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