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The subfamily Mutisioideae (Asteraceae).

Ainsliaea DC.

Prodr. 7: 13. 1838. TYPE: Ainsliaea pteropoda DC. [= Ainsliaea latifolia (D. Don) Sch. Bip.]. 6 of Figs. 3-6.

Diaspananthus Miq., Ann. Mus. Bot. Lugduno-Batavi 2: 186. 1866, as Diaspanthus. TYPE: Diaspananthus uniflorus (Miq.) Kitam. [-Ainsliaea uniflora Sch. Bip.].

Etymology. Named after the British physician Sir Whitelaw Ainslie (1767-1836), author of Materia medica Indica.

Herbs perennial, scapose or rarely small shrubs. Leaves usually rosulate or clustered at the median part of the stem; sessile to distinctly petiolate with petioles winged to wingless; blades linear, elliptic, obovate, deltoid to subrounded, palmately, pinnately, three- to five-veined, margin entire to partite, glabrous to pubescent on both surfaces or only beneath. Capitulescences unilaterally spicate, racemose to paniculate; capitula sessile to pedunculate, homogamous, ligulate, one- to five-flowered; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, chasmogamous or cleistogamous; chasmogamous florets with corolla ligulate, limb deeply five-lobed, with one deeper split and unilateral lobes; cleistogamous florets with corolla pseudoligulate, tubular, four- to five-lobed, with one deeper slit but without an expanded lip; anther apical appendages rounded to apiculate, tails slightly to long-papillose; style bilobed, branches dorsally papillose. Cypselae glabrous to pubescent; pappus of plumose, capillary bristles, occasionally absent.

Pollen Description. Ainsliaea (82 of Figs. 80-85) (Telleria & Katinas, 2005). Pollen spheroidal to prolate, medium to large size, tricolpate (e.g., A. lancangensis Y. Y. Qian, A. latifolia, and A. macroclinidioides Hayata) or tricolporate (e.g., A. acerifolia Sch. Bip., A. pertyoides Franch., A. spanocephala Y. C. Tseng, and A. uniflora), exine Ainsliaea type or Mutisia type, microechinate (see also Lin et al., 2005).

Habitat and Distribution. Genus of 49 Asian species distributed in Afghanistan, Bhutan, China, India, Indonesia, Japan, Kashmir, Korea, Union of Myanmar, Nepal, Pakistan, Philippines, Yaiwan, Thailand, and Vietnam.

Species list

1. Ainsliaea acerifolia Sch. Bip., Jahresber. Pollichia 18-19: 188. 1861. TYPE: Japan: Sine loc., H. Burger & H. Zollinger 272 (holotype, P!). Additional specimens examined: Japan. Kyushu: Kund-sho-jan, C. J. Maximowicz s.n. (MO); Honshu: Miyagi Prefecture, Boufford 19871 (CM*).

2. Ainsliaea angustata C. C. Chang, Sin. Contr. 5: 158. 1934. TYPE: China. Szechuan: Ma-Ping hsien, W. P. Fang 1641 (isotype, PE not seen, PE photo at LP!). Additional specimens examined: China. Sichuan: Szechuan, Tien-chuanhsien, K. L. Chu 2273 (PE, W).

3. Ainsliaea apiculata Sch. Bip., Jahresber. Pollichia 18-19: 188. 1861. TYPE: Japan: sine loc., H. Burger & H. Zollinger 266 (lectotype, designated by Freire, 2007:162, P!). Additional specimens examined: Japan. Honshu: Aichi Prefecture, Mikawa, Kita-shitara-gun, Nagura-mura, Oo-nagura, along Kamsa R., R. M. Furuse s.n. (S); Kyushu: Kagoshina Prefecture, Mt. Kaimon, Saito 1846 (US*).

4. Ainsliaea aptera DC., Prodr. 7: 14. 1838. TYPE: Nepal. India orientale, 1833, Royle s.n. (lectotype, designated by Freire, 2007: 105, K!). Additional specimen examined: India. Punjab: Simla, J. R. Drummond 22635 (UC).

5. Ainsliaea apteroides (C. C. Chang) Y. C. Tseng, Acta Phytotax. Sin. 31: 363. 1993. Basionym: Ainsliaea pteropoda DC. var. apteroides C. C. Chang. TYPE: China. Yunnan: Mi leh district, A. Henry 9907A (holotype, K not seen; isotype, US!). Additional specimens examined: China. Yunnan: Mengtze, A. Henry 9907 (MO, NY).

6. Ainsliaea bonatii Beauverd, Bull. Soc. Bot. Geneve 1: 377. 1909. TYPE: China. Yunnan: Tsong-chan, F. Ducloux 3646 (type, NY!). Additional specimens examined: China. Yunnan: Mi le, A. Henry 9955 (MO, NY).

7. Ainsliaea brandisiana Kurz, J. Asiat. Soc. Bengal 41: 318. 1872. TYPE: Myanmar. Martaban, S. Kurz 865 (holotype, G!).

8. Ainsliaea caesia Hand.-Mazz., Beih. Bot. Centralbl. 56: 469. 1937. TYPE: China. Guangdong: Kwangtung, R. Mell 874 (holotype, WU!).

