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
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FLORA OF PERU
FAMILY COMPOSITAE: PART IV
TRIBE CARDUEAE
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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: 2 C. 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
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