UNIVERSITY or

ILLINOIS LIBRARY

AT URBANA-CHAMPAIGN

NATURAL HI8T 8UBVL

FIFI DIANA

Botany

Published by Field Museum of Natural History

NJo. 10

FLORA OF PERU

J. FRANCIS MACB!

COLLABORATORS

FAMILY COMPOSITAE: PART IV

TRIBE CARDUEAE

MICHAEL O. DILI

•mber 24, 1982

Publical

FLORA OF PERU

FAMILY COMPOSITAE: PART IV

TRIBE CARDUEAE

~D GI G» GI

xi iT-i en 4*

0

Fl "7 "T HT IT" IT

x x x x x

!-•• •>:• -•:" V V 'x'

o ro

en

TJ

W

n GI GI GI GI G« G« iz

en en en r n •' n «' n n'i i'i"i

-T -» -r -^ -r _ ro "

o o o o o 5! en

o o o o o uri co

HH I--I i--l K < K< I'fl p'l

"

co

pi

~n

I-H

m

^-^

X;

-J-.

X'

/n >.£i

--I

^ en

CO o:«

r

^-^

m

73

•c

o

CO

I-H

en H

Xi

X"

f

fc* l'> C

n o i/;

L.I en

iTi

ro G'

GI en

CO

rn

CO

G'

•

en

-q

h-l

CO

m

Tl

n

en

-

.

^

a Cfc^-r^Bj J

- •

**-&O ' • rV_>?

I

^^ :

FIELDIANA

Botany

Published by Field Museum of Natural History

New Series, No. 10

FLORA OF PERU

J. FRANCIS MACBRIDE

AND COLLABORATORS

FAMILY COMPOSITAE: PART IV

TRIBE CARDUEAE

MICHAEL O. DILLON

Assistant Curator

Department of Botany

Field Museum of Natural History

Accepted for publication March 31, 1982

September 24, 1982

Publication 1335

Library of Congress Catalog Card No.: 80-66384

ISSN 0015-0746

PRINTED IN THE UNITED STATES OF AMERICA

CONTENTS

Tribe Cardueae

Key to Genera of Cardueae 2

I. Centaurea 2

1. Centaurea melitensis 2

II. Cirsium *

1. Cirsium vulgare 4

HI. Silybum 6

1. Silybum marianum 6

Acknowledgment 8

LIST OF ILLUSTRATIONS

1 . Centaurea melitensis 3

2. Cirsium vulgare 5

3. Silybum marianum 6

Tribe CARDUEAE

Cardueae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 155. 1819. TYPE: Carduus L.

Carlineae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 152, 1819. TYPE: Carlina L.

Centaureae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 154. 1819. TYPE: Centaurea L.

Echinopsideae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 157. 1819. TYPE: Echinops L.

Cynareae Less., Linnaea 5: 128. 1830. TYPE: Cynara L.

Annual, biennial, or perennial herbs, rarely small shrubs; stems simple or branched,

often spiny. Leaves alternate, usually pinnatifid or incised, rarely entire, the margins

usually dentate, spinose, often densely tomentose or lanate. Capitula homogamous and

discoid, the corolla lobes linear, equal to subequal, heterogamous and disciform or

radiate, the marginal florets pistillate or neuter, the corollas ligulate or rarely 2-labiate, the

inner florets with corollas tubular, deeply to shallowly 5-lobed, homochromous, yel-

lowish or purplish, rarely white, the anthers basally sagittate or caudate, the auricles

entire, irregularly lacinate, or marginally pinnatifid, the terminal appendages elongate,

acute, the styles slightly thickened or with a ring of minute hairs near the point of

bifurcation, the branches separate or connate, short, obtuse. Achenes ± terete or com-

pressed, obovate, unbeaked, sometimes umbonate, mostly glabrous or sparsely covered

with one-celled trichomes; pappus a crown a multiseriate bristles, rarely paleaceous, or of

uniserate bristles or paleae fused basally.

