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The subfamily Mutisioideae (Asteraceae).

Leibnitzia Cass.

Dict. sci. nat. 25: 420. Nov. 1822. TYPE: Tussilago anandria L. [= Leibnitzia anandria (L.) Turcz.]. 28 of Figs. 27-30.

Anandria Less., Linnaea 5: 241. 1830, nom. illeg.

Cleistanthium Kunze, Bot. Zeitung (Berlin) 9: 40. 1851. TYPE: Cleistanthium nepalense Kunze [= Leibnitzia nepalensis (Kunze) Kitam.].

Etymology. In honor of the German scientist Gottfried Wilhelm Leibnitz (1646-1716), an eminent eighteenth century philosopher and mathematician interested in botany.

Herbs perennial, acaulescent, scapose, with plant dimorphism: a vernal (spring) stage with reduced or no leaves and chasmogamous capitula, and an autumnal (summer and fall) stage with fully developed leaves and cleistogamous capitula. Leaves rosulate; pseudopetiolate, blades lanceolate, elliptical to obovate, pinnately veined, margin denticulate to lyrate, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, disciform or radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic with a gradual transition from marginal to central florets, marginal florets female, corolla true ray, filiform to bilabiate with a minute inner lip, outer lip expanded or not, with or without staminodes; central florets bisexual, bilabiate to tubular-bilabiate; anther apical appendages acute, tails smooth; style bilobed, branches dorsally papillose. Cypselae rostrate at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Leibnitzia occimadrensis G. L. Nesom. Pollen spheroidal, circular in polar view, medium size, P x E = (35-42x36-40) [micro]m. Tricolporate, colpi long with microgranulate membrane. Exine Mutisia type, microechinate, 7 [micro]m thick at the equator and 4-5 [micro]m thick at the poles, ectosexine slightly columellate. Ratio ectosexine/endosexine: 1:2. Nexine ca. 2 [micro]m thick (see also Lin et al., 2005).

Habitat and Distribution. Genus of six species distributed in North and Central America and Asia.

Species list

1. Leibnitzia anandria (L.) Turcz., Ukaz. Otkryt. 8: 404. 1831. Basionym: Tussilago anandria L. TYPE: Sine loc., s.d., s. leg. (holotype, LINN 995 not seen). Additional specimen examined: Russia. Siberia, Tuva Autonomous Republic, near Lake Chagiti and Tanv Ola mtn. range, T. Elias et al. 8183 (US).

2. Leibnitzia knorringiana (B. Fedtsch.) Pobed., Fl. U.S.S.R. 28: 594. 1963. Basionym: Gerbera knorringiana B. Fedtsch. TYPE: Soviet Central Asia. Tien Shah: Lake Sary-Chilek, s.d., s. leg. (type, probably LE).

3. Leibnitzia nepalensis (Kunze) Kitam., J. Jap. Bot. 14: 297. 1938. Basionym: Cleistanthium nepalense Kunze. TYPE: Sine loc., Flor. Aug. 1850, s. leg. (holotype, based on a cultivated specimen in LZ not seen).

4. Leibnitzia occimadrensis G. L. Nesom, Brittonia 35: 130. 1983. TYPE: Mexico. State Sinaloa: Sierra Surutato, Los Pucheros, H. S. Gentry 7189 (isotype, GH!).

5. Leibnitzia ruficoma (Franch.) Kitam., J. Jap. Bot. 14: 297. 1938. Basionym: Gerbera ruficoma Franch. TYPE: China. Che-tcho-tze, 10 Oct 1882, Delavay s.n. (holotype, P? not seen). Additional specimen examined: China. Sikang Province: Sacred Mt., Karwar-kar-boo, Tsa-wa-rung, C. W. Wang 66451 (GH).

6. Leibnitzia seemannii (Sch. Bip.) G. L. Nesom, Brittonia 35: 127. 1983. Basionym: Chaptalia seemannii Sch. Bip. TYPE: Mexico. Durango: Sierra Madre, Seemann 1988 (isotype, K not seen, K photo at LP!); Federal District, Cima Station, C. G. Pringle 13544 (WIS!).

Literature. Nesom (1983), Hansen (1988).

Lulia Zardini

Bol. Soc. Argent. Bot. 19: 255. 1980. TYPE: Trichocline nervosa Less. [= Lulia nervosa (Less.) Zardini]. 29 of Figs. 27-30.

Etymology. In honor of the Spanish botanist, resident in Argentina, Angel Lulio Cabrera (1908-1999) who made enormous contributions to the study of Asteraceae.

Herbs caulescent, perennial. Leaves alternate; sessile; blades linear, graminiform, long, parallel-veined, margin entire, glabrous. Capitulescences monocephalous, terminal; capitula long-pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with well developed staminodes, corolla bilabiate, with an expanded outer lip; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bilobed, branches dorsally papillose. Cypselae rostrate at the apex, pubescent; pappus with barbellate, capillary bristles.

Pollen Description (Telleria & Katinas, 2004) (98 of Figs. 95-100). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate.

Habitat and Distribution. Monotypic genus endemic to Sphagnum U bogs in southern Brazil.

Species list

1. Lulia nervosa (Less.) Zardini, Bol. Soc. Argent. Bot. 19: 257. 1980. Basionym: Trichocline nervosa Less. TYPE: Brazil. State Silo Paulo: Campos da Bocaina, A. F. M. Glaziou 8110 (neotype, designated by Zardini, 1980: 258, P not seen). Additional specimen examined: Brazil. State Santa Catarina: Mun. Campo Alegre, fazenda de Ernesto Scheide, S. B. Smith & R. M. Klein 10576 (LP*).

Literature. Zardini (1975, 1980).

Lycoseris Cass.

Dict. sci. nat. 33: 463. 1824. TYPE: Atractylis mexicana L. f. [= Lycoseris mexicana (L. f.) Cass.]. 30 of Figs. 27-30.

Diazeuxis D. Don, Trans. Linn. Soc. London 16: 251. 1830. TYPE: Diazeuxis mutisiana D. Don [= Lycoseris mexicana (L. f.) Cass.].

Etymology. From the Greek lykos, wolf, and seris, chicory, lettuce.

Subshrubs dioecious. Leaves alternate; subsessile; blades elliptic to ovate, threeveined, margin entire to minutely dentate, glabrescent to glabrous above, floccose to dense-arachnoid beneath. Capitulescences monocephalous or paniculate to corymbose, terminal; capitula pedunculate, homogamous or heterogamous, radiate; receptacle epaleate, setose; involucre multiseriate. Florets of female capitula dimorphic, with staminodes, style sometimes widened above, bifid, branches dorsally papillose; marginal florets neuter, corolla tree ray three-dentate, bilabiate to sub-bilabiate (3+1, 4+1 corolla lips), style rudimentary; central florets female, corolla actinomorphic, narrowly tubular, shallowly five-lobed, or sub-zigomorphic with unequal insicions; male capitula with florets dimorphic; marginal florets neuter, corolla true ray threedentate to sub-bilabiate (3+1 corolla lips), with staminodes, style rudimentary, branches dorsally papillose; central florets functionally male, corolla tubular-bilabiate to actinomorphic, tubular-funnelform, shallowly five-lobed, anther apical appendage acute, tails smooth, style widened above, dorsally papillose, papillae longer than in the female capitula. Cypselae glabrous; pappus of numerous, scabrid bristles in female florets, and flatter, seta-like, and relatively few scabrid bristles in male florets.

Pollen Description. Lycoseris trinervis (Ruiz & Pay. ex D. Don) S. F. Blake (99 of Figs. 95-100). Pollen subprolate prolate, elliptic in equatorial view, circular in polar view, medium to large size, P x E = (47-55 x 33-39.5) [micro]m. Tricolporate, colpi long with psilate membrane. Exine Mutisia type, microechinate, 5-6 [micro]m thick, slightly slender at the poles. Ratio ectosexine/endosexine: 1:1; 1.5:1. Nexine ca. 1 [micro]m thick (see also Egerod & Stahl, 1991).

Habitat and Distribution. Genus of 11 species of tropical and subtropical America, from Guatemala to Brazil and Bolivia.

Species list

1. Lycoseris boliviana Britton, Bull. Torrey Bot. Club 19: 266. 1892. TYPE: Bolivia. Dept. La Paz: Guanay, H. H. Rusby 1557 (lectotype, designated by Egerod & Stahl, 1991: 569, NY not seen, NY photo at LP!)

2. Lycoseris colombiana K. Egerod, Nordic J. Bot. 11: 564. 1991. TYPE: Colombia. Dept. Cauca: El Bordo, Archer 3388 (holotype, US not seen).

3. Lycoseris crocata (Bertol.) S. F. Blake, Bull. Torrey Bot. Club 53: 218. 1926. Basionym: Aster crocatus Bertol. TYPE: Guatemala. Vulcano d' acqua, 1837, Velasquez s.n. (holotype, BOLO not seen). Additional specimens examined: Colombia. Dept. Atlantico: cerca a Usiacuri, J. Araque Molina & F. A. Barkley 19 At063 (LP); Puerto Colombia, Bro. Paul 931 (LP).

4. Lycoseris eggersii Hieron., Bot. Jahrb. Syst. 29: 75. 1901. TYPE: Ecuador. Prov. Manabi: El Recreo, H. F. A. Eggers 14952 (isotype, LP!). Additional specimens examined: Ecuador. Prov. Manabi: south of Santa Ana, O. Haught 3462 (LP). Bolivia. Dept. Santa Cruz: Prov. Velazco, San Miguelito, 23 km N de San Ignacio, A. Krapovickas & A. Schinini 32419 (LP).

5. Lycoseris grandis Benth., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn: 111. 1852. TYPE: Costa Rica. San Matheo, A. S. Oersted 10517 (isotype, US not seen, US photo at LP!).

6. Lycoseris mexicana (L. f.) Cass., Dict. sci. nat. 33: 474. 1824. Basionym: Atractylis mexicana L. f. TYPE: Sine loc., s.d., J. C. Mutis s.n. (holotype, LINN 971 not seen).

7. Lycoseris minor K. Egerod, Nordic J. Bot. 11: 559. 1991. TYPE: Colombia. Dept. Huila: Cordillera Oriental, valley of Rio Fortalecillas, 32-33 km of Neiva, F. R. Fosberg 19778 (holotype, US not seen, US photo at LP!).

8. Lycoseris peruviana K. Egerod, Nordic J. Bot. 11: 564. 1991. TYPE: Peru. Dept. San Martin: Pasaraya, near Saposoa, F. Woytkowski 5080 (isotype, NY not seen, NY photo at LP!).

9. Lycoseris retroflexa J. Kost., Blumea 5: 664. 1945. TYPE: Bolivia. Dept. Santa Cruz: near Paliza, T. Herzog 1263 (holotype, L not seen).

10. Lycoseris trinervis (Ruiz & Pav. ex D. Don) S. F. Blake, Contr. U. S. Natl. Herb. 22: 653. 1924. Basionym: Diazeuxis trinervis Ruiz & Pay. ex D. Don. TYPE: Ecuador. Huayaquil, s.d., Tafalla s.n. (lectotype, designated by Egerod & Stahl, 1991: 559, BM not seen). Additional specimens examined: Peru. Junin: San Ramon, Chanchamayo, L. Diers 1249 (LP*); id., Woytkowski 7402 (us).

11. Lycoseris triplinervia Less., Linnaea 5: 257. 1830. TYPE: Colombia. Santa Marta, yon Rohr 27 (holotype, C not seen). Additional specimen examined: Venezuela. State Barinas: 80 km from Barinas, along road to San Cristobal, bank of Rio Bumbum, F. J. Breteler 4414 (LP).

Observation. Lycoseris displays some style characters that are not common in the tribe Mutisieae such as apically somewhat widened styles and style branches of male capitula covered by long papillae. These modifications of the style are presumably due to the dioecious condition of this genus. The pollen features, on the other hand, reveal the typical structure of most members of the tribe Mutisieae.

Literature. Egerod and Stahl (1991), Diaz-Piedrahita and Velez-Nauer (1993), Ferreyra (1995).

Macroclinidium Maxim.

Bull. Acad. Imp. Sci. Saint-Petersbourg, set. 3, 15: 375. 1871. TYPE: Macroclinidium robustum Maxim. 31 of Figs. 31-34.

Pertya sect. Macroclinidium (Maxim.) Makino, Bot. Mag. (Tokyo) 14: 144. 1900. TYPE: Macroclinidium robustum Maxim.

Etymology. From the Greek makros, long, large, and cline, a bed, klinidion, little bed, probably referring to the large receptacle.

Herbs perennial, unbranched. Leaves clustered near the median part of the stem; sessile to long-petiolate; blades ovate, obovate, three-lobed, palmately, pinnately, or three-veined to subpalmately veined, margin dentate to serrate, glabrous to slightly pilose especially beneath. Capitulescences spicate to paniculate; capitula sessile, homogamous, ligulate, one- to ten-flowered; receptacle epaleate, glabrous to setose; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally papillose. Cypselae glabrous to pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Macroclinidium (100 of Figs. 95-100). Pollen spheroidal or suboblate, circular in polar view, medium or large size. Tricolporate, colpi very short with microgranulate membrane. Exine Macroclinidium type, equally thickened in the whole grain. Note: the Macroclinidium exine type is similar to the Cardueae exine type; they differ in the morphology of the spines, being solid in the Macroclinidium type and with an apical channel in the Cardueae type. Macroclinidium rigidulum (Miq.) Makino. Pollen spheroidal, large size, P x E = (51-60 x 52-56) [micro]m. Colpi ca. 23 [micro]m length. Exine ca. 14 [micro]m thick in the whole grain, spines ca. 5 [micro]m length. Nexine 2-2.5 [micro]m thick. Macroclinidium robustum. Pollen suboblate, medium size, p x E = (44-50x50-64) [micro]m. Colpi ca. 16 [micro]m length, with psilate membrane. Exine 9-12 [micro]m thick, spines 4-5 [micro]m length. Nexine ca. 1.5 [micro]m thick (see also Lin et al., 2005).

Habitat and Distribution. Genus of six species endemic to Japan.

Species list

1. Macroclinidium hybridum (Makino) Matsum., Index pl. jap. 3: 658. 1912. Basionym: Pertya hybrida Makino. TYPE: Japan. Musashi: Omiya-hachiman in Wada-mura, 1897, Makino s.n. (type, MAK not seen, MAK photo at LP!).

2. Macroclinidium koribanum, Nakai, Bot. Mag. (Tokyo) 26: 249. 1912. TYPE: Japan. Sendai: Taihakusan, 1901, K. Koriba s.n. (type, probably TI).

3. Macroclinidium rigidulum (Miq.) Makino, Bot. Mag. (Tokyo) 12: 194. 1898. Basionym: Eupatorium rigidulum Miq. TYPE: not localized. Additional specimens examined: Japan. Honshu: Ishikawa, Nunohashi-machi, Komatsushi, Tsugaru & Takeuchi 7324 (NY); id., Deguchi et al. 6382 (MO*).

4. Macroclinidium robustum Maxim., Bull. Acad. Imp. Sci. Saint-Petersbourg, ser. 3, 15: 376. 1871. TYPE: Japan. Jokohama, Wawra 1527 (type, NY!).

5. Macroclinidium suzuki Kitam., J. Jap. Bot. 14: 385. 1938. TYPE: Japan. Hondo: Shirakawa, Kinshoji, 20 Oct 1930, S. Suzuki s.n. (type, probably TI).

6. Macroclinidium trilobum (Makino) Makino, Bot. Mag. (Tokyo) 7: 302. 1894; Bot. Mag. (Tokyo) 12: 194. 1989. Basionym: Ainsliaea triloba Makino. TYPE: not localized. Additional specimen examined: Japan. Rikugen, 1926, Masamune s.n. (NY).

[FIGURES 31-34 OMITTED]

Literature. Kitamura (1937), Ohwi (1965).

Mutisia L. f

Suppl. pl.: 57. 1781 (1782). TYPE: Mutisia clematis L. f. 32 of Figs. 31-34.

Aplophyllum Cass., Dict. sci. nat. 33: 472. 1824. TYPE: not designated, based on the three species Mutisia decurrens Cav., M. ilicifolia Cav., and M. linearifolia Cav.

Guariruma Cass., Dict. sci. nat. 33: 472. 1824. [= Mutisia sect. Guariruma (Cass.) Cabrera, Opera Lilloana 13: 138. 1965]. TYPE: Mutisia hastata Cav. (lectotype designated by Cabrera, 1965).

Etymology. In honor of the Spanish botanist Jose Celestino Bruno Mutis y Bosio (1732-1808), physician, clergyman, and first director of the Real Expedicion del Nuevo Reino de Granada.

