca. 3-seriate, oblong- 
lanceolate to obovate, 1.5-2.0 mm long, ca. 1 mm wide, apically obtuse to rounded, the 
margins scarious; marginal florets multiseriate, pistillate, the ligules white, bilabiate, the 
outer oval, ca. 0.2-0.4 mm long, entire to retuse, the inner minute, cleft; disc florets 20-^40, 
the corollas funnelform, 1.0-1.5 mm long, 5-lobed, yellow. Achenes obconical, subcom- 
pressed laterally, ca. 1 mm long, glandular, glabrous; pappus absent. 

Plagiocheilus frigidus appears to be restricted to the type locality, in the vicinity 
of the Lago de Junin in Depto. Pasco (ca. 4,100 m). 

PASCO. Pasco: vicinity of Lago de Junin, between Junin & Cerro de Pasco, 
Gentry et al. 19226 (MO). 

3. Plagiocheilus solivaeformis DC., Prodr. 6: 142. 1837. TYPE: Bolivia, no exact 
locality, 1829, Pentland s.n. (G-DC, holotype, not seen, IDC Microfiche 800. 
1035: I. 4). 

Prostrate herbs; stems stoloniform. Leaves ovate-spatulate in outline, 1.0-2.5 cm long 
(including petiole), 6-10 mm wide, 2-pinnatifid, the ultimate segments unequally 2-5- 



MACBRIDE: FLORA OF PERU 15 

lobed, sparsely to densely pilose with flaccid, uniseriate trichomes; peholes 5-15 mm long, 
ciliate. Inflorescences solitary; peduncles 0.5-6.0 cm long, 1-3-bracteolate, the bracteoles 
linear to oblanceolate, entire to pinna tifid. Cap i tula 3-4 mm high, ca. 5 mm wide; phyllaries 
2-seriate, oblong to obovate, ca. 2 mm long, 1.0-1.2 mm wide, apically obtuse to rounded, 
the margins scarious; marginal florets multiseriate, pistillate, the ligules white, bilabiate, 
the outer oval, ca. 0.5 mm long, entire to retuse, the inner minute, cleft; disc florets 20-50, 
the corollas funnelform, ca. 2 mm long, 5-lobed, yellow. Achenes obconical, subcom- 
pressed laterally, ca. 1 mm long, glandular, glabrous; pappus absent. 

Plagiocheilus solivaeformis is distributed from Colombia to Bolivia in moist, 
montane habitats (2,700-4,500 m). It has considerable morphological variation 
within its range and several of these morphs have been afforded specific status 
(P. peduncularis in Ecuador, and P. ciliata in Bolivia). In Colombia, Cuatrecasas 
(1954) has recognized two subspecies, P. solivaeformis subsp. colombianus Cuatr. 
and subsp. multiflorus Cuatr., based upon the degree of leaf division and capitula 
size. The Peruvian material examined in the present study appears typical for 
the species. 

AMAZON AS. Chachapoyas: uppermost slopes and summit of Cerros de 
Calla-Calla, near kms. 403-407 of Balsas-Leimebamba road, 3,400-3,550 m, Wur- 
dack 1712 (F). HUANUCO. Huanuco: forests cerca Pillao, 2,700 m, Woytkowski 
34103 (F, MO); 6 miles S of Mito, ca. 3,353 m, Macbride & Featherstone 1888 (F). 
LA LIBERTAD. Sanchez Carrion: no exact locality, Sagastegui 9442 (F, MO). 



IX. SANTOLINA 
Santolina L., Sp. PL 842. 1753. TYPE: S. chamaecyparissus L. 

Shrubs to subshrubs, rarely herbaceous, strongly aromatic; stems decumbent or as- 
cending, much-branched, leafy. Leaves alternate, pinnatifid, the segments pectinate or 
conferted, sessile. Inflorescences solitary. Capitula discoid, homogamous; involucres 
hemispherical; phyllaries 3-4-seriate, imbricate, the margins scarious; receptacles convex, 
paleaceous; disc florets hermaphroditic, the corollas tubular, 5-lobed, expanded basally 
and encircling the top of the achenes, yellow to white, the anthers basally obtuse, the 
style branches flattened, truncate, penicillate. Achenes 3-4(-5)-angled in section, glabrous; 
pappus absent. Chromosome number: n = 9 (x = 9). 

A genus of some 18 species from western Mediterranean regions. Various 
members are cultivated for ornament and often become established in waste 
places. 