9. Ainsliaea cavaleriei H. Lev., Fl. Kouy-Tcheou: 82. 1914. TYPE: China. Sud de Tin-fan, J. Cavalerie 1907 (holotype, E!).

10. Ainsliaea chapaensis Merr., J. Arnold Arbor. 21: 387. 1940. TYPE: Vietnam. Tonkin: Chapa, A. Petelot 2068 (isotype, MO!).

11. Ainsliaea cordifolia Franch. & Sav., Enum. pl. Jap. 2: 416. 1879. TYPE: Japan. Honshu: Mie Prefecture, 7 Oct. 1956, M. Furuse s.n. (epitype, designated by Freire, 2007:167, S!). Additional specimen examined: Japan. Chiba Prefecture, Awa, Awa-gun, Amatsu-machi, Mt. Kiyosumi on S side, M. Furuse s.n. (S).

12. Ainsliaea crassifolia C. C. Chang, Sin. Contr. 6: 549. 1935. TYPE: China. Yunnan: Yung-pe Mtns., G. Forrest 11070 (lectotype, designated by Freire, 2007: 113, E not seen, E photo at LP!, PE!).

13. Ainsliaea dissecta Franch. & Say., Enum. pl. Jap. 2: 416. 1879. TYPE: Japan. Honshu: Tottori Prefecture, Mt. Hakko, J. Sugimoto 10912 (epitype, designated by Freire, 2007: 169, S!). Additional specimen examined: Japan. Honshu: Aichi Prefecture, Prov. Mikawa, Mt. Hooraiji, Hooraiji-machi, Minami-shitaragun, M. Furuse s.n. (S).

14. Ainsliaea elegans Hemsl., Icon. pl. 28: t. 2747. 1902. TYPE: China. Yunnan: Mengtze, A. Henry 9108 (lectotype, designated by Freire, 2007: 116, K!). Additional specimen examined: China. Yunnan: Mar-li-po, Kwang-jin-in, K. M. Feng 12964 (KUN).

15. Ainsliaea foliosa Hand.-Mazz., Acta Horti Gothob. 12: 348. 1938. TYPE: China. [Sichuan?]: Sikang, "inter Taining (Ngata) et Taofu (Dawo), Sunglingku," H. Smith 12120 (holotype, W!).

16. Ainsliaea fragrans Champ. ex Benth., Hooker's J. Bot. Kew Gard. Misc. 4: 236. 1852. TYPE: China. Guangdong: Hong Kong, J. G. Champion 161 (holotype, K!).

17. Ainsliaea fulvipes Jeffrey & W. W. Sm., Notes Roy. Bot. Gard. Edinburgh 8: 175. 1914. TYPE: China. Yunnan: Teng-yueh, G. Forrest 7862 (lectotype, designated by Freire, 2007: 123, S!).

18. Ainsliaea glabra Hemsl., J. Linn. Soc., Bot. 23: 471. 1888. TYPE: China. Sichuan: Szechuan, Mt. Omei, E. Faber 391 (lectotype, designated by Freire, 2007: 123, K!).

19. Ainsliaea glumacea (Fr.) Sch. Bip., Jahresber. Pollichia 20-21: 407. 1863. Basionym: Hieracium glumaceum Fr. TYPE: India. Silhet, 1830, Wallich s.n. (lectotype, designated by Freire, 2007, K-WALL not seen). Additional specimen examined: India. Assam: Tserapflndzi till near Mairong, ex herb. Schlagintweit, Cat. No. 391 (US).

20. Ainsliaea gracilis Franch., J. Bot. (Morot) 8: 297. 1894. TYPE: China. Sichuan: Su-tchuen oriental, Tchen-keou-tin, R. P. Farges 1034A (type, P!).

21. Ainsliaea grossedentata Franch., J. Bot. (Morot) 8: 297. 1894. TYPE: China. Sichuan: Su-tchuen oriental, Tchen- keou-tin, R. P. Farges 333 (isotype, P!).

22. Ainsliaea henryi Diels, Bot. Jahrb. Syst. 29: 628. 1901. TYPE: China. Hubei: Hupeh, A. Henry 6639 (type, K!). Additional specimen examined: China. Hubei: Hupeh, A. Henry 7461 (NY).

23. Ainsliaea hypoleuca Diels ex H. Limpr., Repert. Spec. Nov. Regni Veg. 12: 514. 1922. TYPE: China. Sichuan: Szetchuan, Kwan Hsien, H. Limpricht 1251 (lectotype, designated by Freire, 2007: 131, W!).

24. Ainsliaea kawakamii Hayata, Icon. pl. formos. 8: 72. 1919. TYPE: China. Taiwan: Formosa, s.d., T. Kawakami s.n. (holotype, TI not seen, TI photo at LP!).

25. Ainsliaea lancangensis Y. Y. Qian, J. Trop. Subtrop. Bot. 8: 161. 2000. TYPE: China. Yunnan: Lancang, Y. Y. Qian 3185 (holotype, HITBC not seen). Additional specimen examined: China. Yunnan: Lancang, Y. Y. Qian 3343 (SMAO*).