Shrubs or subshrubs stout to trailing, stem winged to unwinged. Leaves alternate; sessile to short-petiolate; blades simple to pinnately compound, with or without terminal tendrils, linear, subulate, lanceolate to ovate, pinnately to one-veined, margin entire to runcinate, sometimes revolute, glabrous to pubescent. Capitulescences monocephalous, terminal, erect to pendulous; capitula pedunculate, heterogamous, rarely homogamous, discoid or radiate; receptacle epaleate; involucre multiseriate, phyllaries erect to recurved downward. Florets dimorphic or rarely isomorphic; marginal florets female, with staminodes, corolla bilabiate, outer lip expanded or not, inner lip developed to very reduced, or true ray; central florets bisexual, corolla bilabiate; or all the florets bisexual, corolla bilabiate, sub-bilabiate (4+1 corolla lips), or ligulate; anther apical appendages acute, tails smooth; style bilobed, dorsally papillose. Cypselae truncate to attenuate at the apex, glabrous; pappus of plumose, capillary bristles.

Pollen Description. Mutisia (101 of Figs. 101-106) (Telleria & Katinas, 2004). The two species examined (M. kurtzii R. E. Fr. and M. retrorsa Cav.) have pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate (see also Cabrera, 1965; Skvarla & Turner, 1966; Heusser, 1971; Parra & Marticorena, 1972; Skvarla et al., 1977). Note: pollen of Mutisia is highly variable in size and exine thickness (Telleria & Katinas, in press).

Habitat and Distribution. Genus of 63 South American species, from the Andes of Colombia to southern Argentina and Chile, also southern Brazil and adjacent regions.

Species list

1. Mutisia acerosa Poepp. ex Less., Syn. gen. Compos.: 107. 1832. TYPE: Chile. Near Las Cruces et La Guardia de los Hornillos, E. F. Poeppig 26 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, El Maiten, C. Jiles 2451 (LP).

2. Mutisia acuminata Ruiz & Pav., Syst. veg. ft. peruv, chil. 1: 192. 1798. TYPE: Peru. Ceuchin, May 1779, s. leg. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Volcan, A. L. Cabrera et al. 13894 (LP).

3. Mutisia alata Hieron., Bot. Jahrb. Syst. 19: 73. 1894. TYPE: Ecuador. Inter Huasi-huaico et Contrayerba, Lehmann 4571 (isotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cajamarca: Cajamarca, Huambos, Ochoa 1594 (LP).

4. Mutisia andersonii Sodiro ex Hieron., Bot. Jahrb. Syst. 29: 80. 1900. TYPE: Ecuador. Prov. Pichincha: Cerro Pichincha, E. Sodiro s.n. (neotype, designated by Harling, 1991: 74, W not seen). Additional specimen examined: Ecuador. Prov. Chimborazo: camino Rio Bamba a Huamboya- De Pungala a Cusfiipaccha, R. Scolnick 1532 (LP).

5. Mutisia araucana Phil., Anales Univ. Chile 41: 737. 1872. TYPE: Chile. Prov. Arauco: pinares de Nahuelbuta, s.d., H. Volckmann s.n. (holotype, SGO not seen, SGO photo at LP!).

6. Mutisia arequipensis Cabrera, Opera Lilloana 13: 171. 1965. TYPE: Peru. Dept. Arequipa: A. Weberbauer 6857 (isotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Arequipa: Prov. Cailloma, distr. Coporaque, valle de Colca, ca. 100 km N de Arequipa, J. Treacy 840 (WIS).

7. Mutisia brachyantha Phil., Anales Univ. Chile 85: 825. 1894. TYPE: Chile. Andes, Talca, s.d., L. H. Silva s.n. (type, probably SGO). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Morro Blanco, C. Jiles 1590 (LP).

8. Mutisia burkartii Cabrera, Darwiniana 20: 210. 1976. TYPE: Argentina. Prov. Neuquen: Nuco Pehuen, 25 km al W de Loncopue, M. Gentili 215 (holotype, LP!).

9. Mutisia campanulata Less., Linnaea 5: 269. 1830. TYPE: Brazil: s.d., Sello s.n. (holotype, B not seen, B photo F n. 15925 at LP!). Additional specimen examined: Argentina. Prov. Misiones: El Paraiso, M. A. Torres 67 (LP).

10. Mutisia cana Poepp. & Endl., Nov. gen. sp. pl. 1: 17. 1835. TYPE: Chile. Rio Colorado, Santa Rosa, s.d., s. leg. (type, not localized). Additional specimen examined: Chile. Prov. Santiago: Penalolen, E. Navas 959 (LP).

11. Mutisia clematis L. f., Suppl. pl.: 373. 1781 (1782). TYPE: Habitat in Nova Granada, J. C. Mutis 21 (holotype, LINN 1004.1 not seen). Additional specimen examined: Colombia. Dept. Boyaca: Rio Gomeza (Rio Arzobispo), 7-10 km al E de Socha, F. R. Fosberg 22294 (LP).

12. Mutisia coccinea A. St.-Hil., Voy. distr, diam. 1: 386. 1833. TYPE: Brazil. State Silo Paulo: d'Hytu, St. Hilaire 1474bis (holotype, P not seen). Additional specimen examined: Argentina. Prov. Misiones: Loreto, G. Gruner 516 (LP).

13. Mutisia cochabambensis Hieron., Bot. Jahrb. Syst. 19: 74. 1894. TYPE: Bolivia. Cochabamba, M. Bang 881 (isotype, WIS).

14. Mutisia comptoniaefolia Rusby, Mere. Torrey Bot. Club 3: 65. 1893. TYPE: Bolivia. Talca Chugiaguillo, M. Bang 795 (isotype, WIS!).

15. Mutisia decurrens Cav., Icon. 5: 65. 1799. TYPE: Chile et Cordillera del Planchon, s.d., M. Nee s.n. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Neuquen: Canadon de la Mosca, Pampa del Toro, G. Dawson 3287 (LP).

16. Mutisia discoidea Harling, Fl. Ecuador 42: 70. 1991. TYPE: Ecuador. Prov. Canar: road Canar-El Tambo, ca. 4 km from Canar, J. Madsen 50281 (holotype, AAU not seen).

17. Mutisia friesiana Cabrera, Notas Mus. La Plata, Bot. 13: 19. 1948. TYPE: Argentina. Prov. Salta: Dept. Poma, Quebrada de Cobres, A. L. Cabrera 8716 (holotype, LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Susques, Quebrada de Potrero, L. Catalano 25 (LP).

18. Mutisia grandiflora Humb. & Bonpl., P1. aequinoct. 1: 177. 1808 TYPE: Colombia. Andibus Quinduensibus, inter urbem Ibague et Carthago, s.d., F. Humboldt & A. Bonpland s.n. (holotype, P not seen). Additional specimen examined: Colombia. Dept. del Valle: cordillera central, Hoya del Rio Bugalagrande, Loma de Barragan, entre la poblacidn y Albania, J. Cuatrecasas 20917 (F).

19. Mutisia hamata Reiche, Anales Univ. Chile 115: 103. 1904. TYPE: Chile. Prov. Tarapaca: Amincha, s.d., s. leg. (type, probably SGO). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, camino de E1 Angosto al Chana, A. L. Cabrera et al. 22495 (LP).

20. Mutisia hastata Cav., Icon. 5: 64. 1799. TYPE: Peru. Cordillera del Planchon, s.d., M. Nee s.n. (isotype, CONC!). Additional specimen examined: Peru. Dept. Lima: Prov. Canta, Lachaqui, Toma, G. Vilcapoma S. 1307 (LP).

21. Mutisia hieronymii Sodiro ex Cabrera, Notas Prelim. Mus. La Plata 3: 167. 1934. TYPE: Ecuador. Prov. Napo-Pastaza: Cuyuja-Baeza, Feb 1905, A. Sodiro s.n. (isotype, BAF not seen).

22. Mutisia homoeantha Wedd., Chlor. andina 1: 21. 1855. TYPE: Bolivia. Chuquisaca, M. Weddell 4075 (holotype, P not seen). Additional specimen examined: Argentina. Prov. Salta: Dept. Santa Victoria, Lizoite, T. Meyer 33143 (LP).

23. Mutisia ilicifolia Cav., Icon. 5: 63. 1799. TYPE: Chile. Prov. Coquimbo: Coquimbo, s.d., M. Nee s.n. (lectotype, designated by Cabrera, 1965: 198, MA not seen, MA photo at LP!). Additional specimen examined: Chile. Prov. Curico: Cordillera del Planchon, E. Barros 1751 (LP).

24. Mutisia intermedia Hieron., Bot. Jahrb. Syst. 19: 72. 1894. TYPE: Colombia. Narifio: cerca de Tuquerres, s.d., J. Idrobo & M. Ospina 2421 (neotype, designated by Diaz-Piedrahita & Velez-Nauer, 1993: 96, COL not seen).

25. Mutisia involucrata Phil., Linnaea 28: 710. 1856. TYPE: Chile in collibus subandinis, Talcaregue, Feb 1831, C. Gay 786 (type, probably P). Additional specimen examined: Chile. Prov. Curico: Teno, La Montana, E. Barros 3804 (LP).

26. Mutisia kurtzii R. E. Fr., Nova Acta Regiae Soc. Sci. Upsal. 1: 94. 1905. TYPE: Argentina. Prov. Jujuy: Tilcara, Quebrada de Juella, F. Claren 11732 (isotype, BAF not seen, BAF photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Humahuaca, cerro de Zenta, Feb 1931, E. Budin s.n. (LP*).

27. Mutisia lanata Ruiz &Pav., Syst. veg. ft. peruv, chil. 1: 192. 1798. TYPE: Peru: s.d., s. leg. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Bolivia. Dept. Yungas: Tusilmaya a Chlumani, R. Scolnick & R. Luti 519 (LP).

28. Mutisia lanigera Wedd., Chlor. andina 1: 16. 1855. TYPE: Bolivia. Provincie de Carangas, C. D'Orbigny 1341 (holotype, P not seen, P photo at LP!). Additional specimen examined: Chile. Prov. Antofagasta: Toconce, M. Ricardi & C. Marticorena 4677/1062 (LP).

29. Mutisia latifolia D. Don, Trans. Linn. Soc. London 16: 270. 1830. TYPE: Chile. Prov. Valparaiso: Valparaiso, s.d., D. Bridges s.n. (holotype, G not seen, G photo F n. 28874 at LP!). Additional specimen examined: Chile. Prov. Santiago: Cuesta La Dormida, G. Looser 1323 (LP).

30. Mutisia ledifolia Decne. ex Wedd., Chlor. andina 1: 20. 1855. TYPE: Bolivia. Dept. Potosi: cumbre de la Cuesta de Lagunillas, C. D'Orbigny 1272 (lectotype, designated by Cabrera, 1965: 134, P not seen, P photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Rinconada, Quebrada de Fundiciones, A. L. Cabrera 9438 (LP).

31. Mutisia lehmannii Hieron., Bot. Jahrb. Syst. 19: 73. 1894. TYPE: Ecuador. Prov. Azuay: Cerro Soldados, W of Sayausi, G. Harling et al. 8743 (neotype, designated by Harling, 1991: 67, GB not seen). Additional specimen examined: Ecuador. Prov. Azuay: area nacional de Recreacion "Cajas," Sayausi-Molleturo road W of Cuenca, L. J. Dorr & I. Valdepino 6382 (F).

32. Mutisia linearifolia Cav., Icon. 5: 66. 1799. TYPE: Chile. Cordillera del Planchon, s.d., M. Nee s.n. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Neuquen: Cordillera del Viento, J. Frenguelli 521 (LP).

33. Mutisia linifolia Hook., Bot. misc. l: 12. 1830. TYPE: Argentina. Prov. Mendoza: camino de las Minas de Uspallata, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Las Heras, Quebrada del Toro, 1934, A. Perez Moreau (LP).

34. Mutisia lutzii G. M. Barroso, Mere. Inst. Oswaldo Cruz 53: 357. 1955. TYPE: Brazil. State Minas Geraes: Serra do Caparao, A. Lutz 1200 (holotype, R not seen).

35. Mutisia macrophylla Phil., Anales Univ. Chile 18: 62. 1861. TYPE: Chile. Entre Lota y Chillan, s.d., Pearce s.n. (type, probably SGO). Additional specimen examined: Chile. Prov. Chillan: San Carlos, San Fabian, E. Barros 1772 (LP).

36. Mutisia magnifica C. Ulloa & P. Jorg., Novon 6: 131. 1996. TYPE: Ecuador. Loja: Cerro Uritusinga, Loja-La Palma, km 18.4, P. M. Jorgensen et al. 1010 (holotype, MO!).

37. Mutisia mandoniana Wedd. ex Cabrera, Opera Lilloana 13: 163. 1965. TYPE: Bolivia. Prov. Larecaja: inter Sorata et Lacatia, G. Mandon 8 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Bolivia. Near Tarcacama, Larecaja, M. Cardenas 4869 (F).

38. Mutisiamathewsii Hook. & Am.,Comp. Bot. Mag. 1: 107. 1835. TYPE: Peru: s.d., Mathews 1119 (isotype, P not seen, P photo F n. 38030 at LP!). Additional specimen examined: Peru. Dept. Junin: Yauli, Macbride & Featherstone 907 (LP).

39. Mutisia microcephala Sodiro ex Cabrera, Notas Prelim. Mus. La Plata 3: 168. 1934. TYPE: Ecuador. Cuyuja-Baeza, Feb 1905, A. Sodiro s.n. (isotype, BAF not seen, BAF photo at LP!). Additional specimen examined: Ecuador. Prov. Imbabura: Lake Cuicocha, C. W. Penland & R. H. Summers 731 (F).

40. Mutisia microneura Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 121. 1953. TYPE: Ecuador. Prov. Chanar: Tipacocha, C. W. Penland & R. H. Summers 1007 (holotype, F not seen, F photo at LP!).

41. Mutisia microphylla Willd. ex DC., Prodr. 7: 6. 1838. TYPE: Ecuador. Inter Quito et Pichincha, s.d., F. Humboldt & A. Bonpland s.n. (holotype, G not seen). Additional specimen examined: Ecuador. Prov. Pichincha: faldas de Pichincha, C. Firmin 524 (F).

42. Mutisia ochroleuca Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 122. 1953. TYPE: Colombia. Dept. Narifio: Volcan de Chiles, Tambo, NW shoulder of Volcan de Chiles, J. A. Ewan 16089 (holotype, F!).

43. Mutisia oligodon Poepp. & Endl., Nov. gen. sp. pl. 1: 18. 1835. TYPE: Chile. Chile meridionalis, Antuco, Castillo de Ballenar, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Rio Negro: Bariloche, L. R. Parodi 11564 (LP).

44. Mutisia orbignyana Wedd., Chlor. andina 1: 22. 1855. TYPE: Bolivia. St. Bartolo, C. D'Orbigny 1387 (holotype, P not seen, P photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Valle Grande, Caspala, A. Burkart & N. S. Troncoso 11700 (LP).

45. Mutisia pulcherrima Muschl., Bot. Jahrb. Syst. 50: 102. 1913. TYPE: Peru. Prov. Chachapoyas: inter Tambos Bajazan et Almirante, A. Weberbauer 4444 (holotype, B not seen, B photo F n. 15495 at LP!). Additional specimen examined: Peru. Prov. Chachapoyas: Dept. Amazonas, Jalca de Calla-Calla, A. Sagastegui A. 7469 (LP).

46. Mutisia rauhii Ferreyra, Bol. Soc. Peruana Bot. 8: 75. 1980. TYPE: Peru. Dept. Cuzco: al E de la Cordillera de Salcantay, W. Rauh 1451 (holotype, USM!).

47. Mutisia retrorsa Cav., Icon. 5: 65. 1799. TYPE: Argentina. Prov. Santa Cruz: Puerto Deseado, s.d., M. Nee s.n. (isotype, CONC!). Additional specimen examined: Argentina. Prov. Santa Cruz: Lago Argentino, Jan 1902, R. Hauthal s.n. (LP).

48. Mutisia rimbachii Sodiro ex S. K. Harris, Rhodora 35: 257. 1933. TYPE: Ecuador. Andes, 1850 1857, R. Spruce 5456 (lectotype, designated by Harling, 1991: 59, BM not seen; isolectotype, NY not seen, NY photo at LP!). Additional specimen examined: Ecuador. Prov. Tungurahua: Mocha, alrededores del pueblo, P. Villacres 234 (F).

49. Mutisia rosea Poepp. ex Less., Syn. gen. Compos.: 106. 1832. TYPE: Chile. Inter Las Cruces and Guardia de los Hornillos, E. F. Poeppig 113 (27) (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Santiago: San Gabriel, G. Montero O. 515 (F).