1. Santolina chamaecyparissus L., Sp. PL 842. 1753. TYPE: Europe, exact locality 
and collector unknown (LINN, holotype, not seen, IDC Microfiche 117. 561: 
HI. 2). 

Subshrubs to 20 cm tall; stems erect or ascending, green to gray, tomentose. Leaves 
narrowly linear or oblanceolate in outline, 5-15(-30) mm long, densely pectinate-dentate 
to pinnatifid, often vermiform, tomentose-puberulent, aromatic, the segments to 2 mm 
long, obtuse. Inflorescences solitary; peduncles to 3 cm long. Capitula 5-6 mm high, 5-8 
mm wide; involucres hemispherical, phyllaries 3-seriate, lanceolate to obovate, 3.0-3.5 
mm long, 0.75-1.0 mm wide, carinate, tomentose-puberulent, the outer apically narrowly 
oblong, scarious, lacerate, the inner apically rounded, scarious, lacerate; receptacles con- 
vex, paleaceous, the paleae ca. 3.5 mm long, enfolding the florets, scarious, apically 
puberulent; florets 100-150, the corollas cylindrical to narrowly funnelform, 3.0-3.5 mm 
long, conspicuously exceeding the involucres, the tube of the marginal florets usually 
curved, cream to bright yellow. Achenes narrowly obconical, 1.5-2.0 mm long, 3-4-angled 
in section, glabrous. 



16 FIELDIANA: BOTANY 

This taxon is commonly cultivated for ornamental purposes and was probably 
introduced in Peru from the Mediterranean region where it is native. 

LIMA. Lima: Lima, Soukup 995 (F). 



X. SOLIVA 

Soliva Ruiz & Pavon, Flor. Peruv. Chil. Prodr: 113, t. 24. 1794. TYPE: S. sessilis 
Ruiz & Pavon. 

Annual herbs to 15 cm tall, rosulate, essentially acaulescent; stems stoloniferous. Leaves 
alternate, deeply 3-pinnatifid or rarely 2-pinnatifid, the segments lance-linear or linear- 
deltoid; petioles slender. Inflorescences solitary, sessile in clusters of leaves. Capitula 
disciform, heterogamous; involucres hemispherical; phyllaries 2-seriate, subequal, the 
margins scarious or membranous; receptacles flat to subconical, epaleate; marginal florets 
pistillate, fertile, the corollas absent; disc florets hermaphroditic, functionally male, the 
corollas funnelform, 4-lobed, white, the anthers 4, basally obtuse, the terminal appendages 
ovate, the styles of the marginal florets rigid, spinelike, apically bifurcated, the branches 
filiform, caducous, of the disc florets undivided, truncate, penicillate. Achenes dorsiven- 
trally flattened, convex, often with lateral wings or thickenings, crowned by the persistent 
styles, equaling the achenes in length; pappus absent. Chromosome numbers: n = +55, 
59-60. 

A genus of approximately nine species centered in southern South America, 
but adventive in North America, Europe, Asia, Australia, and New Zealand. 
Only two species have been collected in Peru. 

REFERENCE 

CABRERA, A. L. 1949. Sinopsis del genero Soliva (Compositae). Notas Mus. La Plata, Bot. 
14: 123-139. 



KEY TO SPECIES OF Soliva 

1. Achenes obovate, with thin, planar wings, glabrous 1. S. neglecta. 

1. Achenes narrowly oblong, with narrow, thickened wings, transversely rugose 

2. S. stolonifera. 

1. Soliva neglecta Cabrera, Notas Mus. La Plata, Bot. 14: 128. 1949. TYPE: 
Argentina, Jujuy, Santa Ana, en praderas a 3,100 m, Burkart & Troncoso 11665 
(LP, holotype, not seen; SI, isotype, not seen). 

Annual herbs to 4 cm tall, acaulescent; stems stoloniferous, villous. Leaves rosulate, 
oblanceolate-spatulate in outline, 2-pinnatifid, 1.5-3.0 cm long (including petioles), 4-5 
primary lobes, generally 3-4-parted, the ultimate segments lanceolate, 2-3 mm long, 
0.5-0.7 mm wide, mucronate, villous; petioles to 1 cm long, basally dilated, membranous. 
Inflorescences solitary, sessile in center of leaf rosettes. Capitula 3-4 mm high, 5-6 mm 
wide; phyllaries 2-seriate, ovate to oblong, ca. 3 mm long, 2.0-2.5 mm wide, densely 
villous, apically obtuse to rounded; marginal florets 20-30; disc florets 5-10, the corollas 
1.2-1.4 mm long. Achenes 2.5-3.0 mm long, obovate, dorsally compressed, glabrous, the 
lateral wings terminating in divergent projections, the styles persistent, rigid, spinelike, 
to 1.5 mm long, the apex caducous. 