26. Ainsliaea latifolia (D. Don) Sch. Bip., Jahresber. Pollichia 18: 190. 1861. Basionym: Liatris latifolia D. Don. TYPE: Nepal. Narainhetty Nepalensium, 24 Feb. 1803, Hamilton s.n. (holotype, BM not seen). Additional specimens examined: India, Punjab, Kasauli, J. R. Drummond 20055 (UC). China, Prov. Yunnan, Handel & Mazzetti 8501 (W*).

27. Ainsliaea linearis Makino, Bot. Mag. 23: 250. 1909. TYPE: Japan. Kyushu: Kagoshima Prefecture, Osuma, M. Togasi 1448 (neotype, designated by Freire, 2007: 175, MO!). Additional specimen examined: Japan. Kyushu: Kagoshima Prefecture, Isl. Yakushima, Yaku-cho, from Yodoko Hut along the Yodoko River, D. E. Boufford & S. Mitsuta 20062 (CM).

28. Ainsliaea macrocephala (Mattf.) Y. C. Tseng, Acta Phytotax. Sin. 31: 364. 1993. Basionym: Ainsliaea pteropoda DC. var. macrocephala Mattf. TYPE: China. Yunnan: Likiang, C. Schneider 2328 (lectotype, designated by Freire, 2007: 136, K!).

29. Ainsliaea macroclinidioides Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25, 19: 141. 1908. TYPE: Taiwan, Suizan, Nagasawa 665 (lectotype, designated by Freire, 2007: 177, TI not seen, duplicate, TI photo at LP!). Additional specimen examined: Japan. Ryukyu: Okinawa Prefecture, Isl. Ishigaki, Mt. Omoto-dake, G. Murata 17066 (AAU*).

30. Ainsliaea mairei H. Lev., Le Monde des Plantes 18: 31. 1916. TYPE: China. Yunnan: Mt. Io-Chan, Nov 1912, E. E. Maire s.n. (holotype, E!).

31. Ainsliaea nana Y. C. Tseng, Acta Phytotax. Sin. 31: 365. 1993. TYPE: China. Sichuan: Tianquan, Chung-luen Wu 12307 (holotype, NAS not seen)..

32. Ainsliaea nervosa Franch., Bull. Mus. Hist. Nat. (Paris) 1 : 64. 1895. TYPE: China. Yunnan: Long-Ki, J. M. Delavay 4939 (holotype, P not seen, P photo at LP!).

33. Ainsliaea oblonga Koidz., Bot. Mag. (Tokyo) 28: 150. 1914. TYPE: Japan. Yaeyama archipelago: insl. Nisiomotesima, Aug 1889, Tashiro s.n. (holotype, TI not seen, TI photo at LP!).

34. Ainsliaea parvifolia Merr., Philipp. J. Sci. 12: 110. 1917. TYPE: China. Guangdong: Loh Fau Mtn. (Lofaushan), E. D. Merrill 10237 (holotype, US!).

35. Ainsliaea paueicapitata Hayata, Icon. pl. formos. 8: 71. 1919. TYPE: Taiwan. Nokosan, Jan 1908, U. Mori s.n. (type, TI not seen, TI photo at LP!).

36. Ainsliaea pentaflora S. E. Freire, Novon 12: 451. 2002. TYPE: Vietnam. Tonkin: Massif du Tom Dav, A. Petelot 417 (holotype, C!).

37. Ainsliaea pertyoides Franch., J. Bot. (Morot) 2: 70. 1888. TYPE: China. Yunnan: ad fauces Kien-min-keou, prope Mo-so-yn, J. M. Delavay 1218 (lectotype, designated by Freire, 2007: 143, P!). Additional specimen examined: China. Yunnan, 28 Jan 1889, J. M. Delavay s.n. (UC*).

38. Ainsliaea pingbianensis Y. C. Tseng, Acta Phytotax. Sin. 31: 365. 1993. TYPE: China. Yunnan: Pingbian, P. Mao 4186 (holotype, IBSC not seen, IBSC photo at LP!).

39. Ainsliaea qianiana S. E. Freire, Novon 12: 453. 2002. TYPE: China. Yunnan: Colline de Kin-tchong-chan, E. E. Maire 2529 (holotype, W!; isotypes, NY!, UC!).

40. Ainsliaea ramosa Hemsl., J. Linn. Soc., Bot. 23: 471. 1888. TYPE: China. Hubei: Hupeh, Nanto and mtns. to the N, A. Henry 4422 (holotype, K not seen; isotype, US!).

41. Ainsliaea reflexa MelT., Philipp. J. Sci. 1: 242. 1906. TYPE: Philippines. Luzon: Lepanto, Mt. Data, E. D. Merrill 4571 (holotype, NY!).

42. Ainsliaea rubrinervis C. C. Chang, Sin. Contr. 4: 226. 1934. TYPE: China. Sichuan: Kwan-hsien, W. P. Fang 2076 (holotype, N not seen; isotype, NY!).

43. Ainsliaea smithii Mattf., Acta Horti Gothob. 8: 79. 1933. TYPE: China. Sichuan: Sze-chuan, Drogochi, H. Smith 4698 (isotype, W!).