50. Mutisia saltensis Cabrera, Darwiniana 9: 373. 1951. TYPE: Argentina. Prov. Salta: Dept. Oran, Cerro Santa Barbara, Carmelich et al. 190 (holotype, LP!). 51. Mutisia sinuata Cav., Icon. 5: 66. 1799. TYPE: Chile. Cordilleras del Planchon y Portillo, s.d., M. Nee s.n. (isotype, CONC!). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Tunuyan, Valle del Alto Tunuyan, A. Ruiz Leal 2146 (LP).

52. Mutisia sodiroi Hieron., Bot. Jahrb. Syst. 29: 79. 1900. TYPE: Ecuador. Prov. Pichincha, Cerro Pichincha, s.d., Sodiro s.n. (neotype, designated by Harling, 1991: 62, G not seen). Additional specimen examined: Colombia. Dept. Narifio: cordillera occidental, Oscurana, on S slope of Volcan Gualcala, 14 km E of Piedrancha, F. R. Fosbert 21188 (F).

53. Mutisia speciosa Aiton ex Hook., Bot. Mag. 54: 2705. 1827. TYPE: Brazil: s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Misiones: Arroyo Chapa, San Ignacio, G. Gruner 1077 (LP).

54. Mutisia spectabilis Phil., Linnaea 33: 107. 1864. TYPE: Chile. Prov. Aconcagua: Calabozo, Catemu, Sept 1860, R. A. Philippi s.n. (holotype, SGO not seen, SGO photo at LP!).

55. Mutisia spinosa Ruiz &Pav., Syst. veg. ft. peruv, chil. 1: 193. 1798. TYPE: Chile. Arauco, Colo-Colo, s.d., s. leg. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Argentina. Prov. Rio Negro: Bariloche, A. L. Cabrera & M. M. Job 51 (LP).

56. Mutisia splendens Renjifo, Anales Univ. Chile 65: 299. 1884. TYPE: Chile: Dept. Quillota, Vichiculen, Dec 1881, C Renjifo (type, probably SGO). Additional specimen examined: Chile. Prov. Santiago, Claude-Joseph 974 (US).

57. Mutisia stuebelii Hieron., Bot. Jahrb. Syst. 21: 367. 1895. TYPE: Colombia. Cerro Munchique, cerca de Popayan, A. Stubel 318a (holotype, B not seen, B photo F n. 15951 at LP!). Additional specimen examined: Colombia. Dept. del Cauca: cordillera central, cabeceras del rio Palo, quebrada de Santo Domingo, campamento de los quineros, J. Cuatrecasas 19156 (F).

58. Mutisia subspinosa Cav., Icon. 5: 64. 1799. TYPE: Peru. Guamauranga, Planch6n, s.d., M. Nee s.n. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Las Heras, Estancia San Isidro, cerca del Mogote Aspero, A. Ruiz Leal 22532 (LP).

59. Mutisia subulata Ruiz &Pav., Syst. veg. ft. peruv, chil. 1: 193. 1798. TYPE: Chile. Concepcion, s.d., s. leg. (holotype, MA not seen). Additional specimen examined: Argentina. Prov. Neuquen: Cordillera del Viento, J. Frenguelli 516 (LP).

60. Mutisia tridens Poepp. ex Less., Syn. gen. Compos.: 105. 1832. TYPE: Chile. Prov. Bio Bio: Antuco, E. F. Poeppig 220 (isotype, NY not seen, NY photo at LP!).

61. Mutisia venusta S. F. Blake, Proc. Biol. Soc. Wash. 46: 191. 1933. TYPE: Peru. Cuzco, Pifiasniocj, Panticalla Pass, O. F. Cook & G. B. Gilbert 1235 (holotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cuzco: Prov. Urubamba, entre Piri y Penas, C. Vargas C. 4420 (LP).

62. Mutisia vieia J. Kost., Blumea 5: 670. 1945. TYPE: Bolivia. Comarapa, T. Herzog 1974 (isotype, LP!). Additional specimen examined: Bolivia. Dept. Cochabamba: Pocona, M. Cardenas 2256 (LP).

63. Mutisia wurdackii Cabrera, Opera Lilloana 13: 43. 1965. TYPE: Peru. Dept. Amazonas: Prov. Bongara: 2-4 km WSW of Pomacocha, J. J. Wurdack 840 (holotype, LP!). Additional specimen examined: Peru. Dept. Amazonas: Prov. Chachapoyas, entre Ingenio y Pomacocha, A. Lopez et al. 4303 (LP).

Literature. Cabrera (1965, 1976), Ferreyra (1980), Harling 1991), Ulloa Ulloa and Jorgensen (1996); Tellaria and Katinas (in press).

Myripnois Bunge

Enum. pl. China bor.: 38. 1833 (1831). TYPE: Myripnois dioica Bunge. 33 of Figs. 31-34.

Etymology. From the Greek mvri, ointment or resin, and pnois, to smell, to have an odor, referring to the aromatic compounds in the plant.

Shrubs dioecious. Leaves clustered on brachyblasts; subsessile; blades ovate-elliptic, palmately, pinnately to three-veined, margin entire, slightly pubescent on both surfaces. Capitulescences monocephalous, terminal or axillar; capitula sessile on the brachyblasts, homogamous, ligulate, five- to seven-flowered; receptacle epaleate; involucre uniseriate, phyllaries ca. 5. Florets isomorphic, male or female, corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally papillose. Cypselae pubescent; pappus of scabrid, capillary bristles, absent in male florets.

Pollen Description (102 of Figs. 101-106). Pollen spheroidal-subprolate, spheroidal to elliptic in equatorial view, circular in polar view, medium size, P x E = (38-46 x 30-36) [micro]m. Tricolporate, colpi long with psilate membrane. Exine Mutisia type, 5 6 [micro]m thick. Ratio ectosexine/endosexine: ca. 1:1. Nexine 2.5 [micro]m thick (see also Lin et al., 2005).

Habitat and Distribution. Monotypic genus endemic to China.

Species list

Myripnois dioica Bunge, Enum. pl. China bor.: 38. 1833 (1831). TYPE: not localized. Additional specimens examined: China. Hopei: Peking, Wang 20106 (NY); id., s.d., Sheehan s.n. (NY *).

Literature. Von Bunge (1835).

Neblinaea Maguire & Wurdack

Mem. New York Bot. Gard. 9: 391. 1957. TYPE: Neblinaea promontoriorum Maguire & Wurdack. 34 of Figs. 31-34.

Etymology. Named for Cerro de la Neblina. This mountain, which forms part of the border between Brazil and Venezuela, contains the tallest peak in Brazil.

Shrubs. Leaves alternate, mostly clustered at stem apex; petiolate, petioles clasping; blades oblanceolate to spathulate, pinnately veined, margin entire, partially revolute, glabrous, rarely puberulent beneath. Capitulescences cymose, two- to nine-headed, subterminal to lateral, rarely monocephalous; capitula pedunculate, homogamous, bilabiate, two- to five-flowered; receptacle epaleate; involucre multiseriate, cylindrical, phyllaries glabrous. Florets isomorphic, bisexual, corolla bilabiate, glabrous; anthers tan to brown, often white at the apex, exserted, apical appendages acuminate, tails slightly papillose, connate with the tails of adjacent stamens; style bifid, branches dorsally smooth. Cypselae glabrous; pappus of slightly scabrid bristles, half are capillary, half somewhat paleaceous and longer.

Pollen Description (103 of Figs. 101-106). Pollen subprolate, elliptic in equatorial view, circular in polar view, large size, P x E = (62 70x46-55) [micro]m. Tricolporate, colpi long with psilate membrane. Exine Mutisia type, microechinate, ca. 7 [micro]m thick, ectosexine slightly columellate, slightly thickened at the poles. Ratio ectosexine/endosexine: 1:2. Nexine ca. 2 [micro]m thick. SEM: tectum perforate. Note: pollen similar to that of Chimantaea cinerea, C. huberi, Stifftia uniflora, Stenopadus campestris, S. colveii, and S. huachamacari. (see also Carlquist, 1957b; Telleria, 2008).

Habitat and Distribution. Neblinaea is monotypic and endemic to Cerro de la Neblina, where it occurs on both sides of the Venezuelan-Brazilian frontier.

Species list

1. Neblinaea promontoriorum Maguire & Wurdack, Mere. New York Bot. Gard. 9: 392. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 37009 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, N. T. Silva & Brazzao 60895 (NY*).

Literature. Pruski (1991, 1997).

Nouelia Franch.

J. Bot. (Morot) 2: 67. 1888. TYPE: Nouelia insignis Franch. 35 of Figs. 35-38.

Etymology. In honor of M. A. -A. Nouel (d. 1887), naturalist and director of the Musde d'Orleans, France.

Shrubs or small trees. Leaves alternate; petiolate; blades ovate-elliptic, pinnately veined, margin entire, tomentose beneath. Capitulescences monocephalous, rarely two- to three-headed, terminal; capitula short-pedunculate, homogamous, radiate; receptacle epaleate; involucre multiseriate, six- to ten-seriate. Florets dimorphic, bisexual; marginal florets with corollas bilabiate, outer lip, expanded; central florets with corolla actinomorphic, tubular-campanulate, deeply five-lobed; anther apical appendages apiculate, tails papillose; style bifid, branches dorsally smooth. Cypselae villose; pappus of scabrid, capillary bristles, the outermost shorter.

Pollen Description. Pollen subprolate--prolate, elliptic in equatorial view, circular in polar view, large size, P x E = (62-69 x40-50) [micro]m. Tricolporate, colpi long with margin and psilate membrane, mesoapermrate. Exine Mutisia type, scabrate, 6-8 [micro]m, ectosexine slightly columellate, thick internal tectum. Ratio ectosexine/endosexine: 2:1. Nexine ca. 2.5 [micro]m thick.

Habitat and Distribution. Monotypic genus endemic to Sichuan Yun-nan region in China.

Species list

1. Nouelia insignis Franch., J. Bot. (Morot) 2: 68. 1888. TYPE: China. Yun-Nan: Tapin-tze, J. M. Delavay 2498, 619, 245 (syntypes, probably P). Additional specimen examined: China, Maire 2516 (NY*).

Observation. Most populations of Nouelia insignis are seriously threatened and are even at the brink of extinction in China (Heng, 1995). In recent decades, N. insignis has become endangered due to rapid habitat fragmentation. Most of the extant populations consist of fewer than 50 individuals, and no more than 5,000 individuals survive in the wild (Luan et al., 2006).

Literature. Franchet (1888), Heng (1995), Freire et al. (2002).

Oldenburgia Less.

Linnaca 5: 252. 1830. TYPE: Oldenburgia paradoxa Less. 36 of Figs. 35-38.

Etymology. In honor of the Swedish soldier and traveler Franz Pehr Oldenburg (1740 1774) who accompanied Thunberg and Masson on some of their travels in the Cape in the early 1770s.

Shrubs or small trees, often cushion-like shrubs. Leaves alternate, clustered at the end of the branches; subsessile; blades obovate to elliptic, coriaceous, pinnately veined, margin entire, revolute, glabrous above, tomentose beneath. Capitulescences monocephalous or two- to five-headed, corymbose; capitula sessile to pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate, ovoid, globose-urceolate to cyathiform. Florets dimorphic; marginal florets female, well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages long-apiculate, tails smooth; style bilobed, branches dorsally rugulose to papillose. Cypselae glabrous to villose; pappus of plumose to barbellate capillary bristles, widened at the apex.

[FIGURES 35-38 OMITTED]

Pollen Description. Pollen not seen.

Habitat and Distribution. Genus of four species from southern Africa (Cape Province).

Species list

1. Oldenburgia grandis (Thunb.) Baill., Hist. pl. 8: 97. 1882. Basionym: Arnica grandis Thunb. TYPE: Cap, s.d., Masson s.n., herb. Thunb. (holotype, UPS microfiche not seen). Additional specimen examined: Africa. South Africa, H. Humbert 19298 (MO).

2. Oldenburgia intermedia Bond, S. African J. Bot. 53: 496. 1987. TYPE: Africa. South Africa: Bergriviernek, Jonkershoek, P. Bond 1723 (isotype, K not seen). Additional specimen examined: South Africa, Western Cape Province, Tulbagh District, VoElvlei Mts., Saddle Mt. on the way to Ontongskpo from Voelvlei, S. Ortiz 832a (SANT).

3. Oldenburgia papionum DC., Prodr. 7: 12. 1838. TYPE: Berge bei Tulbagh RIII, Drege 1835 (lectotype, designated by Bond, 1987: 499, G not seen). Additional specimen examined: Africa. South Africa, Western Cape Province, Tulbagh District, VoElvlei Mts., Saddle Mt. on the way to Ontongskpo from Voelvlei, S. Ortiz 831a (SANT).

4. Oldenburgia paradoxa Less., Linnaea 5: 252. 1830. TYPE: Lectotype, designated by Bond, 1987: 496, the illustration in Linnaea 5: Table 3, Figs. 67-70 (1830).

Observation. In the molecular analysis of Funk et al. (2005), Oldenburgia appears nested in the subfamily Carduoideae, sister to Tarchonantheae and Cardueae. Ortiz (2006b) suggested that it could be established as a tribe containing only the genus Oldenbugia. Our observations show that the species of this genus have the typical mutisioid style.

Literature. Bond (1987).

Onoseris Willd.

Sp. pl. 3: 1702. 1803. TYPE: Atractylis purpurea L. f. [= Onoseris purpurea (L. f.) S. F. Blake]. 37 of Figs. 35-38.

Caloseris Benth., Pl. hartw.: 88. 1841. TYPE: Caloseris rupestris Benth. [= Onoseris onoseroides (Kunth) B. L. Rob.].

Cataleuca Koch & Fintelm., Wochenschr. Gartnerei Pflanzenk. 2: 163. 1859, nom. nudum.

Centroclinium D. Don, Trans. Linn. Soc. London 16: 254. 1830. TYPE: Centroclinium albicans D. Don [= Onoseris albicans (D. Don) Ferreyra].

Chaetachlaena D. Don, Trans. Linn. Soc. London 16: 256. 1830. TYPE: Chaetachlaena odorata D. Don [= Onoseris odorata (D. Don) Hook. & Am.].

Cladoseris (Less.) Spach, Hist. nat. veg. 10: 35. 1841. TYPE: Cladoseris annua (Less.) Spach, based on Onoseris annua Less.

Cursonia Nutt., Trans. Amer. Philos. Soc. 7: 422. 1841. TYPE: Cursonia peruviana Nutt. [= Onoseris odorata (D. Don) Hook. & Am.].

Hilairia DC., Prodr. 7: 33. 1838, nom. nudum.

Hipposeris Cass., Dict. sci. nat. 33: 464. 1824. [= Onoseris subgen. Hipposeris

(Cass.) Less., Syn. gen. Compos.: 119. 1832]. TYPE: Onoseris salicifolia Kunth (lectotype designated by Sancho, 2004).

Isotypus Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 8. 1818. TYPE: Isotypus onoseroides Kunth [= Onoseris onoseroides (Kunth) B. L. Rob.].

Pereziopsis J. M. Coult., Bot. Gaz. (London) 20: 52. 1895. TYPE: Pereziopsis donnell-smithii J. M. Coult. [= Onoseris donnell-smithii (J. M. Coult.) Ferreyra].

Rhodoseris Turcz., Bull. Soc. Imp. Naturalistes Moscou 24: 95. 1851. TYPE: Rhodoseris conspicua Turcz. [= Onoseris onoseroides (Kunth) B. L. Rob.].

Schaetzellia Klotzch, Allg. Gartenzeitung. 17: 82. 1849. TYPE: Schaetzellia deckeri Klotzsch [= Onoseris onoseroides (Kunth) B. L. Rob.].

Seris Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 1: 139. 1807. TYPE: Seris onoseroides Willd. [= Onoseris onoseroides (Kunth) B. L. Rob.].

Etymology. From the Greek onos, donkey, and seris, seridos, chicory, lettuce, probably because the rosulate leaves, lanceolate and tomentose beneath, which resemble the ears of a donkey.

Herbs annual or perennial or subshrubs, branched to unbranched. Leaves alternate, often clustered near the base; sessile to long-petiolate; blades ovate to elliptic, linear, hastate to rarely orbicular, pinnately to palmately veined, margin entire to dentate, glabrescent above, densely pubescent beneath. Capitulescences monocephalous, terminal, sometimes on scapes, corymbose to paniculate; capitula heterogamous, radiate or discoid; involucre multiseriate; receptacle epaleate, pilose to setose. Florets usually dimorphic, when isomorphic, all slightly zygomorphic with corollas subbilabiate (4+1 corolla lips), lip not expanded; when dimorphic marginal florets female, with staminodes, corolla bilabiate to sub-bilabiate (3+1 corolla lips), outer lip expanded or not; central florets actinomorphic, tubular-funnelform, corolla shallowly five-lobed, lobes recurved, or slightly zygomorphic, five-lobed, lobes straight, with one or two deeper insicions; anther apical appendages acute, tails smooth to papillose, filaments usually papillose at the base; style bifid, branches dorsally papillose, sometimes glabrescent. Cypselae pubescent to glabrescent; pappus of all capillary, scabrid bristles, unequal in length, sometimes the inner ones longer, wider, and flattened.