This species was previously only known from the type locality in northwestern 
Argentina, some 2,000 km disjunct to the southeast of its locality in Peru. It is 
morphologically closest to Soliva sessilis Ruiz & Pavon (Chile) and S. pterosperma 



MACBRIDE: FLORA OF PERU 



17 




1 cm 



FIG. 3. Soliva neglecta. A, habit; B, achene; C, achene from S. stolonifera. (From Cerrate 
2515, MO.) 

(Juss.) Less. (Argentina, Uruguay, and Brasil), but it differs from these species 
in possessing glabrous achenes. 

ANCASH. Bolognesi: Quero, al E de Huasta, 3,600 m, Cerrate 2515 (MO). 

2. Soliva stolonifera 5 (Brot.) Loudon, Hort. Brit.: 364. 1830. 

Hippia stolonifera Brot., Fl. Lusit. 1: 72. 1801. TYPE: Portugal, exact locality and collector 

unknown (LISU, holotype, not seen). 
Gymnostyles nasturtiifolia]uss., Ann. Mus. Natl. Hist. Nat. 4: 262. 1804. TYPE: Argentina, 

Buenos Aires, Commerson s.n. (P-JU, holotype, not seen, IDC Microfiche 6206. 670: 

II. 6). 
Soliva nasturtiifolia (Juss.) DC., Prodr. 6: 142. 1837. 



5 Other synonyms are listed by Cabrera (1949). Only the names listed have been used 
for Peruvian material. 



18 FIELDIANA: BOTANY 

Annual herbs to 5 cm tall; stems stoloniferous, glabrous. Leaves rosulate, oblanceolate- 
spatulate in outline, 1-2-pinnatifid, 2-4 cm long (including petioles), 3-5 primary lobes, 
entire or 1-3-lobed, the ultimate segments oblanceolate to elliptic, 1.5-5.0 mm long, 0.3-1.0 
mm wide, mucronate, villous; petioles to 1 cm long, basally dilated, membranous. Inflo- 
rescences solitary, sessile in center of leaf rosettes. Capitula 2-3 mm high, 4-6 mm wide; 
phyllaries 2-seriate, lanceolate to oblong, 2-3 mm long, 0.5-1.0 mm wide, sparsely villous, 
apically obtuse to rounded; marginal florets 30-40; disc florets 6-10, the corollas 1.2-1.5 
mm long. Achenes obconical, 1.5-2.5 mm long, dorsally compressed, the wings narrow, 
transversely rugose, terminating in two divergent, lateral projections, the styles persistent, 
rigid, spinelike, to 2 mm long. 

This species is native to Uruguay and Argentina and has become naturalized 
in Europe, the Azores, the Canary Islands, and the United States. It is apparently 
rare in Peru, being represented by only one collection. 

LIMA. Huarochiri: Rio Blanco, Macbride & Featherstone 819 (F). 



XI. TANACETUM 
Tanacetum L., Sp. PI. 843. 1753. TYPE: T. vulgare L. 

Annual or perennial herbs, sometimes scapiform, rarely suffruticose, often rhizomatous, 
aromatic; stems erect or ascending, usually leafy and branched, pubescent to glabrous. 
Leaves alternate, entire, toothed, pinnatifid or 1-3-pinnatisect. Inflorescences solitary or 
more often corymbose. Capitula radiate, heterogamous, rarely disciform, heterogamous, 
or discoid, homogamous; involucres hemispherical or campanulate; phyllaries 3-seriate, 
imbricate, lanceolate or oblong, often with scarious margins and apices; receptacles flat 
to convex, naked or with small, rudimentary paleae; marginal florets usually ligulate (rarely 
absent), uniseriate, usually pistillate, the ligules white, yellow, or pink; disc florets her- 
maphroditic, the corollas tubular, 5-lobed, yellow, the anthers basally obtuse, the terminal 
appendages elongate, obtuse, the styles basally enlarged, the branches apically truncate, 
penicillate. Achenes isomorphic, cylindrical or clavate, 3-10-ribbed, glabrous, often glan- 
dular; pappus a short corona, usually unevenly toothed or lobed, sometimes unilateral 
and developed on the posterior side. 