44. Ainsliaea spanocephala Y. C. Tseng, Acta Phytotax. Sin. 26: 74. 1988. TYPE: Thailand. Loei: Phu Luang, Wildlife Sanctuary, H. Koyama, H. Terao & W. Thawatchai 33699 (isotypes, BKF!, GH!, KYO!). Additional specimens examined: Thailand. Loei: Phu Luang, Wildlife Sanctuary, 1982, T. Smitinand s.n. (BKF 077323*); id., Nuijomdha & Vidal 438 (C*).

45. Ainsliaea spicata Vaniot, Bull. Acad. Int. Geogr. Bot. 12: 117. 1903. TYPE: China. Yunnan: environs de Yun-nan-sen, F. Ducloux 9D (holotype, E!; isotype, W!).

46. Ainsliaea trinervis Y. C. Tseng, Acta Phytotax. Sin. 31: 367. 1993. TYPE: China. Guangxi: Rongshui, S. Q. Chen 16409 (holotype, IBSC not seen; isotype, TI not seen, TI photo at LP!).

47. Ainsliaea uniflora Sch. Bip., Jahresber. Pollichia 18: 187. 1861. TYPE: Japan. H. Burger & H. Zollinger 283 (type, K!). Additional specimen examined: Honshu, Mie, Ise, Yamada, 10 Oct 1911, s. leg. (US 1132181*).

48. Ainsliaea walkeri Hook. f., Bot. Mag. 102: tab. 6225. 1876. TYPE: China. Guangdong: Hong Kong, Dec 1875, Capt. A. L. Wilkes s.n. (holotype, K!).

49. Ainsliaea yunnanensis Franch., J. Bot. (Morot) 2: 70. 1888. TYPE: China. Yunnan: Che-tcho-tze, supra Tapin-tze, J. M. Delavay 608 (type, NY!).

Literature. Freire (2002, 2007), Mitsui et al. (2007).

Aphyllocladus Wedd.

Chlor. andina 1 : 11. 1855. TYPE: Aphyllocladus spartioides Wedd. 7 of Figs. 7-10. Jobaphes Phil., F1. atacam.: 27. 1860. TYPE: Jobaphes virgatus Phil. [= Aphyllocladus denticulatus (J. Remy) Cabrera].

Etymology. From the Greek a, without, phyllon, leaf, and clados, branch, with reference to the aphyllous branches.

Shrubs. Leaves alternate, reduced, early caducous; sessile; blades linear-oblong to linear-ovate, pinnately or three- to five-veined, margin entire, subglabrous to lanuginous on both surfaces. Capitulescences monocephalous, terminal; capitula pedunculate, homogamous or heterogamous, radiate or disciform; receptacle epaleate; involucre multiseriate. Florets dimorphic or isomorphic; marginal florets bisexual or female, corolla bilabiate, outer lip expanded or not; central florets or all florets bisexual, with corolla tubular-funnelform, deeply five-lobed; anther apical appendages truncate, reddish, tails papillose; style bifid, branches dorsally papillose. Cypselae densely long-villose; pappus of scabrid to barbellate bristles, the innermost flat, longer, and wider than the others and barbellate at the apex.

Pollen Description. Aphyllocladus (83 of Figs. 80-85) (Telleria & Katinas, 2004). Pollen prolate to subprolate, large size, tricolporate, exine Mutisia type, microechinate.

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

Species list

1. Aphyllocladus decussatus Hieron., Icon. Descr. Pl.: 28.1855. TYPE: Argentina. Prov. La Rioja: Los Corrales, Sierra de Famatina, G. H. E. W. Hieronymus & G. Niederlein 831 (holotype, B, not seen, B photo F n. 15873 at LP!; isotype, GH not seen).

2. Aphyllocladus denticulatus (J. Remy) Cabrera, Darwiniana 9: 367. 1951. Basionym: Cyclolepis denticulata J. Remy. TYPE: Chile. Copiapo, C. Gay 239 (holotype, P not seen, P photo F n. 38078 at LP!). Additional specimen examined: Chile. Prov. Atacama: Bandurrias, s.d., N. Geisse s.n. (LP).

3. Aphyllocladus ephedroides Cabrera, Darwiniana 9: 370. 1951. TYPE: Argentina. Prov. La Rioja: Dept. General Sarmiento, Vinchina, G. Covas 1229 (holotype, LP!).

4. Aphyllocladus sanmartinianus Molfino, Bol. Soc. Argent. Bot. 5: 30. 1953. TYPE: Argentina. Prov. San Juan: Iglesias, Cuesta de la Casitas, R. A. Spegazzini 230 (holotype, BAB, not seen). Additional specimen examined: Argentina. Prov. San Juan: camino a Las Hornillas, A. L. Cabrera et al. 24290 (LP*).

5. Aphyllocladus spartioides Wedd., Chlor. andina 1: 11. 1855. TYPE: Bolivia. Chuquisaca and Potosi, 1843-1847, H. A. Weddell s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Humahuaca, Humahuaca, A. Ruiz Leal 14386 (LP).

Literature. Cabrera (1951, 1978), Molfino (1953), Freire (1996).

Brachyclados D. Don

Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Brachyclados lycioides D. Don. 8 of Figs. 7-10.