Pollen Description. Onoseris alata Rusby (Telleria & Katinas, 2004). Pollen subprolate--prolate, large size, tricolporate, exine Mutisia type, microechinate. Note: pollen of O. odorata has polar caps.

Habitat and Distribution. Genus of 31 species from southern Mexico to northwestern Argentina, with the highest concentration in the western slopes of the Andes of Peru and few species in Brazil.

Species list

1. Onoseris acerifolia Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 6. 1818. TYPE: Peru. Paramo de Yamoca, F. Humboldt & A. Bonpland 2568 (holotype, P not seen). Additional specimen examined: Bolivia. Santa Cruz, Steinbach 8200 (US).

2. Onoseris alata Rusby, Descr. S. Amer. pl.: 163. 1920. TYPE: Bolivia. Dept. La Paz: Coroico, Yungas, M. Bang 2365 (isotypes, US!, GH, K not seen, GH and K photos at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Humahuaca, M. Venturi 8359 (US).

3. Onoseris albicans (D. Don) Ferreyra, J. Arnold Arbor. 25: 389. 1944. Basionym: Centroclinium albicans D. Don. TYPE: Peru: s.d., H. Ruiz & J. A. Pavon s.n. (holotype, P not seen). Additional specimen examined: Peru. Apurimac, Vargas 5816 (LP).

4. Onoseris amplexicaulis Ferreyra, J. Arnold Arbor. 25: 364. 1944. TYPE: Peru. Dept. Lima: Matucana, J. F. Macbride & W. Featherstone 310 (isotypes US!, GH not seen, GH photo at LP!). Additional specimen examined: Peru. Dept. Lima: Matucana, J. F. Macbride & W. Featherstone 131 (F).

5. Onoseris annua Less., Linnaea 5:341.1830. TYPE: Peru. Lima, s.d., J. Dombey s.n. (isotypes, B, F not seen, B photo F n. 15887 in LP!). Additional specimen examined: Peru. Ancash, R. Ferreyra 73 76 (US).

6. Onoseris brasiliensis Cabrera, Bol. Mus. Bot. Munic. 19: 1. 1975. TYPE: Brazil. Mato Grosso: Rondonopolis, G. Hatschbach 34719 (holotype, LP!).

7. Onoseris cabrerae Ferreyra, Bol. Soc. Argent. Bot. 19: 19. 1980. TYPE: Peru. Dept. Amazonas: Chachapoyas, R. Ferreyra & C. Acleto 15339 (holotype, USM!, USM photo LP!).

8. Onoseris castelnaeana Wedd., Chlor. andina 1: 10. 1855. TYPE: Peru. Cusco, 1847, M. F. Castelnau s.n. (holotype, P not seen, P photo F n. 38105 at LP!). Additional specimen examined: Peru. Apurimac, Vargas 404 (LP).

9. Onoseris chrysactinioides Sagest. & M. O. Dillon, Brittonia 37: 8. 1985. TYPE: Peru. Dept. Cajamarca: Contumaza A. Sagastegui et al. 10839 (holotype, HUT!, HUT photo at LP!; isotype MO!). Additional specimen examined: Peru. Dept. Cajamarca: Contumaza, Becker & Terrones 1011 (US).

10. Onoseris costaricensis Ferreyra, J. Arnold Arbor. 25: 367. 1944. TYPE: Costa Rica. San Ramon, Brenes 6520 (holotype, F not seen). Additional specimen examined: Costa Rica. Pittier 10596 (US).

11. Onoseris cumingii Hook. & Am., Comp. Bot. Mag. 1: 103. 1835. TYPE: Peru. Lima, Cuming 995 (holotype, K not seen, K photo at LP!). Additional specimen examined: Peru. Piura, O. Haught 30 (US).

12. Onoseris donnell-smithii (J. M. Coult.) Ferreyra, J. Arnold Arbor. 25:368.1944. Basionym: Pereziopsis donnell-smithii J. M. Coult. TYPE: Guatemala. Santa Rosa: Rio de los Esclavos, E. T. Heyde & E. J. D. S. Lux 4527 (isotype, US!). Additional specimen examined: E1 Salvador. Santa Ana, Standley 19701 (US).

13. Onoseris drakeana Andre, Rev. Hort.: 181. 1882. TYPE: Colombia. Cauca, E. F. Andre 2917 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Colombia. Narifio, Uribe 5355 (US).

14. Onoseris fraterna S. F. Blake, J. Wash. Acad. Sci. 33: 368. 1943. TYPE: Bolivia. Dept. La Paz: Sur Yungas, B. A. Krukoff 10266 (holotype, US!). Additional specimen examined: Bolivia. Dept. La Paz: Sur Yungas, Fournet 472 (US).

15. Onoseris gnaphalioides Muschl., Bot. Jahrb. Syst. 50: 94. 1913. TYPE: Peru. Ancash, Caraz, A. Weberbauer 3012 (isotype, F not seen). Additional specimen examined: Peru. Ayacucho, Hjerting 1415 (LP).

16. Onoseris hastata Wedd., Chlor. andina 1: 9. 1855. TYPE: Bolivia. Dept. Chuquisaca: Monte Curl, H. A. Weddell 3763 (isotype, G not seen). Additional specimen examined: Argentina. Prov. Jujuy: Tumbaya, Volcan, A. L. Cabrera et al. 18286 (LP).

17. Onoseris humboldtiana Ferreyra, Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, Bot. 11: 1. 1959. TYPE: Peru. Ica: Nazca, Lomas de San Nicolas, R. Ferreyra 13400 (holotype, USM!, USM photo at LP!).

18. Onoseris hyssopifolia Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 7. 1818. TYPE: Ecuador. Imbabura, F. Humboldt & A. Bonpland 2200 (holotype, P not seen; isotype, B not seen, B photo F n. 15893 at LP!). Additional specimen examined: Ecuador. Azuay, King 6648 (US).

19. Onoseris linearifolia Sagest., Phytologia 57: 415. 1985. TYPE: Peru. Dept. Cajamarca: Cajabamba, A. Sagastegui et al. 11264 (holotype, HUT!, HUT photo at LP!). Additional specimen examined: Peru. Dept. Cajamarca: Cajabamba, I. Sanchez Vega et al. 956 (US).

20. Onoseris lopezii Ferreyra, Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, Bot. 11: 4. 1959. TYPE: Peru. La Libertad: Santiago de Chuco, A. Lopez Miranda 439 (holotype, USM!, USM photo at LP!). Peru. Cajamarca, Hudson 1073 (MO).

21. Onoseris macbridei Ferreyra, Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, Bot. 1: 2. 1949. TYPE: Peru. Cajamarca: Yamaluc, R. Ferreyra 846 (holotype, USM!, USM photo at LP!). Additional specimen examined: Peru. Ancash, Lopez et al. 7473 (LP).

22. Onoseris minima Domke, Notizbl. Bot. Gart. Berlin-Dahlem 13: 247. 1936. TYPE: Peru. Dept. Arequipa: Socosani, D. B. Stafford 295a (holotype, K not seen). Additional specimen examined: Peru. Moquegua, A. Weberbauer 7440 (LP).

23. Onoseris odorata (D. Don) Hook. & Am., Comp. Bot. Mag. l: 103. 1835. Basionym: Chaetachlaena odorata D. Don. TYPE: Peru. Guayaquilia Peruvianorum, s.d., J. Tafalla s.n. (holotype, P not seen). Additional specimen examined: Peru. A. Weberbauer 1492 (US).

24. Onoseris onoseroides (Kunth) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Isotypus onoseroides Kunth. TYPE: Venezuela. Near Tui River, F. Humboldt & A. Bonpland s.n. (holotype, P not seen, P photo at LP!). Additional specimen examined: Colombia. Guajira, O. Haught 3937 (LP).

25. Onoseris peruviana Ferreyra, J. Arnold Arbor. 25: 373. 1944. TYPE: Peru. Junin: La Merced, J. F. Macbride 5425 (holotype, US!; isotype, F not seen, F photo at LP!).

26. Onoseris purpurea (L. f.) S. F. Blake, Proc. Biol. Soc. Wash. 38: 85. 1925. Basionym: Atractylis purpurea L. f. TYPE: Colombia. "Nova Granada," J. C. Mutis s.n. (syntype, US!). Colombia. Cundinamarca, Killip et al. 38365 (US).

27. Onoseris sagittata (Rusby) Rusby, Descr. S. Amer. pl.: 164. 1920. Basionym: Seris sagittatus Rusby. TYPE: Bolivia. Dept. Cochabamba: Toredon, M. Bang 1139 (isotypes, MO!, G not seen). Additional specimen examined: Bolivia. Samaipata, Fosberg 28652 (US).

28. Onoseris salicifolia Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 7. 1818. TYPE: Ecuador. Chimborazo: Alausi, F. Humboldt & A. Bonpland 3226 (holotype, P not seen; isotype, B not seen, B photo F n. 15896 at LP!). Additional specimen examined: Ecuador. Chimborazo, O. Haught 3318 (LP).

29. Onoseris silvatica Greenm., Proc. Amer. Acad. Arts 40: 51. 1904. TYPE: Costa Rica. Alto del Rodeo, Pittier 1622 (holotype, GH not seen). Additional specimen examined: Colombia. Cundinamarca, King 6027 (US).

30. Onoseris speciosa Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 6. 1818. TYPE: Ecuador. Chimborazo, Alausi, A. Bonpland 3235 (isotype, F not seen). Additional specimen examined: Peru. Amazonas, King & Bishop 9197 (US).

31. Onoseris weberbaueri Ferreyra, J. Arnold Arbor. 25: 386. 1944. TYPE: Peru. Cajamarca: hills between Tabaconas and Maranon rivers, A. Weberbauer 6163 (holotype, F not seen, F photo n. 49246 at LP!). Additional specimen examined: Peru. Amazonas, Gentry et al. 61362 (MO).

Literature. Ferreyra (1944, 1949, 1959, 1995), Cabrera (1975), Sagastegui Alva and Dillon (1985), Sancho (2004).

Paehylaena D. Don ex Hook. & Arn.

Comp. Bot. Mag. 1: 106. 1835. TYPE: Pachylaena atriplicfolia D. Don ex Hook. & Am. 38 of Figs. 35-38.

Chionoptera DC., Prodr. 7: 14. 1838. TYPE: Chionoptera gayophyta DC. [= Pachylaena atriplicfolia D. Don ex Hook. & Am.].

Etymology. From the Greek pachys, thick, stout, and chlaena, overcoat, blanket, because of the rigid phyllaries surrounding the florets, or with reference to the crass and thick leaves.

Herbs postrate, caulescent to semi-acaulescent, perennial. Leaves" nearly rosulate; pseudopetiolate; blades orbicular to spathulate, crass, rugose, pinnately veined, margin denticulate, glabrous. Capitulescences monocephalous, terminal; capitula pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bilobed, dorsally papillose, papillae occasionally extending below the bifurcation point. Cypselae truncate at the apex, glabrous; pappus of plumose, capillary bristles.

Pollen Description. Pachylaena atriplicifolia (104 of Figs. 101-106) (Telleria & Katinas, 2004). Pollen spheroidal-subprolate, medium to large size, tricolporate, exine Mutisia type, microechinate. Note: pollen similar to that of Urmenetea atacamensis Phil. (see also Parra & Marticorena, 1972).

Habitat and Distribution. Genus of one species distributed in the Andes of Argentina and Chile.

Species list

Pachylaena atriplicifolia D. Don ex Hook. & Am., Comp. Bot. Mag. 1: 106. 1835. TYPE: Argentina. Prov. Mendoza: near Agua del Ceno [Cerro] Pelado, on the ascent to E1 alto de los Manantides [Manantiales], s.d., J. Gillies s.n. (holotype, K not seen, K photo at LP!). Additional specimens examined: Argentina. Prov. Neuquen: Dept. Pehuenches, Cerro Chacayco, A. Chicchi 67 (LP); id., Cerro Piedra Santa, 7 Feb 1944, J. Frenguelli s.n. (LP*).

Literature. Johnston (1929), Cabrera (1971 b), Katinas (1996a, 2008).

Perdicium L.

Pl. rar. afr.: 22. 1760. TYPE: Perdicium capense L. 39 of Figs. 39-42. Idicium Neck., Elem. hot. 1: 28. 1790, nom. illeg.

Pardisium Burro. f., Fl. indica: 26. 1768, an orthographic variant of the name Perdicium.

Etymology. From the Greek perdikion, the name of a plant applied by Theophrastus to a species of Polygonum, perdikion is the diminutive of perdix, partridge.

Herbs perennial, acaulescent, scapose. Leaves rosulate; pseudopetiolate; blades oblong to obovate, pinnately veined, margin sinuate to runcinate, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, disciform; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with staminodes, corolla bilabiate; central florets bisexual, corolla bilabiate to tubular-bilabiate; anther apical appendages acute, tails smooth; style bilobed, branches dorsally papillose. Cypselae constricted to shortly rostrate at the apex, glabrous to pubescent; pappus of scabrid, basally connate, capillary bristles.

[FIGURES 39-42 OMITTED]

Pollen Description. Perdicium capense. Pollen spheroidal to prolate, spheroidal or elliptic in equatorial view, subcircular in polar view, large size, P x E = (50-65 x 41-47) [micro]m. Tricolporate, long colpi with microgranulate membrane. Exine Mutisia type, microechinate, 6-8 [micro]m thick at the equator and 4-5 [micro]m thick at the poles, ectosexine slightly columellate. Ratio ectosexine/endosexine: 1:1; 1:2; 1:3. Nexine ca. 2-3 [micro]m thick.

Habitat and Distribution. Genus of two species endemic to the western Cape in southern Africa.

Species list

1. Perdicium capense L., P1. rar. afr.: 22. 1760. TYPE: Sine loc., s.d., Burmann s.n. (holotype?, G not seen). Additional specimen examined: Africa. South Africa: terra Capensis, regio occidentalis, Tarkinsberg, Schlechter 8616 (MO).

2. Perdicium leiocarpum DC., Prodr. 7: 39. 1838. TYPE: Africa. Africa capensi, Camisberge, s.d., Drege s.n. (holotype, G-DC not seen). Additional specimen examined: Africa. s.d., Drege s.n. (MO).

Literature. Hansen (1985c).

Pertya Sch. Bip.

Bonplandia 10: 109. 1862. TYPE: Pertya glabrescens Sch. Bip. [= Pertya seandens Sch. Bip.]. 40 of Figs. 39-42.

Etymology. In honor of Joseph Anton Maximilian Perty (1804-1884), German botanist and zoologist.

Shrubs or subshrubs, stems branched. Leaves alternate or clustered on brachyblasts; sessile to petiolate; blades commonly ovate, oblong, elliptic, lanceolate to linear, palmately, pinnately to three-veined, margin entire to lobed, subglabrous to pubescent beneath. Capitulescences monocephalous, capitula terminal on the brachyblasts to paniculate; capitula pedunculate, homogamous, ligulate, one-, three- to 16-flowered; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally papillose. Cypselae, glabrous to pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Pertya (105 of Figs. 101-106). Pollen spheroidal to prolate, elliptic or subrectangular in equatorial view, circular in polar view, medium or large size. Tricolporate, colpi long with scabrate-microgranulate membrane. Exine Mutisia type, microechinate, equally thickened in the whole grain or slightly thickened at the poles. Nexine ca. 1.5 [micro]m thick. Pertya cordifolia Mattf. Pollen spheroidal-subprolate, elliptic in equatorial view, medium size, P x E = (41-42 x 38-40) [micro]m. Exine 5.5-7 [micro]m thick. Ectosexine with compact aspect. Ratio ectosexine/ endosexine: 1:1.5. Pertya discolor Rehder. Pollen subprolate-prolate, elliptic in equatorial view, medium size, P x E = (3743 x 29-33). Exine 5-6 [micro]m thick. Ectosexine with granulate aspect or slightly columellate, thick internal tectum. Ratio ectosexine/endosexine: ca. 1.5:1. Pertya ovata Maxim. Pollen spheroidal-subprolate, subrectangular, medium to large size, P x E = (48-52 x 3342) [micro]m. Exine 6-7 [micro]m thick, slightly thickened at the poles. Ratio ectosexine/endosexine: ca. 1 : 1.5 (see also Lin et al., 2005).