A genus of about 70 species of annuals and herbaceous perennials, with 
centers of diversity in southwest Asia and the Caucasus, although various spe- 
cies are found throughout temperate regions of the northern hemisphere. In 
Peru, three introduced species are common escapes from cultivation. 



REFERENCES 

HEYWOOD, V. H. 1954. A revision of the Spanish species of Tanacetum L. subsect. Leucan- 

themopsis. Anales Inst. Bot. Cavanilles 12: 313-377. 
. 1968. Summary of the divisions of Chrysanthemum, Pyrethrum, Leucanthemum and 

Tanacetum, and a key to the English members. Bot. Soc. Brit. Isles Proc. 3: 177-179. 
HEYWOOD, V. H., AND C. J. HUMPHRIES. 1977. Anthemideae Systematic review. In Hey- 

wood, V. H., et al. (eds.), The Biology and Chemistry of the Compositae, pp. 852-898. 

Academic Press, London. 



KEY TO SPECIES OF Tanacetum 

1. Capitula disciform; all corollas yellow 3. T. vulgare. 

1. Capitula radiate; the ligules conspicuous, white; disc corollas yellow 2. 

2. Leaf segments narrowly lanceolate to oblong-lanceolate, densely silvery-sericeous; 

capitula solitary; involucres 10-15 mm in diameter 1. T. cinerariifolium. 

2. Leaf segments oblong to lanceolate, pubescent to subglabrous; capitula 5-30 in lax 
corymbs; involucres 9-11 mm in diameter 2. T. parthenium. 



MACBRIDE: FLORA OF PERU 19 

1. Tanacetum cinerariifolium (Trev.) Sch. Bip., Tanacet. 58. 1844. 

Pyrethrum cinerariifolium Trev., Ind. Sem. Hort. Vratislav. App. 2: 2. 1820. TYPE: not 
seen. 

Perennial herbs to 50 cm tall, silvery-gray, sericeous. Basal leaves oblanceolate-spatulate, 
to 10 cm long (including petiole), pinnatifid to palmatifid, the segments narrowly lanceolate 
to oblong-lanceolate, glandular-punctate, sericeous; cauline leaves lanceolate to oblanceo- 
late, entire to pinnatifid or palmatifid, 3-6 cm long; glandular-punctate, sericeous; petioles 
widely margined, 1-3 cm long. Inflorescences solitary; peduncles to 30 cm long, striate, 
sericeous, bracteate. Capitula radiate, heterogamous, 6-10 mm high, 10-15 mm wide 
(excluding rays); involucres hemispherical; phyllaries 3-seriate, imbricate, the outer ovate- 
lanceolate, 3-5 mm long, 1.0-1.5 mm wide, puberulent, the inner oblong to obovate, 5-6 
mm long, 1.5-2.0 mm wide, the margins scarious, lacerate; receptacles convex, epaleate; 
ray florets 13-21, pistillate, the ligules oblong, 8-16 mm long, 2-4 mm wide, white; disc 
florets ca. 100, the corollas 2-3 mm long, 5-lobed, yellow. Achenes cylindrical, 2.5-3.5 mm 
long, curving inward, 5-ribbed, glandular; pappus an irregularly lobed corona, 0.7-1.0 
mm high. Chromosome number: n = 9. 

This species is native to western Yugoslavia and Albania and cultivated 
throughout southeastern and east-central Europe as an insecticide. In Peru, it 
is cultivated for its insecticidal properties and horticultural purposes. It is called 
"piretro." 

JUNIN. Huancayo: Huancayo, 3,317 m, Soukup 2933 (F). LIMA. Lima: La 
Molina, terrene cultivado, no collector, USM. # 11548 (MO). 



2. Tanacetum parthenium (L.) Sch. Bip., Tanacet. 55. 1844. 

Matricaria parthenium L., Sp. PI. 890. 1753. TYPE: Europe, exact locality and collector 

unknown (BM, holotype, not seen). 
Chrysanthemum parthenium (L.) Bernh., Syst. Verz. Plf. Erfurt 145. 1800. 