Lavidia Phil., Anales Univ. Chile 85: 837. 1894. TYPE: Lavidia caespitosa Phil. [= Brachyclados caespitosus (Phil.) Speg.].

Etymology. From the Greek brachys, short, and klados, branch, with reference to the short shoots or brachyblasts.

Shrubs stout to cushion-like, with brachyblasts and macroblasts. Leaves alternate, rigid; sessile, clasping; blades linear, ovate, subulate to oblong, parallel veins in the sheathing portion and one prominent central vein in the blade, margin entire, revolute, glabrous above, pubescent beneath. Capitulescences monocephalous, terminal; capitula subsessile, 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, branches dorsally papillose. Cypselae truncate to shortly attenuate at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Brachyclados (84 of Figs. 80-85) (Telleria & Katinas, 2004). Pollen subprolate to prolate, large size, tricolporate, exine Mutisia type, microechinate or almost psilate (see also Parra & Marticorena, 1972).

Habitat and Distribution. Genus of three species of central Chile, Patagonia, and western Argentina.

Species list

1. Brachyclados caespitosus (Phil.) Speg., Anales Soc. Ci. Argent. 53: 28. 1902. Basionym: Lavidia caespitosa Phil. TYPE: Chile. Patagonia, Lago Pinto, s.d., H. Ibar s.n. (type, probably SGO). Additional specimen examined: Argentina. Prov. Chubut: cerca de Facundo, A. L. Cabrera et al. 23227 (LP).

2. Brachyclados lycioides D. Don, Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido de Patagones, cerca de Cardenal Cagliero, A. L. Cabrera & H. A. Fabris 14865 (LP*).

3. Brachyclados megalanthus Speg., Anales Soc. Ci. Argent. 53: 29. 1902. TYPE: Argentina. Prov. Buenos Aires: Tres Cerros to Rio Negro, Carmen de Patagones, Feb 1898, C. L. Spegazzini s.n. (syntypes, LP!).

Observation. Brachyclados lycioides constitutes a very important element in the diet composition of the guanaco (Lama guanicoe) (Candia & Dalmasso, 1995).

Literature. Cabrera (1971 b), Katinas (1996a).

Catamixis Thomson

J. Linn. Soc., Bot. 9: 342. 1867. TYPE: Catamixis baecharoides Thomson. 9 of Figs. 7-10.

Etymology. From the Greek katamixis, katameixis, admixture, or kata and mixis, a mixing, mingling because the affinities in Catamixis point in many directions.

Subshrubs branched, erect. Leaves alternate; short-petiolate; blades obovate to oblong, pinnately veined, margin obtusely dentate-serrate, pubescent to glabrate. Capitulescences corymbose; capitula pedunculate, homogamous, ligulate; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb shallowly five-lobed; anther apical appendages acute, tails papillose; style bifid, branches dorsally papillose. Cypselae truncate at the apex, densely pubescent; pappus of scabrid, capillary bristles, unequal in length.

Pollen Description. Pollen not seen.

Habitat and Distribution. Monotypic genus endemic to the narrow range in the Siwalik belt and Theri district of Garwhal Himalayas in India and Nepal.

Species list

1. Catamixis baccharoides Thomson, J. Linn. Soc., Bot. 9: 343. 1867. TYPE: India. Uttaranchal: District Garhwal, Siwalicis Himalaya, s.d., s. leg. (syntypes, K not seen, photos at LP!).

Observation. The ligulate corolla of Catamixis results unusual for Mutisioideae, where there is a predominance of bilabiate corollas (although some species of Mutisia have also ligulate florets). According to Small (1919), all five petals in the corolla of Catamixis are fused and the split is posterior (the posterior lip corresponds to the two-lobed lip in the typical bilabiate corolla) so that something like a ligulate corolla results. But this is just one more variation in the manner of splitting of the variable corollas of Mutisioideae and cannot be taken as an intermediate type leading to the Cichorieae (Small, 1919).

The genus and its only species could be on the verge of extinction (Nayar & Ahmedullah, 1985).

Literature. Nayar and Ahmedullab (1985), Rao (1995).

Chaetanthera Ruiz & Pav.

Fl. peRUv. prodr.: 106. 1794. TYPE: Chaetanthera ciliata Ruiz & Pay. 10 of Figs. 7-10. Aldunantea J. Remy, Fl. chil. 3: 320. 1849. TYPE: not designated, based on the three species Aldunantea chilensis J. Remy, A. gnaphalioides J. Remy, and A. lycopodioides J. Remy.

Carmelita Gay ex DC., Prodr. 7: 14. 1838. [= Chaetanthera subgen. Carmelita (Gay) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 97. 1937]. TYPE: Carmelita formosa Gay ex DC. [= Chaetanthera villosa D. Don].

Cherina Cass., Dict. sci. nat. 8: 437. 1817. TYPE: Cherina micropylla Cass. [= Chaetanthera microphylla (Cass.) Hook. & Arn.].

Chondrochilus Phil., Linnaea 28: 711. 1858. TYPE: Chondrochilus crenatus Phil. [=Chaetanthera renifolia (J. Remy) Cabrera].