Habitat and Distribution. Genus of ca. 22 species distributed in Afghanistan, China, Japan, Taiwan, and Thailand.

Species list

1. Pertya aitchisoni C. B. Clarke, J. Linn. Soc., Bot. 18: 72. 1881. TYPE: Afghanistan. Kurrum Valley, Aitchison 392, 721 (syntypes, P!).

2. Pertya angustifolia Y. C. Tseng, Guihaia 5: 328. 1985. TYPE: China. Sichuan: Daofu, Songlinkou, Guan 50236 (type, SCBI not seen).

3. Pertya berberidoides (Hand.-Mazz.) Y. C. Tseng, Guihaia 5: 328. 1985. Basionym: Pertya bodinieri Vaniot var. berberidoides Hand.-Mazz. TYPE: China. Yunnan: Tonschieferfelsen de str. St. unter Meti uber dem Yangtse sw von Dschungdien, s. leg. n. 7781 (type, WU not seen).

4. Pertya bodinieri Vaniot, Bull. Acad. Int. Geogr. Bot. 12:116. 1903. TYPE: not localized. Additional specimen examined: China. Yunnan: Yun nan sen, Ducloux 257 (UC).

5. Pertya cordifolia Mattf., Notizbl. Bot. Gait. Berlin-Dahlem 11: 103. 1931. TYPE: China. Hunan: Yunshan, S. S. Sin n. S. H. 956 (type, probably B). Additional specimen examined: Kwansi, Steward & Cheo 972 (NY*).

6. Pertya corymbosa Y. C. Tseng, Guihaia 5: 332. 1985. TYPE: China. Guangxi: Rongshui, S. H. Chun 16893 (type, SCBI not seen).

7. Pertya desmocephala Diels, Notizbl. Bot. Gart. Berlin-Dahlem 9: 1032. 1926. TYPE: China. Chekiang: Lung chiung hsien, Hu 409 (type, probably M).

8. Pertya discolor Rehder, J. Arnold Arbor. 10: 135. 1929. TYPE: China. Gansu: en route to Linhua Shan, from Choni via Daozhou, J. F. Rock 12667 (type, S!). Additional specimen examined: Shansi, Chieh-hsiu distr., Mien-shan-ye, Smith 5786 (MO*).

9. Pertya henanensis Y. C. Tseng, Guihaia 5: 330. 1985. TYPE: China. Henan: Songxian County, Forest. Henan Exped. 973 (types, PE not seen, IBSC not seen).

10. Pertya hossei Craib ex Hosseus, Kew Bull.: 402. 1911. TYPE: Thailand. Doi Djieng Dao-Hochland, C. C. Hosseus 405 (type, MO!).

11. Pertya macrophylla Nakai, Icon. pl. koisikav. 4: 77. 1920. TYPE: Japan. Hondo: Sagami, prope Zushi, 19 Oct 1919, Nakai s.n. (type, TI not seen, TI photo at P!).

12. Pertya mattfeldii Bornm., Repert. Spec. Nov. Regni Veg. Beih. 108: 65. 1938. TYPE: not localized. Additional specimen examined: Afghanistan. Nuristan: between Pushuki and Kushimaicot, 1955, Kitamura s.n. (KYO).

13. Pertya monocephala W. W. Sm., Notes Roy. Bot. Gard. Edinburgh 8: 212. 1914. TYPE: China. Yunnan: Jalong, Oui-chu valley, F. Kingdon Ward 1141 (type, E not seen, E photo at LP!).

14. Pertya ovata Maxim., Bull. Acad. Imp. Sci. Saint-Petersbourg. 16: 217. 1871. TYPE: Japan. Yokohama, 1862, Maximowicz s.n. (type, P!). Additional specimen examined: Honshu, Pref. Wakayama, near Mizugamine, Takahashi et al. 1387 (MO*).

15. Pertya phylicoides Jeffrey, Notes Roy. Bot. Gard. Edinburgh 5: 200. 1912. TYPE: China. Yunnan: from Zhongdian plateau to Yangzi, near Jiao-tou, G. Forrest 112 (type, E not seen).

16. Pertyapungens Y. C. Tseng, Guihaia 5: 334. 1985. TYPE: China. Guangdong: Yangchun, Jinshan, Wang 38778 (type, MO!).

17. Pertya scandens (Thunb. ex Murray) Sch. Bip., Bonplandia 10: 109. 1862. Basionym: Erigeron scandens Thunb. ex Murray. TYPE: Sine loc., Faurie 6756 (type of Pertya scandens (Yhunb. ex Murray) Sch. Bip. f. schultziana Franch., P!).

18. Pertya simozawai Masam., Trans. Nat. Hist. Soc. Taiwan 30: 37. 1940. TYPE: not localized. Additional specimen examined: Taipei. Sanshia: Wuliaochien, Peng 8683 (TAI).

19. Pertya sinensis Oliv., Icon. pl. 23: pl. 2214. 1892. TYPE: not localized. Additional specimens examined: China. Hubei: Shennongjia Forerst, S of Jiuhuping Forest Farm along the Jizigou canyon bottom, Sino-Amer. Exped. 1537 (MO, NY).

20. Pertya tsoongiana Ling, Contr. Bot. Surv. N. W. China 1: 40. 1939. TYPE: not localized. Additional specimen examined: China. Szechuan: Fushan, T. P. Wang 10293 (PE).

21. Pertya uniflora (Maxim.) Mattf., Notizbl. Bot. Gart. Berlin-Dahlem 11: 105. 1931. Basionym: Myripnois uniflora Maxim. TYPE: China. Kansu, Przewalski 1880 (type, K not seen).

22. Pertya yakushimensis H. Koyama & Nagam., Acta Phytotax. Geobot. 39: 67. 1988. TYPE: Japan. Kyushu: Kagoshima-ken, Yakushima, Tainoko River, 1984, Miura s.n. (type, KYO!).

Literature. Kitamura (1937), Koyama (1973, 1975), Tseng (1985), Hind (2001a).

Plazia Ruiz & Pay.

Fl. peruv, prodr.: 104. 1794. TYPE: Plazia conferta Ruiz & Pav. 41 of Figs. 39-42.

Aglaodendron J. Remy, Ann. Sci. Nat., Bot. 12: 175. 1849. TYPE: Aglaodendron cheiranthifolium J. Remy [= Plazia cheiranthifolia (J. Remy) Wedd.].

Harthamnus H. Rob., Phytologia 45: 452. 1980. TYPE: Harthamnus boliviensis H. Rob. [= Plazia daphnoides Wedd.].

Etymology. In honor of Juan Plaza (1526-1609), a Spanish botanist and medical doctor.

Shrubs. Leaves alternate, clustered at the end of the branches; sessile; blades obovateoblong, one-veined, margin entire, glabrous to densely pubescent on both surfaces. Capitulescences monocephalous, terminal; capitula sessile, surrounded by leaves, homogamous, radiate; receptacle epaleate; involucre three- to four-seriate. Florets dimorphic, bisexual; marginal florets with corolla bilabiate, often geniculate, outer lip expanded; central florets with corolla tubular sub-campanulate, deeply five-lobed; anther apical appendages reddish, truncate, tails papillose; style bifid, branches dorsally papillose, rounded to truncate at the apex. Cypselae glabrous; pappus of scabrid bristles, some capillary, the innermost flattened and longer.

Pollen Description. Plazia daphnoides (106 of Figs. 101-106) (Telleria & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate (see also Parra & Marticorena, 1972).

Habitat and Distribution. Genus of three species from the Andes of Argentina, Bolivia, Chile, and Peru.

Species list

1. Plazia cheiranthifolia (J. Remy) Wedd., Chlor. andina 1: 12. 1855. Basionym: Aglaodendron cheiranthifolium J. Remy. TYPE: Chile. s.d., C. Gay s.n. (holotype, P not seen, P photo F n. 38107 at LP!).

2. Plazia conferta Ruiz & Pav., Syst. veg. ft. peruv, chil. 1: 187. 1798. TYPE: Peru. Tarma, s.d., H. Ruiz & J. A. Pavon s.n. (holotype, MA not seen).

3. Plazia daphnoides Wedd., Chlor. andina 1: 13. 1855. TYPE: Peru. Cordillera de Tacora, entre Tacna et La Paz, s.d., H. A. Weddell s.n. (lectotype, selected by Ferreyra, 1995: 83, P not seen). Additional specimens examined: Argentina. Mendoza: Paramillo de Uspallata, A. Ruiz Leal 4839 (LP); Prov. Jujuy: Casabindo, A. L. Cabrera 9315 (LP*).

Literature. Cabrera (1951, 1978), Robinson (1980), Ferreyra (1995), Freire (1996).

Salcedoa F. Jimenez Rodr. & Katinas

Syst. Bot. 29: 991. 2004. TYPE: Salcedoa mirabaliarum F. Jimenez Rodr. & Katinas. 42 of Figs. 39-42.

Etymology. For the Salcedo province, the type locality, in the Dominican Republic.

Trees small. Leaves alternate; petiolate; blades oblanceolate to elliptical, pinnately veined, margin entire, glabrous above, tomentose beneath. Capitulescences corymbose; capitula pedunculate, homogamous, bilabiate; receptacle epaleate; involucre multiseriate, outermost phyllaries decurrent onto the stalk. Florets isomorphic, bisexual; corolla bilabiate, deeply incised; anther apical appendages apiculate, tails papillose; style bilobed, branches reddish, dorsally smooth. Cypselae villose; pappus of scabrid bristles, the outermost capillary, the innermost somewhat paleaceous, with tips slightly broadened, and longer.

Pollen Description (Jimenez Rodriguez et al., 2004). Pollen prolate, large size, tricolporate, exine Wunderlichia type, echinate.

Habitat and Distribution. Monotypic genus endemic to Cordillera Septentrional, northern Dominican Republic.

Species list

1. Salcedoa mirabaliarum F. Jimenez Rodr. & Katinas, Syst. Bot. 29: 991. 2004. TYPE: Dominican Republic. Prov. Salcedo: Cordillera Septentrional, el Penon del Nuevo Mundo, F. Jimenez et al. 3345 (isotype, LP!). Additional specimen examined: Dominican Republic. Prov. Salcedo: Mun. Tenares, Distr. Blanco Arrriba, Veloz et al. 2383 (LP*).

Literature. Jimenez Rodriguez et al. (2004).

Trichocline Cass.

Bull. Sci. Soc. Philom. Paris: 13. 1817. TYPE: Doronicum incanum Lam., Encycl. 2: 315. 1786, non L., 1753, nec Hill, 1759, nora. illeg. [= Trichocline incana Cass.]. 43 of Figs. 43-46.

Amblysperma Benth., Enum. pl.: 67. 1837. [= Trichocline sect. Amblysperma (Benth.) Zardini, Darwiniana 19: 657. 1975]. TYPE: Amblysperma scapigera Benth. [= Trichocline spathulata (A. Cunn. ex DC.) J. H. Willis].

Bichenia D. Don, Trans. Linn. Soc. London 16: 236. 1830. TYPE: Bichenia aurea D. Don [= Trichocline aurea (D. Don) Reiche].

Etymology. From the Greek thrix, trichos, hair, and kline, bed in reference to the fimbriate receptacle.

Herbs perennial, acaulescent, usually scapose. Leaves rosulate; sessile to petiolate; blades linear to ovate, pinnately veined, margin entire to pinnatisect, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate; receptacle epaleate, smooth to fimbriate; involucre multiseriate. Florets dimorphic; marginal florets female, with well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails smooth to papillose, filaments smooth to papillose; style bilobed, dorsally papillose to sometimes glabrescent. Cypselae truncate at the apex, shortly pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Trichocline reptans (Wedd.) B. L. Rob. (107 of Figs. 107-111) (Telleria & Katinas, 2004). Pollen prolate, large size, tricolporate, exine Mutisia type, microechinate. Note: pollen similar to that of Lulia nervosa.

Habitat and Distribution. Genus of 22 species most of them in South America from southern Peru to central Argentina and Chile, and one species in Australia (see Observation).

[FIGURES 43-46 OMITTED]

Species list

1. Trichocline aurea (D. Don) Reiche, Anales Univ. Chile 115: 342. 1904. Basionym: Bichenia aurea D. Don. TYPE: Chile. Coquimbo, s.d., Caldcleugh s.n. (type, probably GH). Additional specimen examined: Chile. Prov. Concepcion: La Florida, E. Barros 244 (LP).

2. Trichocline auriculata (Wedd.) Hieron., Bot. Jahrb. Syst. 21: 369. 1896. Basionym: Bichenia auriculata Wedd. TYPE: Bolivia. Environs de Potosi, D'Orbigny 1429 (holotype, P not seen, P photo F n. 38073 at SI not seen). Additional specimen examined: Argentina. Prov. Salta: Dept. Iruya, quebrada de Iruya, Luti & Montana 5358 (LP).

3. Trichocline beckeri H. Rob., Phytologia 65: 47. 1988. TYPE: Peru. Dept. Cajamarca: surroundings of Cajamarca, B. Becker & F. M. Terrones H. 1305 (holotype, US not seen, US photo at LP!).

4. Trichocline boecheri Cabrera, Bol. Soc. Argent. Bot. 15: 122. 1973. TYPE: Argentina. Prov. Mendoza: ruta 40, km 512, T. W. Bocher et al. 1551 (holotype, LP!). Additional specimen examined: Argentina. Prov. San Juan: Dept. Iglesia, Maliman, A. L. Cabrera et al. 24526 (LP).

5. Trichocline catharinensis Cabrera, F1. Ilustr. Catarinense 1: 44. 1973. TYPE: Brazil. State Santa Catarina: Cacador, Fazenda Carneiros, L. B. Smith & R. Reitz 9041 (holotype, LP!). Additional specimen examined: Brazil. State Santa Catarina: Mun. Curitibanos, entre Curitibanos e Lages, Pereira 8398 & Pabst 7673 (LP).

6. Trichocline caulescens Phil., Anales Mus. Nac. Santiago de Chile 8: 1. 1891. TYPE: Chile. Medano, pr. Pica, F. Philippi 2069 (holotype, SGO not seen; isotype, B not seen, B photo F n. 16013 at LP!). Additional specimen examined: Chile. Prov. Tarapaca: camino al Portezuelo de Chapiquina, M. Ricardi et al. 35 (LP).

7. Trichocline cineraria (D. Don) Hook. & Am., Comp. Bot. Mag. 1: 103. 1835. Basionym: Bichenia cineraria D. Don. TYPE: Argentina. Prov. Mendoza: near el Portezuelo de Uspallata, Andes of Mendoza, s.d., J. Gillies s.n. (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Carlos, camino a Laguna del Diamante, Barkley & Cuezzo 20Mz398 (LP).

8. Trichocline dealbata (Hook. & Am.) Benth. & Hook. f. ex Griseb., Abh. Konigl. Ges. Wiss. Gottingen 24: 215. 1879. Basionym: Chaetanthera dealbata Hook. & Arn. TYPE: Argentina. Prov. Mendoza: ascens to Peuquenes, Andes of Mendoza, s.d., J. Gillies s.n. (lectotype, designated by Zardini 1975: 704, K not seen). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Tupungato, Portezuelo Morado, Semper 11842 (LP).

9. Trichocline deserticola Zardini, Darwiniana 20: 304. 1976, nora. novum. Basionym: Onoseris spathulata Phil. TYPE: Chile. Prov. Antofagasta: Ascotan, 23 Feb 1885, F. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Chile. Prov. Antofagasta: camino a Ascotan, NW volcan San Pedro, Martin 449 (LP).

10. Trichocline exscapa Griseb., Abh. Konigl. Ges. Wiss. Gottingen 19: 197. 1874. TYPE: Argentina. Prov. Tucuman: sierra de Tucuman, P G. Lorentz 132 (isotype, LP!). Additional specimen examined: Argentina. Prov. Salta: Dept. Metan, cima del cerro de Las Mesadas Largas (continuacion del Creston), S. Maldonado 2962 (LP).

11. Trichocline heterophylla (Spreng.) Less., Linnaea 5: 289. 1830. Basionym: Onoseris heterophylla Spreng. TYPE: Brazil. Brasilia meridionalis, Sellow 907 (isotype, LP!). Additional specimen examined: Uruguay. Dept. Florida: Cerro Colorado, estancia San Pedro, Gallinal et al. 2810 (LP).

12. Trichocline humilis Less., Linnaea 5: 288. 1830. TYPE: Brazil. Brasilia, s.d., Sellow s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Entre Rios: Dept. Federacion, Chajari, A. L. Cabrera 12361 (LP).