Perennial herbs to 80 cm tall, aromatic; stems erect, densely leafy, prominently angled, 
puberulent to glabrate. Leaves ovate in outline, 3-12(-15) cm long (including petioles), 
1-2-pinnatifid, the primary segments 3-5-paired, 0.7-3.5 cm long, 0.5-2.0 cm wide, usually 
3-7-lobed, the margins dentate or pinnatifid, the ultimate segments oblong to lanceolate, 
obtuse to acute, mucronate, puberulent to glabrous, punctate-glandular beneath; petioles 
0.5-3.5 cm long. Inflorescences laxly corymbose, 5-20(-30)-flowered; peduncles 3-8 cm 
long, puberulent, bracteate. Capitula radiate, heterogamous, 3-5 mm high, 9-11 mm wide 
(excluding rays); involucres hemispherical; phyllaries 3-seriate, imbricate, lanceolate, 
2.0-3.5 mm long, 0.3-0.6 mm wide, keeled, stramineous, puberulent, glandular, the mar- 
gins scarious, ciliolate; receptacles convex, sparsely paleaceous; paleae oblanceolate, cil- 
iolate; ray florets 13-21, the ligules 4-6(-9) mm long, 2-4 mm wide, oval to oblong, 3- 
denticulate, white, the tube ca. 1 mm long, dorsiventrally compressed, enlarged near the 
base; disc florets 150-300, the corollas cylindrical, 1.5-2.0 mm long, 5-lobed, yellow. 
Achenes cylindrical, 1.0-1.6 mm long, curving inward, 5-8-ribbed, glandular; pappus 
absent or an irregularly lobed corona, 0.1-0.3 mm high. Chromosome number: n = 9. 

This species is native to the Balkan peninsula and has long been cultivated 
for ornament and as a medicinal plant throughout Europe. It is a common 
introduction throughout the New World and often escapes from cultivation. 
Label datum indicates that it is used in Peru medicinally as follows (Stork & 
Horton 10265): "Infusion of leaves and stems, as well as flowers, applied exter- 
nally to relieve stomach trouble." It is called "Santa Maria" in Latin America. 

AMAZONAS. Bongara: Pomacochas, 2,200-2,300 m, Ferreyra 15425 (MO); 
Chachapoyas: Chachapoyas, 2,700 m, Williams 7543 (F), 7550 (F). CUZCO. Anta: 
Santa Ana, El Chaccan, 3,511 m, Brunei 550 (F, MO); Cuzco: Picchu, cerca Cuzco, 
3,440 m, Brunei 17 (F, MO). HUANCAVELICA. Tayacaja: Salcabamba, 3,250 m, 



20 FIELDIANA: BOTANY 

Stork & Morton 10265 (F). HUANUCO. Huanuco: Acomayo, 2,500 m, Ridoutt s.n. 
(MO). LA LIBERTAD. Otuzco: Hacienda San Ignacio, Sinsicap, 3,150 m, Lopez 
1122 (MO); Pataz: Huancaspata, 3,100 m, Lopez & Sagastegui 8237 (MO). LIMA. 
Lima: Museo de Historia Natural, Cornjejo s.n. (MO). PASCO. Daniel Carrion: 
Yanahuanca, 3,048 m, Macbride & Featherstone 1251 (F). PUNO. Huancane: Con- 
ima, 3,900 m, Aguilar 428 (MO); Puno: Puno, Soukup 468 (F); Sandia: Cuyocuyo, 
3,500-3,600 m, Ferreyra 16612 (MO). SAN MARTIN. Lamas: San Roque, 
1,350-1,500 m, Williams 7177 (F); San Martin: Tatapoto, 750 m, Williams 5921 (F). 

3. Tanacetum vulgare L., Sp. PI. 844. 1753. TYPE: Europe, exact locality and 
collector unknown (BM, holotype, not seen). 

Perennial herbs to 1.5 m tall, aromatic; stems sparsely pubescent, glandular. Basal leaves 
oblanceolate-spatulate in outline, to 15 cm long (including petiole), 2-pinnatifid, the pri- 
mary segments 7-10-paired, oblong to oblanceolate, 2-5 cm long, the ultimate segments 
oblong to lanceolate, serrate or irregularly toothed; petioles 3-5 cm long; cauline leaves 
ovate in outline, to 10 cm long, sessile, 2-pinnatifid, the primary segments 9-12-paired, 
oblong to oblanceolate, 2-5 cm long, the ultimate segments oblong to lanceolate, serrate 
or irregularly toothed, to 1 cm long, glabrous to sparsely pubescent. Inflorescences cor- 
ymbose, 20-200-flowered. Capitula disciform, heterogamous, 4-5 mm high, 5-10 mm 
wide; involucres hemispherical; phyllaries 3-seriate, imbricate, the outer linear to lanceo- 
late, 2.0-2.5 mm long, 0.6-1.2 mm wide, the inner ovate to oblong, 2.5-3.0 mm long, 
1.0-1.5 mm wide, all with whitish, scarious margins; marginal florets 20-50, 1-seriate, 
pistillate, the corollas cylindrical, ca. 1.5 mm long, zygomorphic, yellow, apically 3-lobed, 
the middle lobe shorter; disc florets 200-300, the corollas cylindrical, 2-3 mm long, 5-lobed, 
yellow. Achenes cylindrical, 1.2-2.0 mm long, 5-ribbed, glandular; pappus an irregularly 
lobed corona, 0.1-^0.3 mm high. Chromosome number: n = 9. 