Egania J. Remy, Fl. chil. 3: 324. 1849. [= Chaetanthera subgen. Egania (J. Remy) Reiche, Fl. chil. 4: 355. 1905]. TYPE: not designated, based on the three species Egania acerosa J. Remy, E. apiculata J. Remy, and E. dioica J. Remy.

Elachia DC., Prodr. 7: 256. 1838. TYPE: Elachia euphrasioides DC. [= Chaetanthera euphrasioides (DC.) F. Meigen].

Euthrixia D. Don, Trans. Linn. Soc. London 16: 257. 1830. TYPE: Euthrixia salsoloides D. Don [= Chaetanthera microphylla (Cass.) Hook. & Arn.].

Minythodes Phil., Anales Univ. Chile 115: 317. 1904, nom. nudum.

Oriastrum Poepp. & Endl., Nov. gen. sp. pl. 3: 50. 1842. [= Chaetanthera subgen. Oriastrum (Poepp. & Endl.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 96. 1937]. TYPE: Oriastrum pusillum Poepp. & Endl. [= Chaetanthera pulvinata (Phil.) Hauman].

Proselia D. Don, Trans. Linn. Soc. London 16: 234. 1830. [= Chaetanthera subgen. Proselia D. Don, 1. c.]. TYPE: Proselia serrata D. Don [= Chaetanthera serrata Ruiz & Pav.].

Tiltilia Phil., Anales Univ. Chile 115: 323. 1904, nom. nudum.

Tylloma D. Don, Trans. Linn. Soc. London 16: 238. 1830. [= Chaetanthera subgen. Tylloma (D. Don) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 97. 1937]. TYPE: Tylloma limbatum D. Don [= Chaetanthera limbata (D. Don) Less.].

Etymology. From the Greek chaite, bristle, long hair and anthera, anther, alluding to the papillose tails of the anthers.

Herbs caulescent, stout to cushion-like, annual or perennial, rarely shrubs, monoecious, gynodioecious, or dioecious. Leaves alternate, sparse to densely imbricate; sessile to clasping; blades linear, subulate, oblanceolate to spathulate, pinnately, three-veined, or with a conspicuous central vein, margin entire, ciliate to dentate, glabrous to densely pubescent. Capitulescences monocephalous, terminal; capitula sessile to pedunculate, homogamous or heterogamous, discoid, disciform, or radiate; receptacle epaleate; involucre multiseriate, phyllaries acute at the apex or with an apical, showy appendage. Florets isomorphic in dioecious and gynodioecious plants, or dimorphic in monoecious plants; marginal florets female or neuter, with or without staminodes, corolla bilabiate, outer lip expanded or not, inner lip developed to absent, sub-bilabiate (3+1 corolla lips), or true ray; central florets bisexual or male, corolla bilabiate to tubular-bilabiate; female capitula with all florets female, bilabiate; anther apical appendages acute, tails papillose; style bilobed to bifid, branches dorsally papillose. Cypselae truncate at the apex, glabrous to pubescent; pappus of scabrid, simple to barbellate, capillary bristles.

Pollen Description. Chaetanthera (85 of Figs. 80-85) (Telleria & Katinas, 2004). Pollen spheroidal--subprolate to prolate, size medium to large, tricolporate, with endocingulus, exine Mutisia type, microechinate. Two pollen types were recognized, Type I (found in 14 species) with pollen subrectangular in equatorial view, subcircular or subtriangular in polar view, medium size, nexine outline elliptic to subrectangular in shape, exine 5-7 [micro]m thick; and Type II (found in 25 species) with pollen elliptic in equatorial view, large size, nexine outline dumbbell-shaped, exine 11-18 [micro]m thick. In the type II, two subtypes were recognized (see also Parra & Marticorena, 1972; Markgraf & D'Antoni, 1978; Wingenroth & Heusser, 1983).

Habitat and Distribution. Genus of 45 South American species distributed mainly in the Andes, from Peru to southern Argentina and Chile.

Species list

1. Chaetanthera acerosa (J. Remy) Benth. & Hook. f. ex Griseb., Symb. ft. argent.: 214. 1879. Basionym: Egania acerosa J. Remy. TYPE: Sine loc., s.d., C. Gay s.n. (types probably GH, P). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Rafael, valle de Las Lenas, A. Ruiz Leal 24661 (LP).

2. Chaetanthera acheno-hirsuta (Tombesi) Arroyo, A. M. R. Davies & Till-Bottr., Gayana, Bot. 61: 28. 2004. Basionym: Chaetanthera pulvinata (Phil.) Hauman var. acheno-hirsuta Tombesi. TYPE: Argentina. Prov. San Juan: Dept. Iglesia, Mina Las Carachas, E. Nicora et al. 8217 (holotype, SI!).

3. Chaetanthera apiculata (J. Remy) F. Meigen, Bot. Jahrb. Syst. 18: 456. 1894. Basionym: Egania apiculata J. Remy. TYPE: Chile. Coquimbo, Pasto Blanco, Los Patos, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Prov. Santiago: cordillera Rio San Francisco, Fierro Carrera, E. Werdermann 627 (LP).

4. Chaetanthera australis Cabrera, Notas Mus. La Plata, Bot. l : 64. 1935. TYPE: Argentina. Prov. Rio Negro: San Carlos de Bariloche, arroyo Nirico, A. Burkart 6599 (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquen: near San Martin de los Andes, Bocher et al. 1658 (LP).