The Cardueae sensu lato includes approximately 80 genera, primarily Eur-

asian, chiefly Mediterranean, and southwest to central Asia. Dittrich (1977) has

proposed following Cassini's (1817, 1819) splitting of the tribe into presumably

more natural units by removing the tribes Echinopsideae Cass.1 (2 genera/ca. 120

species) and Carlineae Cass. (11 genera/ca. 90 species) and recognizing two

subtribes within the Cardueae, the Carduinae Dumort. (40 genera/ca. 1,700

species) and the Centaureinae Dumort. (27 genera/ca. 813 species). While the

delineation of natural taxonomic units is utilitarian, the fragmentation of an

obviously monophyletic tribe seems unnecessary.

The tribe is represented in Peru by three species, all native to the Old World

and adventive weeds in the New World.

This study was assisted by National Science Foundation Grant DEB-79-05078 (Alwyn

H. Gentry, principal investigator).

'Cited as Echinopeae Cass. by Dittrich (1977).

1

FIELDIANA: BOTANY

REFERENCES

CASSINI, H. 1817. Cinarochephales. In Leurault, F. G. (ed.), Dictionnaire des Sciences

Naturelles, 9: 226-227. Strasbourg & Paris.

— . 1819. Sixieme memoire sur 1'ordre des synantherees, contenant les caracteres des

tribus. J. Phys. Chim. Hist. Nat. Arts 88: 152-159.

DITTRICH, M. 1977. Cynareae-systematic review. In Heywood, V. H., et al. (eds.), The

Biology and Chemistry of the Compositae, pp. 999-1015. Academic Press, London.

KEY TO GENERA OF CARDUEAE

1. Leaves without spines; achenes articulating laterally I. Centaurea.

1. Leaves spiniferous; achenes articulating basally 2.

2. Phyllaries spine-tipped, lacking marginal or lateral spines II. Cirsium.

2. Phyllaries spinescent, marginal or lateral spines present HI. Silybum.

I. CENTAUREA

Centaurea L., Sp. PI. 909. 1753. TYPE: C. centaurium L.

Annual or perennial herbs, rarely dwarf shrubs. Leaves alternate or basal, entire to

pinnatifid. Inflorescences solitary or cymose-paniculate. Capitula heterogamous, dis-

ciform or radiate, or rarely homogamous, discoid; involucres cylindrical to globose;

phyllaries imbricate, multiseriate, coriaceous or scarious, often spinose; receptacles

plane, bristly; marginal florets sterile (sometimes with staminodes), the corollas funnel-

form, deeply 5(-8)-lobed, ± zygomorphic; disc florets hermaphrodite, tjie corollas tubu-

lar, deeply 5-lobed, yellow, violet, purple, blue, or white, the anthers basally sagittate,

the styles bifid, separate or connate, often with a ring of short hairs below point of

bifurcation, the branches linear. Achenes obovoid to oblong, compressed or tetragonal,

glabrous or puberulent with unicellular trichomes of varying lengths, articulating later-

ally; pappus persistent or rarely caducous, of 2 to several rows of scabrid to plumose setae,

or oblong to linear squamellae, the inner longer.

A genus of 500-600 species, primarily distributed in the Mediterranean re-

gion and southwest Asia, with a few from tropical Africa and North and South

America. In Peru only the following introduced weed has been collected.

1. Centaurea melitensis L., Sp. PI. 917. 1753. TYPE: Europe, exact locality and

collector unknown (LINN, holotype, not seen, IDC Microfiche 117. 619: III. 1).