This species is native to Eurasia and is adventive throughout the New World. 
Although it has long been employed as an herbal remedy to rid the body of 
worms, it is uncertain if it is employed as such in Peru. 

CUZCO. Anta: Llamaponga, El Chaccan, 3,580 m, Brunei 104 (F, MO). 

ACKNOWLEDGMENTS 

Figures were prepared by Marlene Werner, Department of Exhibition, Field 
Museum of Natural History. 



INDEX OF LATIN NAMES 

Names in boldface refer to new species; names in Roman type refer to valid 
species; names in italics refer to synonyms. Numbers in boldface refer to de- 
scriptions; numbers in italics refer to illustrations. 



Abrotanella 12 
Achillea 1, 2 

millefolium 2 
Ambrosia 4 
Anacyclus australis 6 
Anthemidaceae 1 
Anthemideae 1 
Anthemis 1, 11 
Artemisia 2, 3 

absinthium 3 

annua 3, 4 

vulgaris 3 
Centipeda 12 
Chrysanthemum 2, 4 

carinatum 4 

coronarium 4, 5 

indicum 9 

leucanthemum 10 

morifolium 9, 10 

parthenium 19 

segetum 4 
Cotula 1, 2, 5, 12 

australis 6 

cabrera 7 

coronopifolia 5, 6 

mexicana 5, 7, 8 

minuta 7 

pedicellata 7 

pygmaea 7 
Dendranthema 2, 4, 9 

indicum 9, 10 

japonense 10 

makinai 10 

morifolium 1, 9, 10 

ornatum 10 
Gymnostyles minuta 7 

nasturtiifolia 17 

peruviana 7 
Hippia bogotensis 13 

minuta 7 



peduncularis 12 

stolonifera 17 
Lancisia minuta 7 
Leptinella bogotensis 13 
Leucanthemum 2, 4, 10 

maximum 11 

vulgare 10, 11 
Matricaria 2, 11 

courrantiana 12 

parthenium 19 

recutita 11, 12 
Plagiocheilus 1, 2, 12 

bogotensis 13 

cilita 15 

frigidus 13, 14 

peduncularis 12, 15 

prostratus 13 

solivaeformis 13, 14, 15 
subsp. colombianus 15 
subsp. multiflorus 15 

tanacetoides 12 
Pyrethrum cinerariifolium 19 
Santolina 1, 15 

chamaecyparissus 15 
Soliva 1, 2, 12, 16 

mexicana 7 

minuta 7 

nasturtiifolia 17 

neglecta 16, 17 

pedicellata 7 

pedunculata 7 

pterosperma 16 

pygmaea 7 

sessilis 16 

stolonifera 16, 17 
Tanacetum 2, 4, 18 

cinerariifolium 18, 19 

parthenium 1, 18, 19 

vulgare 2, 18, 20 
Tripleurospermum 11 



21 



ECUADOR 



COLOMBIA 



'*'' \ 
8 r-A-. 



BRAZIL 



1 TUMBES 

2 PIURA 

3 CAJAMARCA 

4 AMAZONAS 

5 LORETO 

6 SAN MARTIN 

7 LAMBAYEQUE 

8 LA LIBERTAD 

9 ANCASH 

HUANUCO 

1 LIMA 

2 PASCO 

3 JUNIN 

4 HUANCAVELICA 

5 ICA 

6 AYACUCHO 

7 APURIMAC 

8 CUZCO 

9 MADRE DE DIGS 

20 AREQUIPA 

21 PUNO 

22 MOQUEGUA 

23 TACNA 






km PERU 
Provinces 



CHILE 




Field Museum of Natural History 
Roosevelt Road at Lake Shore Drive 
Chicago, Illinois 60605 
Telephone: (312) 922-9410