5. Chaetanthera aymarae Martic. & Quezada, Bol. Soc. Biol. Concepcion 48: 107. 1974. TYPE: Chile. Prov. Tarapaca Dept. Arica, camino de Zaphahuira a Putre, Quebrada de Socoroma, M. Ricardi et al. 166 (holotype, CONC!).

6. Chaetanthera boliviensis J. Kost., Blumea 5: 673. 1945. TYPE: Bolivia. Choquecatachico, T. Herzog 2339 (type, probably M).

7. Chaetanthera brachylepis Phil., Anales Univ. Chile 87: 12. 1894. TYPE: Chile. Prov. Arauco: Cordillera de Nahuelbuta, vega de Rucapillan, Jan 1877, R. Philippi s.n. (holotype, S not seen, S photo F n. 15960 at LP!). Additional specimen examined: Chile. Prov. Curico: Potrero Grande, Lomas Blancas, E. Barros 7443 (LP*).

8. Chaetanthera chilensis (Willd.) DC., Ann. Mus. Natl. Hist. Nat. 19: 70. 1812. Basionym: Perdicium chilense Willd. TYPE: Chile. Prov. Nuble: Cucha Cucha, s.d., s. leg., herb. Willdenow n. 16092 (holotype, BD not seen, BD photo at LP!). Additional specimen examined: Chile. Prov. Nuble: Chillan, Bureo, E. Barros 7441 (LP).

9. Chaetanthera chiquianensis Ferreyra, Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, Bot. 6: 5. 1953. TYPE: Peru. Dept. Ancash: Prov. Bolognesi, cerro al S de Chiquian, E. Cerrate 1323 (isotype, LP!).

10. Chaetanthera ciliata Ruiz & Pav., Syst. veg. ft. peruv, chil.: 190. 1798. TYPE: Chile. Prov. Maule: Huiquilemu, s.d., s. leg. (type, probably MA). Additional specimen examined: Chile. Prov. Nuble, Brochers 1071c (LP).

11. Chaetanthera cochlearifolia (A. Gray) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Oriastrum cochlear(/hilum A. Gray. TYPE: Peru. Alpamarca, s.d., leg. U. S. Explor. Exped. under Cap. Wilkes s.n. (holotype, GH not seen). Additional specimen examined: Peru. Prov. Junin: Casapalca, Macbride & Featherstone 845 (LP).

12. Chaetanthera dioica (J. Remy) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Egania dioica J. Remy. TYPE: Chile. Cordilleras de Coquimbo, cerca de Pasto Blanco, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Argentina. Prov. La Rioja: Sierra de Famatina, La Mesada, Rio Amarillo superior, F. Kurtz 13717 (LP).

13. Chaetanthera elegans Phil., Linnaea 28: 712. 1856. TYPE: Chile. Prov. Concepcion: Los Angeles, C. Gay 806 (isotype, S not seen). Additional specimen examined: Chile. Prov. Linares: Cordillera de Linares, Los Hualis, E. Barros 2410 (LP).

14. Chaetanthera euphrasioides (DC.) F. Meigen, Bot. Jahrb. Syst. 17: 284. 1893. Basionym: Elachia euphrasioides DC. TYPE: Chile. Altos Andes, s.d., C. Gay s.n. (type, probably GH, P). Additional specimen examined: Chile. Prov. Santiago: Quebrada de Jorquera, cerca del Refugio de Farellones, cordillera de Las Condes, G. Looser 5751 (LP).

15. Chaetanthera flabellata D. Don, Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Chile. Cordilleras, Cummings" 291 (type, probably GH). Additional specimen examined: Chile. Prov. Santiago: Piuquencillos, valle del Rio Colorado, Pisano et al. 1641 (LP).

16. Chaetanthera flabellifolia Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 153. 1937, nora. novum. Basionym: Pachylaena elegans Phil. TYPE: Chile. Prov. Coquimbo, Bafios del Toro, 1860-1861, H. Volckmann s.n. (holotype, S not seen). Additional specimen examined: Chile. Prov. Coquimbo: Banos del Toro, E. Werdermann 189 (LP).

17. Chaetanthera glabrata (DC.) F. Meigen, Bot. Jahrb. Syst. 23: 456. 1894. Basionym: Tylloma glabratum DC. TYPE: Chile. Collina, s.d., C. Macrae s.n. (type, probably K). Additional specimen examined: Chile. Prov. Elqui: Dept. Elqui, Vicuna, Cuesta de Elqui, G. Looser 4389 (LP).

18. Chaetanthera glandulosa J. Remy, Fl. chil. 3:311. 1847. TYPE: Chile: s.d., C. Gay s.n. (isotype, B not seen, B photo F n. 15968 at LP!). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Morro Blanco, Cabreria, C. Jiles 1237 (LP).

19. Chaetanthera gnaphalioides (J. Remy) I. M. Johnst., Physis (Buenos Aires) 9: 325. 1929. Basionym: Aldunatea gnaphalioides J. Remy. TYPE: Chile. Prov. Coquimbo: cerros de las cordilleras de Dona Ana, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Dept. Elqui: 3 km al interior de Rivadavia, en el camino a Huanta, Wagenknecht 4388 (LP).