Annual or biennial herbs to 80 cm tall; stems erect, winged above, striate, arachnoid,

crispate-puberulent. Lower leaves oblanceolate in outline, 10-15 cm long, 1.5-3.0 cm

wide, lyrate-pinnatifid to sinuately lobed, apically obtuse, basally cuneate, the upper

oblong to oblanceolate, 2-6 cm long, 0.5-2 cm wide, entire to irregularly undulate-

dentate, decurrent on the stems, both sides green, crispate-puberulent, arachnoid,

glandular-punctate. Inflorescences of solitary capitula at the tips of branchlets or weakly

cymose. Capitula homogamous, discoid; involucres ovoid-globose, 8-12 mm wide, ca. 12

mm high; outer phyllaries ovate, 4-8 mm long, 2-3 mm wide, glabrous to arachnoid-

tomentose, scarious, indurate, stramineous, apically and marginally spinulose, squar-

rose, the inner lance-ovate to lanceolate, 8-9 mm long, 1.5-2.0 mm wide, glabrous, scari-

ous, stramineous, apically spinulose, all apices reddish; receptacles plane; corollas yel-

low, tubular, ca. 1 cm long, glandular, 5-lobed, the lobes 1-2 mm long, linear. Achenes

oblong, 2.5-3.0 mm long, puberulent to glabrate; pappus multiseriate, the outer of narrow

squamellae, 0.4-1.6 mm long, the inner of elongate scabrid setae, ca. 3 mm long. Chromo-

some numbers: n = 11, 12, 18.

This species is a native of Europe and North Africa and is an adventive weed

in North and South America. It occurs in a wide range of habitats throughout

Peru, but most frequently in disturbed areas. No vernacular names have been

recorded for this species in Peru; however, it is known as "abre puno amarillo"

in Argentina and "cizana" in Chile.

1cm

2cm

2cm

FIG. 1. Centaurea melitensis. A, habit; B, floret; C, basal leaf. (A, B from Macbride &

Featherstone 1029, F; C from Sagtistegui & Araujo 7785, F.)

4 FIELDIANA: BOTANY

Centaurea solstitialis L. is the only other member of this genus, with yellow

corollas, known to occur in South America (Argentina), and it is easily distin-

guished by its much larger involucral spines (10-25 mm long).

CAJAMARCA. Contumaza: alrededores de Contumaza, 2,600 m, Sagdstegui et

al. 9856 (F, MO). CUZCO. Calca: alturas de Taray, 3,050 m, Vargas 4259 (F).

JUNlN. Tarma: 4 km W of Tarma, Edwin & Schunke 3890 (F); Tarma, Macbride &

Featherstone 1029 (F). LA LIBERTAD. Trujillo: alrededores de Trujillo (Santa

Rosa), 50 m, Sagdstegui & Araujo 7785 (F, MO).

II. CIRSIUM

Cirsium Mill., Card. Diet. Abr. ed. 4. 1754. LECTOTYPE:2C. heterophyllum (L.)

Hill (Carduus heterophyllus L.)

Perennial or biennial herbs, rarely annuals; stems simple or branched, rarely acaules-

cent. Cauline leaves alternate, often decurrent, serrate or pinnatifid, the lobes dentate,

spinose. Inflorescences solitary or corymbose to paniculate, rarely sessile among basal

leaves. Capitula homogamous, discoid; involucres ovoid or globose; phyllaries mul-

tiseriate, imbricate, graduate, narrow, appressed basally, usually spinescent, sometimes

vittate; receptacles flat to subconic, bristly; florets hermaphrodite (rarely functionally

gynodioecious or dioecious), the corollas purplish, white, pinkish, or rarely yellowish,

tubular, the limb symmetrical or oblique, 5-lobed, the lobes linear, the anthers basally

caudate, the auricles fringed, the apices long-appendaged, the styles with a thickened,

often hairy ring below the branches, the branches linear or filiform, subobtuse, connate

except distally. Achenes ± oblong, gibbous, compressed or 4-angulate, articulating ba-

sally, smooth or costate, glabrous, truncate or umbonate apically; pappus bristles mul-

tiseriate, equal to subequal, plumose, affixed to basal annulus, deciduous in a ring or ±

persistent.