13. Trichocline incana Cass., Dict. sci. nat. 55: 216. 1826, nom. novum. Basionym: Doronicum incanum Lam., nom. illeg. TYPE: Uruguay. Montevideo, s.d., D. Commerson s.n. (holotype, P not seen, P photo F n. 22642 at SI not seen). Additional specimen examined: Uruguay. Dept. Maldonado: Punta del Este, Scott de Biraben 24 (LP).

14. Trichocline linearifolia Malme, Kongl. Svenska Vetensk. Acad. Handl. 12: 114. 1933. TYPE: Brazil. State Parana: Tamandua, P. Dusen 7714 (isotype, LP!). Additional specimen examined: Brazil. State Parana: Mun. Curitiba, Rio Atuba, Kummrow 198 (LP).

15. Trichocline macrocephala Less., Linnaea 5: 288. 1830. TYPE: Brazil. Brasilia, s. d., Sellow s.n. (neotype, designated by Zardini, 1975: 666, K not seen). Additional specimen examined: Brazil. State Santa Catarina: Mun. Joacaba, campos of Palmas, 62 km W of Cacador, S. B. Smith & R. M. Klein 11409 (LP).

16. Trichocline macrorhiza Cabrera, Darwiniana 9: 51. 1949. TYPE: Argentina. Prov. Jujuy: Tumbaya, Volcan, Cerro Bola Loma, 21 July 1948, s. leg. (holotype, BAB not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, subida de Purmamarca, Abra de Pives, A. L. Cabrera et al. 26403 (LP).

17. Trichocline maxima Less., Linnaea 5: 290. 1830. TYPE: Brazil. Brasilia, s.d., Sellow s.n. (isotype, K not seen, K photo at LP!). Additional specimen examined: Uruguay. Dept. Rivera: Paso Ataques, J. Chebataroff 9112 (LP).

18. Trichocline plicata Hook. & Am., Comp. Bot. Mag. 1: 103. 1835. TYPE: Argentina. Prov. San Luis: Portezuela and Cerro de Achiras, s.d., s. leg. (holotype, OXF not seen, OXF photo at LP!). Additional specimen examined: Argentina. Prov. Cordoba: Dept. Pocho, Sierra de Pocho, A. L. Cabrera et al. 29649 (LP).

19. Trichocline reptans (Wedd.) Hieron., Bot. Jahrb. Syst. 21: 369. 1896. Basionym: Bichenia reptans Wedd. TYPE: Bolivia. Environs de Chuquisaca, s.d., s. leg. (type, probably P). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Santa Barbara, Abra de los Morteros, A. L. Cabrera et al. 24099 (LP*).

20. Trichocline sinuata (D. Don) Cabrera, Man. Fl. Alrededores Buenos Aires: 531. 1953. Basionym: Bichenia sinuata D. Don. TYPE: Argentina. Las Pampas del rio Desaguadero, s.d., J. Gillies s.n. (lectotype, designated by Zardini, 1975: 673, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido de Patagones, Igarzabal, A. L. Cabrera & H. A. Fabris 14847 (LP*).

21. Trichocline spathulata (A. Cunn. ex DC.) J. H. Willis, W. Austral. Naturalist 10: 157. 1967. Basionym: Celmisia spathulata A. Cunn. ex DC. TYPE: Australia. SW coast Australia, Oyster Harbour, King George's Sound, Jan 1818, A. Cunningham s.n. (holotype, G-DC not seen, G-DC photo at LP!). Additional specimen examined: Australia. Murray District, Pritzel 140 (LP).

22. Trichocline speciosa Less., Syn. gen. Compos.: 117. 1832. TYPE: Brazil. Brasilia, s.d., Sellow s.n. (neotype, designated by Zardini, 1975: 696, P not seen). Additional specimen examined: Brazil. State Parana: Mun. R Grossa, Parque V. Velha, G. Hatschbach 16048 (LP).

Observation. A new species of Amblysperma, A. minor Keighery, was described for western Australia (Keighery, 2005). The genus Amblysperma was reduced to the synonymy of Trichocline (Katinas, 2004b), and therefore this new species could belong to Trichocline. However, because the protologue of this new species lacks a description and a detailed illustration of corollas and style, we prefer to maintain it as a dubious species until we analyze this taxon in more detail.

Literature. Zardini (1975), Robinson (1988), Hind (2001 b), Katinas (2004b).

Uechtritzia Freyn

Oesterr. Bot. Z. 42: 240. 1892. TYPE: Uechtritzia armena Freyn & Sint. 44 of Figs. 43-46.

Etymology. In honor of the German (Silesian) botanist Rudolf Karl von Uechtritz (1838-1886).

Herbs perennial, acaulescent, scapose. Leaves rosulate; pseudopetiolate; blades elliptic to ovate, pinnately veined, margin entire to lyrate or pinnatifid, glabrous to pubescent. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate; receptacle fimbriate-laciniate, involucre multiseriate. Florets dimorphic; marginal florets female, with well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bilobed, branches dorsally papillose. Cypselae truncate at the apex, villose with shaggy hairs; pappus of scabrid, capillary bristles.

Pollen Description. Uechtritzia kokanica (Regel & Schmalh.) Pobed. (Telleria & Katinas, 2004). Pollen prolate, large size, tricolporate, exine Mutisia type, scabrate.

Habitat and Distribution. Genus of three species distributed in southern Russia, north to Afghanistan, Kashmir, northwestern China, Armenia, and northwestern India.

Species list

1. Uechtritzia armena Freyn & Sint., Oesterr. Bot. Z. 42: 241. 1892. TYPE: Armenia. Sipikordagh, Sinentis 3185 (holotype, BRNU not seen).

2. Uechtritzia kokanica (Regel & Schmalh.) Pobed., Fl. U.S.S.R. 28: 597. 1963. Basionym: Gerbera kokanica Regel & Schmalh. TYPE: Bangladesh. In Kokania, prope Schahimardan, Karassu, s.d., O. Fedtschenko s.n. (holotype?, LE not seen). Additional specimen examined: Russia. Tadzhikistania: Hissaricum, Unou, 13 Aug 1934, F. Zaprjagaev s.n. (WIS).

3. Uechtritzia lacei (Watt) C. Jeffrey, Kew Bull. 21: 213. 1967. Basionym: Gerbera lacei Watt. TYPE: India. Nachar forest, Lace 935 (lectotype, designated by Hansen, 1988: 74, K not seen).

Literature. Hansen (1988).

Urmenetea Phil.

Fl. atacam.: 26. 1860. TYPE: Urmenetea atacamensis Phil. 45 of Figs. 43-46.

Etymology. In honor of Hieronymus Urmeneta (1816-1881) who promoted agricultural science in Chile.

Herbs perennial, subscapose. Leaves nearly rosulate; petiolate; blades spathulate to orbicular, palmately veined, margin serrate, rugose above, woolly beneath. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with reduced, filiform staminodes, corollas bilabiate, outer lip expanded; central florets with corollas tubular-bilabiate, shallowly five-lobed, lobes straight, one incision deeper than the others; anther apical appendages acute to apiculate, tails papillose, filaments papillose at the base; style bifid, branches dorsally papillose. Cypselae glabrous; pappus of scabrid, capillary bristles, a few inner ones flattened and wider.

Pollen Description (Telleria & Katinas, 2004) (108 of Figs. 107-111). Pollen spheroidal, large size, tricolporate, exine Mutisia type, microechinate. Note: pollen similar to that of Pachylaena atriplicifolia (see also Parra & Marticorena, 1972).

Habitat and Distribution. Monotypic genus from the Andes of Chile and northwestern Argentina.

Species list

1 Urmenetea atacamensis Phil., F1. atacam.: 26. 1860. TYPE: Chile. Prov. Atacama: cerca de la poblacion de San Pedro de Atacama, s.d., R. Philippi s.n. (type, probably SGO). Additional specimen examined: Argentina. Prov. Salta: Los Andes, Abra de Navarro, A. L. Cabrera & 14. Schwabe 80 (LP*).

Literature. Cabrera (1978), Freire (1996), Sancho (2004).

Tribe Nassauvieae Cass.

H. Cassini, J. Phys. Chim. Hist. Nat. Arts. 88: 198. 1819. Nassauvioideae (Cass.) Lindl. in Loudon, Encycl. pl.: 1074. 1829. Nassauviinae Dumort., Anal. fam. pl.: 31. 1829, as "Nassauvieae"; Less., Linnaea 5: 2. 1830. Nassauviaceae Burmeist., Handb. Naturgesch. I: 290. 1837. TYPE: Nassauvia Comm. ex Juss.

Trixideae (Cass.) Lindl. in Loudon, Encycl. pl.: 1074. 1829. Trixidinae Less., Linnaea 5: 6. 1830, as "Trixidea." TYPE: Trixis P. Browne.

Jungeae D. Don, Trans. Linn. Soc. London 16: 224. 1830, as "Jungieae." TYPE: Jungia L. f.

Polyachyreae D. Don, Trans. Linn. Soc. London 16: 229. 1830. Polyachyrinae Endl., Gen. pl. 3: 489. 1838. TYPE: Polyachyrus Lag.

Herbs, subshmbs, shrubs or vines. Leaves alternate to rosulate, entire to pinnatisect, rarely spiny, glabrous to pubescent. Capitulescences monocephalous, laxly to densely corymbose, racemose, paniculate to glomemlose or pseudocephalia. Capitula homogamous, discoid or radiate by extension of the outer lip in marginal corollas, rarely disciform; involucre uniseriate to multiseriate; receptacle epaleate to paleate, glabrous to pubescent. Florets isomorphic, rarely dimorphic, bisexual, corolla bilabiate with the outer lip three-toothed and the inner bifid, rarely tubular; anthers with apical appendage lanceolate, acute, long, smooth to papillose tails; style cleft into two truncate branches, each branch with an apical crown of collector or sweeping hairs, rarely branches dorsally papillose and papillae not in a crown. Cypselae truncate to rostrate at the apex, glabrous to pubescent; pappus of capillary bristles to paleaceous, scabrose to plumose, equal in length, rarely pappus absent. Pollen microechinate, exine distinctly bilayered, with the ectosexine and the endosexine clearly columellate. n=4, 11, 12, 20, 22, 27, 28; 2n=16, 24, 36, 47 [+ or -] 3, 54.

Characters that define the tribe are in italics.

Tribe with 25 genera and ca. 320 species of the Neotropical biogeographic province (Cabrera & Willink, 1973). Plants belonging to this tribe are an important component of the Andean and Patagonian flora.

Genera: Acourtia D. Don, Ameghinoa Speg., Berylsimpsonia B. L. Turner, Burkartia Crisci, Calopappus Meyen, Cephalopappus Nees & Mart., Criscia Katinas, Dolichlasium Lag., Holocheilus Cass., Jungia L. f., Leucheria Lag., Leunisia Phil., Lophopappus Rusby, Macrachaenium Hook. f., Marticorenia Crisci, Moscharia Ruiz &Pav., Nassauvia Comm. ex Juss., Oxyphyllum Phil., Panphalea Lag., Perezia Lag., Pleocarphus D. Don, Polyachyrus Lag., Proustia Lag., Triptilion Ruiz &Pav., Trixis P. Browne.

Literature. Crisci (1974a, 1980), Cabrera (1977), Katinas (1995).
Key to Genera

1a. Paleae embracing all or only the central florets of the capitula.

2a. Leaves linear to linear-lanceolate Pleocarphus

2b. Leaves ovate-elliptic, suborbicular to triangular-ovate.

3a. Leaves ovate-elliptic, pinnately veined, sessile, decurrent to
clasping Marticorenia

3b. Leaves elliptic to orbicular-cordate, palmately veined, petiolate
(except Jungia stuebelii with leaves ovate, pinnately veined, sessile)
 Jungia
1b. Paleae only in the marginal florets of the capitula, or completely
absent.

4a. Pappus absent.

5a. Leaves spathulate, entire; cypselae constricted to rostrate at the
apex Cephalopappus

5b. Leaves linear-lanceolate, orbicular, entire to lyrate to
pinnatisect; cypselae truncate at the apex Panphalea

4b. Pappus present.

6a. Pappus bristles narrowly paleaceous, as long as the florets or
longer (except in Nassauvia coronipappa with crown-like pappus).

7a. Pappus elements longitudinally plicate, apically plumose
Triptilion

7b. Pappus elements flat, ciliate to laciniate throughout their
length.

8a. Involucre multiseriate; capitulescence monocephalous Calopappus

8b. Involucre two-seriate; capitulescences monocephalous, racemose,
spicate to pseudocephalium Nassauvia

6b. Pappus capillary, if paleaceous very short.

9a. Capitula arranged in pseudocephalia.

10a. Pseudocephalia spiciform Polyachyrus

10b. Pseudocephalia capituliform Moscharia

9b. Capitulescences monocephalous or cymose.

11a. Most leaves reduced to three-parted spines; paleae subtending all
the marginal florets Oxyphyllum

11b. Leaves not reduced to three-parted spines; capitula with or
without paleae.

12a. Style branches dorsally papillose.

13a. Shrubs or little trees.

14a. Plants with thorns to unarmed; capitula in
racemes to panicles Proustia

14b. Plants unarmed; capitula solitary or few at the
apex of the branches Lophopappus

13b. Perennial, acaulescent herbs Macrachaenium

12b. Style branches smooth, crowned by a tuft of collector
hairs.

15a. Shrubs or subshrubs.

16a. Leaves pinnatisect Dolichlasium

16b. Leaves entire to lobulate.

17a. Clambering plants, with recurved, bifurcate,
pseudostipulate spines Berylsimpsonia

17b. Erect plants, without spines or with thorns
formed by foreshortened stems.

18a. Pappus paleaceous-setaceous, unequally
fused at the base Leunisia

18b. Pappus bristles scabrid to plumose

19a. Dwarf shrubs with subulate leaves;
capitulescences monocephalous or
two-headed Burkartia

19b. Erect plants with broad leaves;
capitulescences monocephalous to
cymose with numerous capitula.

20a. Stems with brachyblasts and
macroblasts; cypselae usually
truncate at the apex Ameghinoa

20b. Stems without brachyblasts;
cypselae usually rostrate Trixis

15b. Perennial leafy caulescent or scapose herbs.

21a. Plants covered with a woolly to silky white
pubescence Leucheria

21b. Plants glabrous, subglabrous or with a reddish
pubescence.

22a. Corollas and pappus orange-red.

23a. Scapose herbs; capitulescences monocephalous Criscia

23b. Caulescent herbs; capitulescences corymbose,
paniculate to spicate Trixis

22b. Corollas pink, purple, white, rarely yellow;
pappus white, brown, reddish, bronzepurplish.

24a. Involucre one- to two-seriate Holocheilus

24b. Involucre with three to more series of
phyllaries.

25a. Stem base with a tuft of rust-colored
woolly hairs; capitula
discoid Acourtia

25b. Stem base glabrous or at least
not densely rust-colored woolly
hairs; capitula radiate, occasionally
capitula discoid Perezia


Acourtia D. Don

Trans. Linn. Soc. London 16: 203. 1830. TYPE: Acourtia formosa D. Don [= Acourtia fruticosa (La Llave & Lex.) B. L. Turner]. 46 of Figs. 43-46. Neoshinneria B. L. Turner, Phytologia 38: 456. 1978, nom. nudum.

Etymology. In honor of Mrs. Mary Elizabeth Catherine Gibbes A'Court (1792-1878), a British amateur botanist.

Herbs perennial or subshrubs, caulescent or acaulescent, base of the stem with a tuft of rust-colored woolly hairs. Leaves rosulate to spreading; sessile, petiolate, pseudopetiolate to clasping; blades lanceolate, broadly orbicular to cordate, pinnately veined, margin usually spinulose, entire to lyrate, glabrous to pubescent. Capitulescences monocephalous, or corymbose to paniculate; capitula pedunculate to subsessile, homogamous, discoid; receptacle epaleate, occasionally fimbrillate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate, occasionally tubular-funnel-form five-lobed; anther apical appendages acute to slightly apiculate, tails smooth; style bifid, branches linear, apically rounded, crowned by collector hairs. Cypselae truncate at the apex, glabrous to pubescent; pappus of simple to scabrid bristles, soft to rigid.

Pollen Description. Acourtia (Crisci, 1974a; Cabrera-Rodriguez, 1992 sub Rzedowskiela; Cabrera-Rodriguez & Dieringer, 2003). Pollen subprolate, oblate-spheroidal, prolate-spheroidal or spheroidal, medium size, tricolporate, exine Trixis and Proustia type, slightly microechinate.

Habitat and Distribution. Genus of ca. 85 species distributed from southern United States to Central America reaching El Salvador, with a high concentration in the forested middle and higher slopes of the Sierra Madre in Mexico and the high plateaus east of the mountains in central and northern Mexico.

