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 ANTHEMIDEAE1

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 paleaceous2 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 recutita4 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.

4Other 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 stolonifera5 (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.

5Other 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

0 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
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