A genus of 250-300 species of temperate Eurasia and North America, with

several species becoming naturalized weeds. In Peru, only the European weed

Cirsium vulgare is represented.

REFERENCE

AIRY-SHAW, H. K. 1938. On the correct names of three European species of Cirsium.

Feddes Repert. Spec. Nov. Regni Veg. 43: 302-315.

1. Cirsium vulgare (Savi) Tenor, Fl. Napol. 5: 209. 1835-36.

Carduus lanceolatus L., Sp. PI. 821. 1753. TYPE: Europe, exact locality and collector

unknown (LINN, holotype, not seen, IDC Microfiche 117. 548: I. 4).

Cirsium lanceolatum (L.) Scop., F. Carniol. ed. 2. 2:130. 1772. (non C. lanceolatum Hill,

Veg. Syst. 4: 27. 1762), nom. illeg.

Carduus vulgaris Savi, Fl. Pis. 2: 241. 1798. TYPE: not seen.

Cirsium vulgare (Savi) Airy-Shaw, Feddes Repert. Spec. Nov. Regni Veg. 43: 304. 1938,

nom. illeg., superfluous combination.

Annual or biennial herbs to 1.5 m tall; stems branched above, winged, arachnoid to

spreading-hirsute. Cauline leaves lanceolate, to 25 cm long, to 10 cm wide, pinnatifid, the

lobes toothed or lobed, all spinescent, decurrent for the entire internode, scabrous-hispid

above, thinly white tomentulose to green and hirsute below. Inflorescences solitary to

weakly corymbose or paniculate, pedunculate. Capitula homogamous, discoid; in-

volucres obovoid, to 40 mm high, to 50 mm wide; phyllaries lance-linear, spinescent,

designated by Britton & Brown, 111. Fl. N. U.S. ed. 2. 3: 548. 1913. Airy-Shaw (1938) has

shown this taxon to be conspecific with Cirsium helenioides (L.) Hill (Carduus helenioides

L.).

1cm

FIG. 2. Cirsium vulgare. A, habit; B, floret. (From Brunei 679, MO.)

5

6 FIELDIANA: BOTANY

sparsely arachnoid, the outer ca. 10 mm long, 1-2 mm wide, the inner to 5 mm long, 1-2

mm wide; corollas purple, tubular, 26-36 mm long, mostly 5-lobed, the lobes 5-6 mm

long, linear. Achenes obovoid-elliptic, to 5 mm long, glabrous, apically truncate with an

annular thickening; pappus bristles multiseriate, plumose, to 30 mm long, deciduous in a

ring. Chromosome number: n = 34.

This species is native to Europe and naturalized throughout temperate re-

gions of the world. It occurs throughout South America, from Ecuador to

Argentina. In Peru, it is known by the following Quecha names, "alq'u kiska"

and "h lauli."

CUZCO. Anta: Santa Ana, El Chaccan, 3,493 m, Brunei s.n. (MO), Huaypo

Grande, 3,711 m, Brunei 679 (MO).

III. SILYBUM

Silybum Adans., Fam. PI. 2: 116, 605. 1763. Nom. cons. TYPE: Carduus marianus

L. = Silybum marianum (L.) Gaertn.

Annual or biennial herbs; stems erect from a basal rosette, simple or sparingly

branched, unwinged. Leaves alternate, white veined or variegated, margins spinose.

Inflorescences solitary. Capitula homogamous, discoid; involucres ovoid to subglobose;

phyllaries multiseriate, imbricate, spiny-margined, strongly spinose; receptacles flat,

densely setose; florets hermaphrodite, the corollas purple, tubular, the limb slightly ex-

panded, deeply 5-lobed, the tube slender, the anthers shortly tailed, the style branches

usually connate. Achenes subcompressed, obovoid-oblong, articulating basally, smooth,

glabrous, the inner shorter, all connate basally, deciduous in a ring. Chromosome

number: n = 17.