Species list

1. Acourtia acevedoi M. Gonzalez, Phytologia 61: 117. 1986. TYPE: Mexico. Durango: Municipio Vicente Guerrero, Rincon de las Mulas, 3 km al SW de San Isidro, S. Acevedo 163 (isotype, NY, NY photo at LP!).

2. Acourtia aspera (Bacig.) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia aspera Bacig. TYPE: Mexico. Coahuila: Saltillo, Bro. Arsene 3395 (isotype, MO, MO photo at LP!).

3. Acourtia belizeana B. L. Turner, Phytologia 74:138.1993. TYPE: Belize. Toledo District: lower part of Richardson Creek, affluent of Bladen Branch, lower part of Maya Mtns., G. Davidse & A. E. Brant 31904 (isotype, US not seen).

4. Acourtia bravohollisiana Rzed., Bol. Soc. Bot. Mexico 45: 98. 1983. TYPE: Mexico. Guerrero: camino a la torte de microondas Tuxpan, [+ or -] 10 km al NE de Iguala, J. Rzedowski 37162 (isotype, NY, NY photo at LP!).

5. Acourtia butandae R. L. Cabrera, Brittonia 53: 423. 2001. TYPE: Mexico. Chihuahua: 18 km of Villa Matamoros, hwy 45, L. Cabrera & G. Dieringer 650 (isotype, US not seen, US photo at LP!).

6. Acourtia caltepecana B. L. Turner, Phytologia 74: 386. 1993. TYPE: Mexico. Puebla: Mpio. Caltepec, Cerro El Tambor, al NE de Caltepec, P. Tenorio 7634 (holotype, TEX not seen).

7. Acourtia carpholepis (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia carpholepis A. Gray. TYPE: Mexico. Puebla,: Chapulco, Liebmann 351 (lectotype, designated by Turner, 1993a: 387, GH, not seen).

8. Acourtia carranzae R. L. Cabrera, Brittonia 53: 416. 2001. TYPE: Mexico. Querrtaro: Mun. Arroyo Seco, 1 km al S de Arroyo Seco, E. Carranza 1488 (holotype, IEB not seen).

9. Acourtia ciprianoi Panero & Villasenor, Brittonia 51: 87. 1999. TYPE: Mexico. Oaxaca: Dto. Tehuantepec, Mun. Santiago Astata, carretera Pochutla-Salina Cruz, C. Martinez 1978 (isotype, TEX not seen).

10. Acourtia collina (S. Watson) Crisci, J. Arnold Arbor. 55: 605. 1974. Basionym: Perezia collina S. Watson. TYPE: Mexico. Jalisco: near Guadalajara, C. G. Pringle 2123 (holotype, GH not seen).

11. Acourtia cordata (Cerv.) B. L. Turner, Phytologia 74: 388. 1993. Basionym: Perdicium cordatum Cerv. TYPE: Mexico. DF?, montibus del desierto Mexico vicinis, M. Sesse & J. M. Mocino 3736 (lectotype, designated by Turner, 1993a: 388, M, not seen).

12. Acourtia coulteri (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia coulteri A. Gray. TYPE: Mexico. State Hidalgo: Zimapan, Coulter 234 (holotype, GH not seen).

13. Acourtia cuernavacana (B. L. Rob. & Greenm.) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia cuernavacana B. L. Rob. & Greenm. TYPE: Mexico. Morelos: above Cuernavaca, C. G. Pringle 6196 (isotype, MO, MO photo at LP!).

14. Acourtia dieringeri R. L. Cabrera, Sida 13: 419. 1989. TYPE: Mexico. Chihuahua: 20 km N of Basaseachic, along the Basaseachic-Yepachic dirt road, L. Cabrera & G. Dieringer 628 (isotype, GH not seen).

15. Acourtia discolor Rzed., Bol. Soc. Bot. Mexico 45: 100. 1983. TYPE: Mexico. Oaxaca: 30 km al NNW de Telixtlahauaca, sobre la carretera de Cuicatlan, J. Rzedowski 37181 (holotype, ENCB not seen).

16. Acourtia dissiticeps (Bacig.) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia dissiticeps Bacig. TYPE: Mexico. San Luis Potosi: Sierra Tablon, Minas de San Rafael, C. A. Purpus 5018 (isotype, MO, MO photo at LP!).

17. Acourtia dugesii (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia dugesii A. Gray. TYPE: Mexico. State Guanajuato: Guanajuato, s.d., A. Duges s.n. (holotype, GH not seen).

18. Acourtia durangensis B. L. Turner, Phytologia 74: 389. 1993. TYPE: Mexico. State Durango: Cienega bottomland near small lake, 40 mi N of Cd. Durango, H. Gentry 8594 (holotype, MICH not seen).

19. Acourtia elizabethiae Rzed. & Calderon, Acta Bot. Mex. 12: 16. 1990. TYPE: Mexico. Queretaro: alrededores de Maconi, municipio de Cadereyta, J. Rzedowski 47625 (holotype, ENCB not seen).

20. Acourtia erioloma (S. F. Blake) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia erioloma S. F. Blake. TYPE: Mexico. Oaxaca: near Tlaxiaco, W. H. Camp 2225 (holotype, NY not seen, NY photo at LP!).

21. Acourtia fragrans Rzed., Bol. Soc. Bot. Mexico 45: 102. 1983. TYPE: Mexico. Puebla: Salinas Grandes, cerca de Zapotitlan de las Salinas, J. Rzedowski 37174 (isotype, NY not seen, NY photo at LP!).

22. Acourtia fruticosa (La Llave & Lex.) B. L. Turner, Phytologia 74: 390. 1993. Basionym: Perezia fruticosa La Llave & Lex. TYPE: Mexico. Michoacan: Jesus del Monte near Morelia, C. G. Pringle 10410 (neotype, designated by Turner, 1993a: 390, LL not seen; isoneotype, US not seen, US photo at LP!).

23. Acourtia gentryi R. L. Cabrera, Acta Bot. Mex. 20: 35. 1992. TYPE: Mexico. Sinaloa: above la Jolla, Sierra Surotato, 14. S. Gentry 7258 (isotype, NY not seen, NY photo at LP!).

24. Acourtia glandulifera (D. L. Nash) B. L. Turner, Phytologia 38: 461. 1978. Basionym: Perezia glandulifera D. L. Nash. TYPE: Guatemala. Dept. Huehuetenango: canyon of Rio Selequa, 40 km NW of Huehuetenango, L. O. Williams et al. 41167 (isotype, NY not seen, NY photo at LP!).

25. Acourtia glomeriflora (A. Gray) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Gochnatia glomer!flora A. Gray. TYPE: Mexico. Cuernavaca, D. Bilimek 512 (holotype, GH not seen).

26. Acourtia gracilis R. L. Cabrera, Acta Bot. Mex. 20: 36. 1992. TYPE: Mexico. Puebla: Barranca de Patla, Villa Juarez, E. Matuda 38750 (holotype, MICH not seen).

27. Acourtia grandifolia (S. Watson) Reveal & R. M. King, Phytologia 27: 229. 1973. Basionym: Perezia grandifolia S. Watson. TYPE: Mexico. Jalisco: hillsides near Guadalajara, C. G. Pringle 1858 (isotype, MO, MO photo at LP!).

28. Acourtia guatemalensis B. L. Turner, Phytologia 74: 393. 1993. TYPE: Guatemala. Dept. Solola: above Lake Atitlan, ca. 3-5 km W of Panojachel, L. O. Williams et al. 25318 (isotype, US not seen, US photo at LP!).

29. Acourtia hidalgoana B. L. Turner, Phytologia 74: 394. 1993. TYPE: Mexico. State Hidalgo: 7 km al NE de Mesquititlan, sobre la carretera a Zacualtipan, J. Rzedowski 32510 (holotype, MEXU not seen).

30. Aeourtia hondurana B. L. Turner, Phytologia 38: 466. 1978. TYPE: Honduras. El Paraiso: cerros El Zapotillo, road from Zamorano to Guinope, G. Webster et al. 11982 (holotype, MICH not seen).

31. Aeourtia hooveri (McVaugh) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia hooveri McVaugh. TYPE: Mexico. Jalisco: Nevada de Colima, 11 mi. from Atenquique on the Tonila, R. McVaugh 11750 (isotype, NY not seen, NY photo at LP!).

32. Acourtia-huajuapana B. L. Turner, Phytologia 38: 455. 1978. TYPE: Mexico. State Oaxaca: 37 mi NW of Huajuapan de Leon, along highway 125, B. L. Turner P-59 (holotype, LL not seen).

33. Acourtia humboldtii (Less.) B. L. Turner, Phytologia 74: 395. 1993. Basionym: Dumerilia humboldtii Less. TYPE: Mexico. sine loc., s.d., A. Humboldt s.n. (holotype B, microfiche Herb. Willdenow 16095 not seen).

34. Acourtia intermedia R. L. Cabrera, Brittonia 53: 418. 2001. TYPE: Mexico. Durango: 166 mi NW of Durango, along hwy 145, L. Cabrera & G. Dieringer 653 (isotype, US not seen, US photo at LP!).

35. Acourtia joaquinensis R. L. Cabrera, Brittonia 53: 423. 2001. TYPE: Mexico. Queretaro: Mun. San Joaquin, ca. 1 km al S de Santo Tomas, Huerta 1324 (holotype, IEB not seen).

36. Acourtia lepidopoda (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia lepidopoda B. L. Rob. TYPE: Mexico. Morelos: near Cuernavaca, C. G. Pringle 9253 (holotype, GH not seen).

37. Acourtia lobulata (Bacig.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia lobulata Bacig. TYPE: Mexico. Oaxaca: San Juan del Estado, L. C. Smith 888 (holotype, GH not seen, photo in Bacigalupi, 1931: plate 2).

38. Acourtia longifolia (S. F. Blake) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia longifolia S. F. Blake. TYPE: Mexico. Jalisco: Calabaza, B. P. Reko 4872 (holotype, US not seen).

39. Acourtia lozanii (Greenm.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia lozanii Greenm. TYPE: Mexico. Hidalgo: between Metepec and Zontecomate stations, C. G. Pringle 8871 (isotype, MO, MO photo at LP!).

40. Acourtia macvaughii B. L. Turner, Phytologia 74: 398. 1993. TYPE: Mexico. Michoacan: 8-12 km SE of Aserradero Dos Aguas and nearly W of Aguililla, R. McVaugh 22789 (isotype, NY not seen, NY photo at LP!).

41. Acourtia macrocephala Sch. Bip. ex Seeman, Bot. Voy. Herald. 55: 315. 1856. TYPE: Mexico. Durango?: Sierra Madre, Dec-Feb 1838-1839, Seeman s.n. (holotype, K, not seen).

42. Acourtia mexiae R. L. Cabrera, Acta Bot. Mex. 20: 32. 1992. TYPE: Mexico. Jalisco: Real Alto, trail to El Tajo de Santiago, Y. Mexia 1749 (isotype, NY not seen, NY photo at LP!).

43. Acourtia mexicana (Lag. ex D. Don) H. Rob., Phytologia 69: 106. 1990. TYPE: Mexico. s.d., M. Sesse & J. M. Mocino s.n. (holotype, G not seen).

44. Acourtia michoacana (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia michoacana B. L. Rob. TYPE: Mexico. Michoacan: near Patzcuaro, C. G. Pringle 3988 (isotype, MO, MO photo at LP!).

45. Acourtia microcephala DC., Prodr. 7: 66. 1838. TYPE: United States. California: Douglas 62 (holotype, GH not seen).

46. Acourtia moctezumae Rzed. & Calderon, Acta Bot. Mex. 12: 14. 1990. TYPE: Mexico. Queretaro: 4 km al SE de Matzacintla, municipio de Landa, L. M. Chavez 217 (holotype, ENCB not seen).

47. Acourtia molinana B. L. Turner, Phytologia 38: 461. 1978. TYPE: Honduras. Dept. Morazan, Rio Guarabuqui, A. Molina R. 3048 (holotype, US not seen).

48. Acourtia moschata (La Llave & Lex.) DC., Prodr. 7: 66. 1838. Basionym: Perezia moschata La Llave & Lex. TYPE: Mexico. Michoacan de Ocampo: 24 km W of Morelia, H. D. D. Ripley & R. C. Barneby 14849 (neotype, designated by Turner, 1993a: 398, NY not seen, NY photo at LP!).

49. Acourtia nana (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia nana A. Gray. TYPE: Mexico. Chihuahua: high, dry valley near Chihuahua, J. Gregg 45 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Sine loc., s.d., Supernaugh s.n. (LP 303725).

50. Acourtia nelsonii (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia nelsonii B. L. Rob. TYPE: Mexico. State Jalisco: mountains near Talpa, E. Nelson 403 7 (holotype, GH not seen).

51. Acourtia nudicaulis (A. Gray) B. L. Turner, Phytologia 38: 467. 1978. Basionym: Perezia nudicaulis A. Gray. TYPE: Guatemala. Sine loc., s.d., Skinner s.n. (holotype, GH not seen).

52. Acourtia nudiuscula (B. L. Rob.) B. L. Turner, Phytologia 38: 467. 1978. Basionym: Perezia nudiuscula B. L. Rob. TYPE: Mexico. Nayarit: Tepic, E. Palmer 2018 (isotype, NY not seen, NY photo at LP!).

53. Acourtia oaxaeana R. L. Cabrera, Acta Bot. Mex. 20: 31. 1992. TYPE: Mexico. Oaxaca: 62 km al SE de Oaxaca, sobre la carretera a Tehuantepec, J. Panero & A. Salinas 2495 (isotype, US not seen).

54. Acourtia ovatifolia R. L. Cabrera, Sida 14: 141. 1990. TYPE: Mexico. Guerrero: 9 mi by road N of Iguala, W. R. Anderson & C. Anderson 5656 (holotype, MICH not seen).

55. Acourtia oxylepis (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia oxylepis A. Gray. TYPE: Mexico. San Luis Potosi, C. C. Parry & E. Palmer 547 (isotype, MO, MO photo at LP!).

56. Acourtia palmeri (S. Watson) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia palmeri S. Watson. TYPE: Mexico. Baja California: near Los Angeles Bay, Gulf of California, E. Palmer 527 (isotype, NY not seen, NY photo at LP!).

57. Acourtia parryi (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia parryi A. Gray. TYPE: Mexico. San Luis Potosi, C. C. Parry & E. Palmer 545 (isotype, MO, MO photo at LP!).

58. Acourtia patens (A. Gray) Reveal & R. M. King, Phytologia 27: 230. 1973. Basionym: Perezia patens A. Gray. TYPE: Mexico. Sine loc., 1839-40, Seeman s.n. (lectotype, designated by Turner 1993a: 401, GH not seen).

59. Acourtia pilulosa (Bacig.) B. L. Turner, Phytologia 74: 402. 1993. Basionym: Perezia dugesii A. Gray var. pilulosa Bacig. TYPE: Mexico. Oaxaca: below Jayacatlin, L. Smith 373 (holotype, GH not seen).

60. Acourtia pinetorum (Brandegee) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia pinetorum Brandegee. TYPE: Mexico. Baja California: Sierra de la Laguna, 22 Jan 1899, T. S. Brandegee s.n. (isotype, GH not seen).

61. Acourtia platyphylla (A. Gray) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia platyphylla A. Gray. TYPE: Mexico. Chihuahua, A. Wislizenus 165 (isotype, MO, MO photo at LP!).

62. Acourtia platyptera (B. L. Rob.) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia platyptera B. L. Rob. TYPE: Mexico. Michoacan or Guerrero: Sierra Madre, E. Langlasse 773 (isotype, US not seen).

63. Acourtia potosina R. L. Cabrera, Brittonia 53: 420. 2001. TYPE: Mexico. San Luis Potosi: ca. 32.2 km E of San Luis Potosi, 1.1 km S of jct, with Rt. 70 along road to microwave station, A. Prather & J. Soule 914 (isotype, TEX not seen).

64. Acourtia pringlei (B. L. Rob. & Greenm.) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia pringlei B. L. Rob. & Greenm. TYPE: Mexico. Michoacan: near Morelia, C. G. Pringle 5464 (holotype, GH not seen).

65. Acourtia pulchella R. L. Cabrera, Acta Bot. Mex. 20: 33. 1992. TYPE: Mexico. Aguascalientes: 3 km al E de La Congoja, sobre el camino a San Jose de Gracia, J. Rzedowski & R. McVaugh 897 (holotype, MICH not seen).

66. Acourtia purpusii (Brandegee) Reveal & R. M. King, Phytologia 27:231. 1973. Basionym: Perezia purpusii Brandegee. TYPE: Mexico. San Luis Potosi: Minas de San Rafael, C. A. Purpus 4787 (isotype, MO, MO photo at LP!).

67. Acourtia queretarana B. L. Turner, Phytologia 74: 402. 1993. TYPE: Mexico. Queretaro: Mpio. Pinal de Amoles, 13 km al NE de Pinal de Amoles, sobre la carretera a Jalpan, J. Rzedowski 48108 (holotype, TEX not seen).

68. Acourtia reticulata (Lag. ex D. Don) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Proustia reticulata Lag. ex. D. Don. TYPE: Mexico. Sine loc., 1787-1804, M. Sesse & M. Mocino 3082 (lectotype, designated by Turner 1993a: 403, M not seen).

69. Acourtia runcinata (Lag. ex D. Don) B. L. Turner, Phytologia 38: 460. 1978. Basionym: Clarionia runcinata D. Don. TYPE: Mexico. Sine loc., s.d., M. Sesse & M. Mocino s.n. (holotype, BM not seen). Additional specimen examined: United States. Texas: Neueces county, Corpus Christy bay, A. Heller 1537 (LP).

70. Acourtia rzedowskii B. L. Turner, Phytologia 74: 405. 1993. TYPE: Mexico. Puebla: 8 km al NNE de Azumbilla, sobre la carretera a Esperanza, J. Rzedowski 37173 (isotype, US not seen).

71. Acourtia scapiformis (Bacig.) B. L. Turner, Phytologia 38: 463. 1978. Basionym: Perezia scapiformis Bacig. TYPE: Mexico. Oaxaca: Las Sedas, C. G. Pringle 6015 (isotype, MO, MO photo at LP!).

72. Acourtia scaposa (S. F. Blake) B. L. Turner, Phytologia 38: 463. 1978. Basionym: Perezia scaposa S. F. Blake. TYPE: Mexico. Michoacan: Distrito Coalcoman, Aquila, G. B. Hinton et al. 15838 (holotype, US not seen).

73. Acourtia simulata (S. F. Blake) Reveal & R. M. King, Phytologia 27: 231. 1973. Basionym: Perezia simulata S. F. Blake. TYPE: Mexico. Michoacan: Distrito [District] Coalcoman, G. B. Hinton 13654 (isotype, NY not seen, NY photo at LP!).

74. Acourtia sinaloana B. L. Turner, Phytologia 74: 406. 1993. TYPE: Mexico. Sinaloa: summit of Sierra Tacuichamona, H. S. Gentry 5679 (isotype, NY not seen, NY photo at LP!).

75. Acourtia souleana B. L. Turner, Phytologia 74: 406. 1993. TYPE: Mexico. Oaxaca: 7.5 mi NW of Huahuapfin de Leon, 4.5 mi SE of Puebla, J. A. Soule & L. A. Prather 3187 (holotype, TEX not seen).

76. Acourtia tenoriensis B. L. Turner, Phytologia 58: 225. 1985. TYPE: Mexico. Pueblo: Mpio. Tamazulapan, Cerro Pericon, al NW de San Pedro Nopala, Tenorio & Romero de T. 7871 (holotype, TEX not seen).

77. Acourtia thurberi (A. Gray) Reveal & R. M. King, Phytologia 27: 231. 1973. TYPE: Mexico. State Sonora: Santa Cruz, Thurber 939 (holotype, GH not seen).

78. Acourtia tomentosa (Brandegee) Reveal & R. M. King, Phytologia 27:231. 1973. Basionym: Perezia tomentosa Brandegee. TYPE: Mexico. Puebla: Esperanza, C A. Purpus 2632 (isotype, MO, MO photo at LP!).

79. Acourtia turbinata (La Llave & Lex.) Reveal & R. M. King, Phytologia 27: 232. 1973. Basionym: Perezia turbinata La Llave & Lex. TYPE: Mexico. State Michoacfin: Municipio Senguio, 5 km al S de Chincua, J. Rzedowski 48261 (neotype, designated by Turner 1993a: 407, TEX not seen).

80. Acourtia umbratalis (B. L. Rob. & Greenm.) B. L. Turner, Phytologia 38: 465. 1978. Basionym: Perezia umbratalis B. L. Rob. & Greenm. TYPE: Mexico. Oaxaca: Tomellin Cation, C. G. Pringle 5966 (holotype, GH not seen).

81. Acourtia venturae R. L. Cabrera, Brittonia 53: 420. 2001. TYPE: Mexico. Guanajuato: Mun. Victoria, Puerto del Aire, por la Sierra de Victoria, E. Ventura & E. Lopez 9797 (holotype, IEB not seen).

82. Acourtia veracruzana B. L. Turner, Phytologia 74: 408. 1993. TYPE: Mexico. State Veracruz: Maltrata, E. Matuda 1236 (holotype, MEXU not seen).

83. Acourtia wislizenii (A. Gray) Reveal & R. M. King, Phytologia 27: 232. 1973. Basionym: Perezia wislizenii A. Gray. TYPE: Mexico. Chihuahua: base of Sierra Madre, Wislizenus 198 (isotype, MO, MO photo at LP!).

84. Acourtia wrightii (A. Gray) Reveal & R. M. King, Phytologia 27: 232. 1973. Basionym: Perezia wrightii A. Gray. TYPE: United States. Texas: on the Rio Seco and westward, also on the Rio Grande, C Wright 411 (holotype, GH not seen).

85. Acourtia zacatecana B. L. Turner, Phytologia 74: 411. 1993. TYPE: Mexico. Zacatecas: 6 km antes de Monte Escobedo, a 36 km de Huejucar, J. D. Garcia P. & S. Delgado 856 (isotype, NY not seen, NY photo at LP!).

Literature. Bacigalupi (1931), Reveal and King (1973), Cabrera-Rodriguez (1992, 2001), Cabrera-Rodriguez and Dieringer (2003).

Ameghinoa Speg.

Revista Fac. Agron. Univ. Nac. La Plata 3: 539. 1897. TYPE: Ameghinoa patagonica Speg. 47 of Figs. 47-50.

Etymology. In honor of the Argentine geologist Carlos Ameghino (1865-1936) who collected plants in Patagonia.

Shrubs stout, with brachyblasts and macroblasts. Leaves alternate; short-petiolate; blades obovate, coriaceous, palmately veined, margin dentate to lobate, glabrescent. Capitulescences corymbose; capitula short-pedunculate, homogamous, discoid; receptacle epaleate; involucre one- to three-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate to slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description (Telleria & Forcone, 2002) (109 of Figs. 107-111). Pollen prolate to subprolate, medium size, tricolporate, exine Trixis type, slightly microechinate, with polar caps (see also Crisci, 1974a).

Habitat and Distribution. Monotypic genus endemic to the Patagonian steppe in Argentina.

Species list

1. Ameghinoa patagonica Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 539. 1897. TYPE: Argentina. Prov. Santa Cruz: Dept. Deseado, Puerto Deseado, 1894, C. Ameghino s.n. (holotype, LP!). Additional specimens examined: Argentina. Prov. Neuquen: Cerro Negro, R. Maldonado 681 (LP); Prov. Chubut: Dept. Rio Senguerr, Rio Senguerr, A. Soriano 3208 (LP*).

Literature. Cabrera (1971b), Katinas (1995).

Berylsimpsonia B. L. Turner

Phytologia 74:351. 1993. TYPE: Proustia vanillosma C. Wright [= Berylsimpsonia vanillosma (C. Wright) B. L. Turner]. 48 of Figs. 47-50.

Etymology. In honor of the botanist Beryl Simpson (b. 1942), from United States, who made important contributions to the study of Asteraceae.

[FIGURES 47-50 OMITTED]

Shrubs clambering, with bifurcate, short, pseudostipulate, recurved spines. Leaves alternate; short-petiolate; blades oblong, cuneate to suborbicular, pinnately veined, margin serrulate to spinulose, pubescent. Capitulescences densely racemose to paniculate; capitula short-pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate to rounded, crowned by collector hairs. Cypselae truncate or slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Pollen not seen.

Habitat and Distribution. Genus of two species of Cuba, Haiti, Puerto Rico, and Dominican Republic.

Species list

1. Berylsimpsonia crassinervis (Urb.) B. L. Turner, Phytologia 74: 352. 1993. Basionym: Proustia crassinervis Urb. TYPE: Unknown data (type, probably B). Additional specimen examined: Haiti. Dept. du Nord: vicinity of St. Michel de l'Atalaye, E. Leonard 7754 (US).

2. Berylsimpsonia vanillosma (C. Wright) B. L. Turner, Phytologia 74: 352. 1993. Basionym: Proustia vanillosma C. Wright. TYPE: Cuba. C. Wright 3616 (isotype, US not seen, US photo at LP!). Additional specimen examined: Puerto Rico. Municipio de Salinas: Barrio Lapa, Tetas de Cayey, R. King & G. Proctor 10601 (US).

Literature. Fabris (1968), Turner (1993b).

Burkartia Crisci

Bol. Soc. Argent. Bot. 17: 242. 1976. TYPE: Perezia lanigera Hook. & Am. [= Burkartia lanigera (Hook. & Am.) Crisci]. 49 of Figs. 47-50.

Etymology. In honor of the Argentine botanist Arturo Burkart (1906-1975) who made important contributions to the knowledge of the flora of Argentina.

Shrubs dwarf, cushion-like. Leaves densely imbricate; sessile, clasping; blades subulate, coriaceous, three to five parallel veins in the sheathing portion, reduced to one vein in the blade, margin entire, revolute, pubescent. Capitulescences monocephalous or two-headed; capitula sessile, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Pollen spheroidal, elliptic in equatorial view, circular in polar view, medium size, P x E = (43-47 x 34-37) [micro]m. Tricolporate, colpi long with microgranulate membrane. Exine Trixis type, 5 [micro]m at the equator, slightly slender at the poles, occasionally with polar elevations. Nexine 1-1.5 [micro]m thick (see also Crisci, 1974a sub Perezia lanigera).

Habitat and Distribution. Monotypic genus endemic to the Patagonian steppe in Argentina.

Species list

1. Burkartia lanigera (Hook. & Am.) Crisci, Bol. Soc. Argent. Bot. 17: 243. 1976. Basionym: Perezia lanigera Hook. & Am. TYPE: Argentina. Prov. Santa Cruz: Puerto Deseado ("Port Desire"), C. Darwin 314 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Chubut: Comodoro Rivadavia, Canadon del Lagarto, A. Ruiz Leal 14704 (LP).

Literature. Crisci (1976a).

Calopappus Meyen

Reise Chile l: 315. 1834. TYPE: Calopappus acerosus Meyen. 50 of Figs. 47-50.

Etymology. From the Greek kalos, beauty, and pappos, apical tuff of bristles on the fruit, by the appearance of the pappus.

Shrubs dwarf, cushion-like. Leaves densely imbricate; sessile, clasping; blades subulate, coriaceous, three to five parallel veins in the sheathing portion, reduced to one vein in the blade, margin entire to dentate-spiny, glabrous to pubescent. Capitulescences monocephalous, terminal; capitula sessile, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae slightly constricted at the apex, glabrous; pappus of scabrid, narrowly paleaceous bristles.

Pollen Description (Crisci, 1974a; Crisci & Freire, 1986). Pollen spheroidal-prolate to spheroidal-oblate, large pollen, tricolporate, exine Calopappus type.

Habitat and Distribution. Monotypic genus endemic to the arid biogeographic region of Central Chile.

Species list

1. Calopappus acerosus Meyen, Reise Chile 1: 315. 1834. TYPE: Chile. Prov. Colchagua: Talcaregue, 19..?, J. Gay s.n. (holotype, P not seen, P photo F n. 38131 at LP!). Additional specimen examined: Chile. Prov. Aconcagua: Dept. Los Andes, Saladillo, Rio Blanco, O. Zollner 1406 (CONC).

Literature. Crisci and Freire (1986).

Cephalopappus Nees & Mart.

Nova Acta Phys.-Me& Acad. Caes. Leop.-Carol. Nat. Cur. 12: 5. 1824. TYPE: Cephalopappus sonchifolius Nees & Mart. 51 of Figs. 51-54.

Etymology. From the Greek kephale, head, and pappos, apical tuft of bristles on the fruit, probably in reference to the cypsela lacking a pappus.

Herbs perennial, acaulescent. Leaves rosulate; pseudopetiolate to petiolate; blades spathulate, pinnately veined, margin sinuate-denticulate, glabrous to pubescent. Capitulescences loosely racemose; capitula long-pedunculate, sometimes scapiform, homogamous, discoid; receptacle epaleate; involucre one- to three-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted to rostrate at the apex, glabrous, tuberculate; pappus absent.

Pollen Description (110 of Figs. 107-111). Pollen spheroidal, circular in polar view, medium size, P = E = (33-38) [micro]m. Tricolporate, colpi long with microgranulate membrane. Exine Cephalopappus type, 6 [micro]m thick, slightly slender at the poles. Nexine ca.l.5 [micro]m thick (see also Crisci, 1974a).

Habitat and Distribution. Monotypic genus endemic to eastern Brazil, generally growing close to the rivers.

Species list

1. Cephalopappus sonchifolius Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 5, tab. 1. 1824. TYPE: Brazil. Minas Gerais: ad flumen Ilheos, Dec 1816, C. Martius s.n. (holotype, M not seen, M photo F n. 20737 at LP!). Additional specimen examined: Brazil. State Rio de Janeiro: Rio de Janeiro, Jardin Botanico, A. L. Cabrera 12243 (LP*).

Literature. Baker (1884).

Criscia Katinas

Bol. Soc. Argent. Bot. 30: 60. 1994. TYPE: Onoseris stricta Spreng. [= Criscia stricta (Spreng.) Katinas]. 52 of Figs. 51-54.

[FIGURES 51-54 OMITTED]

Etymology. In honor of the Argentine botanist Jorge V. Crisci (b. 1945) for his important contributions to our knowledge of the tribe Nassauvieae.

Herbs perennial, acaulescent, scapose to scapiform. Leaves rosulate; pseudopetiolate; blades obovate to orbicular, pinnately veined, margin entire to denticulate, pubescent. Capitulescences monocephalous or few capitula, on scapes; capitula homogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically rounded, crowned by collector hairs. Cypselae rostrate at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description (111 of Figs. 107-111). Pollen prolate, medium to large size, tricolporate, exine Trixis type (see also Crisci, 1974a).

Habitat and Distribution. Monotypic genus from rocky and sandy places in northeastern Argentina, southern Brazil, and Uruguay.

Species list

1. Criscia stricta (Spreng.) Katinas, Bol. Soc. Argent. Bot. 30: 62. 1994. Basionym: Onoseris stricta Spreng. TYPE: Uruguay. Montevideo, s.d., Sello s.n. (lectotype, designated by Katinas, 1994: 62, BMI). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Tandil, Villa del Lago, J. V. Crisci 545 (LP*).

Literature. Katinas (1994).

Dolichlasium Lag.

Amen. nat. Espan.: 33. 1811. TYPE: Dolichlasium lagascae D. Don. 53 of Figs. 51-54.

Etymology. From Greek dolichos, long, and lasios, woolly, shaggy, because of the long, papillose anther tails.

Shrubs dwarf. Leaves alternate; petiolate; blades oblong to elliptic, pinnatisect, pinnately veined, lobes palmatifid, glabrescent. Capitulescences monocephalous, terminal; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bifid, branches apically truncate, crowned by collector hairs. Cypselae rostrate at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Exine Trixis type (see Crisci 1974a).

Habitat and Distribution. Monotypic genus from the Andes of western Argentina.

Species list

Dolichlasium lagascae D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Probably from Argentina. Prov. Mendoza: s.d., J. Gillies s.n. (types probably at BM, K, not seen). Additional specimen examined: Argentina. Prov. Mendoza: serranias de Uspallata, C. L. Spegazzini 1484 (LP).

Literature. Cabrera (1936, 1971b).

Liliana Katinas (1,4) John Pruski (2) Gisela Sancho (1) Maria Cristina Telleria (3)

(1) Division Plantas Vasculares, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, Argentina

(2) Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, USA

(3) Laboratorio de Sistematica y Biologia Evolutiva (LASBE), Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, Argentina

(4) Author for Correspondence; e-mail: katinas@fcnym.unlp.edu.ar

Published online: 9 December 2008
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Title Annotation:Part 3
Author:Katinas, Liliana; Pruski, John; Sancho, Gisela; Telleria, Maria Cristina
Publication:The Botanical Review
Article Type:Report
Geographic Code:1USA
Date:Dec 1, 2008
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The subfamily Mutisioideae (Asteraceae).
The subfamily Mutisioideae (Asteraceae).
The subfamily Mutisioideae (Asteraceae).
The subfamily Mutisioideae (Asteraceae).

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