A genus of two species, with the following one, adventive throughout the

New World.

REFERENCE

VARMA, P. N., S. K. TALWAR, AND G. P. GARG. 1980. Chemical investigations of Silybum

marianum. PI. Med. 38: 377-378.

1. Silybum marianum (L.) Saertn., Fruct. et Semin Plant. 2: 398. 1791. Carduus

marianus L., Sp. PL 2: 823. 1753. TYPE: Europe, exact locality and collector

unknown (LINN, holotype, not seen, IDC Microfiche 117. 549: III. 4).

Annual herbs to 2 m tall; stems erect, glabrous or slightly arachnoid-pubescent, striate.

Basal leaves in a rosette, obovate, 10-50(-80) cm long, 5-15 cm wide, pinnatifid, glabrous

or glabrescent, petiolate, the upper leaves sessile, auriculate-amplexicaul, spinescent.

Inflorescences solitary. Capitula discoid; involucres globose, to 4 cm wide, 4-6 cm high;

phyllaries basally ovate to lanceolate, widening abruptly to an ovate-subulate appendage

distally, the outer 1-2 cm long, often lacking an appendage, the inner 4-5 cm long (in-

cluding appendage), all ca. 1 cm wide, the margins and apices spinose, reflexed to erect;

corollas ca. 30 mm long, the lobes ca. 6 mm long, linear. Achenes ovoid to obovoid, 6-8

mm long; pappus 15-20 mm long. Chromosome number: n = 17.

This species is widely distributed within Peru, primarily at elevations above

3,000 m. It is known throughout Latin America by many vernacular names,

including "cardo, cardo manchado, cardo mariano, cardo bianco, cardo santo,"

and in Peru as "cardo de la alamenda."

Silybum marianum has been used in homoeopathic medicine in India, where

its seeds are used in the control of hemorrhage, jaundice, cirrhosis of the liver,

and related liver and gallstone conditions (Varma et al., 1980). At present, no

record exists of this plant species being used medicinally in Peru.

1cm

B

2cm

FIG. 3. Silybum marianum. A, habit; B, floret. (From Dillon b Turner 1569, F.)

7

FIELDIANA: BOTANY

CAJAMARCA. Cajamarca: ca. 16 km NE of Cajamarca, on road to Celendin,

ca. 3,000 m, Dillon & Turner 1569 (F, MO, TEX, USM). JUNfN. Huancayo: 16.5

km W of Huancayo, Soukup 2932 (F). LA LIBERTAD. Otuzco: Salpo, ca. 3,400

m, Lopez 1850 (F); Pataz: alrededores de Huancaspata, 3,100 m, Lopez & Sagds-

tegui 8241 (MO). CERRO DE PASCO. Pasco: Huariaca, ca. 3,050 m, Macbride &

Featherstone 2406 (F). PUNO. Puno: Amantani, 3,900 m, Aguilar s.n. (MO).

ACKNOWLEDGMENT

Figures were prepared by Marlene Werner, Department of Exhibition, Field

Museum of Natural History.

1 TUMBES

2 PIURA

3 CAJAMARCA

4 AMAZONAS

5 LORETO

6 SAN MARTIN

7 LAMBAYEOUE

8 LA LIBERTAD

9 ANCASH

10 HUANUCO

1 1 LIMA

12 PASCO

13 JUNIN

14 HUANCAVELICA

15 ICA

16 AYACUCHO

17 APURIMAC

18 CUZCO

19 MADRE OE DIOS

20 AREOUlPA

21 PUNO

22 MOOUEGUA

23 TACNA

UNIVERSITY OF ILLINOIS-URBANA

580.5FBN.S. C005

FIELDIANA. BOTANY CHGO

101982

Field Museum of Natural History

Roosevelt Road at Lake Shore Drive

Chicago, Illinois 60605-2496

Telephone: (312) 922-9410

UNIVERSITY OF ILLINOIS-URBANA