20. Chaetanthera incana Poepp. ex Less., Linnaea 5: 284. 1830. TYPE: Chile. Prov. Aconcagua: Concon, E. F. Poeppig 218 (holotype, B not seen, B photo F n. 15969 at LP!; isotype, BD not seen). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Illapel, al N de Los Vilos, C. Jiles 2326 (LP).

21. Chaetanthera kalinae A. M. R. Davies, Novon 16: 51. 2006. TYPE: Chile. Prov. Elqui: Vicuna, Embalse La Laguna, C. Ehrhart 61 (isotype, MO not seen).

22. Chaetanthera lanata (Phil.) I. M. Johnst., Physis (Buenos Aires) 9: 325. 1929. Basionym: Tylloma lanatum Phil. TYPE: Chile. Prov. Atacama: Vaquillas, s.d., R. Philippi s.n. (isotype, S not seen). Additional specimen examined: Chile. Prov. Coquimbo: Banos del Toro, A. L. Cabrera 3550 (LP!).

23. Chaetanthera leptocephala Cabrera, Notas Mus. La Plata, Bot. 17: 78. 1954. TYPE: Chile. Prov. Coquimbo: Quebrada del Toro, 20 km E de Hurtado, hacienda E1 Bosque, E. Wagenknecht 18490 (holotype, CONC!). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Illapel, hacienda Cuncumen, Cuncumen, a 77 km al E de Illapel, Munoz & Johnson 2192 (LP).

24. Chaetanthera limbata (D. Don) Less., Syn. gen. Compos.: 116. 1832. Basionym: Tylloma limbatum D. Don. TYPE: Chile. Prov. Coquimbo: Coquimbo, s.d., Caldcleugh s.n. (holotype, G not seen, G photo F n. 28856 at LP!). Additional specimen examined: Chile. Prov. El Loa: Atacama, Estacion Cuba, E. Barros 2402 (LP).

25. Chaetanthera linearis Poepp. ex Less., Syn. gen. Compos.: 112. 1832. TYPE: Chile: s.d., C. Bertero 912 (type, probably GH). Additional specimen examined: Chile. Prov. Santiago: cordillera de Santiago, Refugio de Farellones, G. Looser 5214 (LP).

26. Chaetanthera lycopodioides (J. Remy) Cabrera, Revista Centro Estud. Doct. Ci. Nat. 1: 58. 1935. Basionym: Aldunantea lycopodioides J. Remy. TYPE: Chile. Santiago, Cordillera de Los Patos, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Prov. Los Andes: Potrero Escondido, O. Boeleke 2468 (LP).

27. Chaetanthera microphylla (Cass.) Hook. & Am., Comp. Bot. Mag. 1: 104. 1835. Basionym: Cherina microphylla Cass. TYPE: Chile, s.d., s. leg. herb. M. de Jussieu s.n. (type, probably P). Additional specimen examined: Argentina. Prov. Neuquen: Andacollo, Mina Guaraco, A. L. Cabrera 11120 (LP).

28. Chaetanthera minuta (Phil.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 134. 1937. Basionym: Tylloma minutum Phil. TYPE: Chile. Prov. Atacama: Pastos Largos, 6 Jan 1885, F. Philippi s.n. (isotype, S not seen). Additional specimen examined: Chile. Prov. Atacama: Dept. Copiapo, Mondaca, A. Krapovickas & J. H. Hunziker 5 700 (LP).

29. Chaetanthera moenchioides Less., Syn. gen. Compos.: 113. 1832. TYPE: Chile. Prov. Concepcion: Antuco, s.d., E. F. Poeppig 208 (holotype, BD not seen, BD photo F n. 15974 at LP!). Additional specimen examined: Chile. Chacabuco: Colina, s.d., s. leg. (LP).

30. Chaetanthera pentacaenoides (Phil.) Hauman, Anales Soc. Ci. Argent. 86: 317. 1918. Basionym: Oriastrum pentacaenoides Phil. TYPE: Argentina. Prov. Mendoza: entre Mendoza y Santa Rosa, 1869, s. leg. (holotype, S not seen). Additional specimen examined: Argentina. Prov. Mendoza: Dept. Las Heras, Alto de los Manantiales, A. Ruiz Leal & F. Roig 23617 (LP).

31. Chaetanthera perpusilla (Wedd.) A. Anderb. & S. E. Freire, Taxon 39: 431. 1990. Basionym: Luciliopsis petpusilla Wedd. TYPE: Bolivia. Dept. La Paz: Ravin de Chuquiaguillo, La Lancha, 1851, H. A. Weddell s.n. (holotype, P!).

32. Chaetantheraperuviana A. Gray, Proc. Amer. Acad. Arts 5: 144. 1861. TYPE: Peru. Andes, Banos, s.d., leg. U. S. Explor. Exped. under Capt. Wilkes s.n. (holotype, GH not seen). Additional specimen examined: Peru. Dept. Ancash: Prov. Bolognesi, Rumipufuio, E. Cerrate et al. 6490 (LP).

33. Chaetanthera planiseta Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 128, nom. novum. Basionym: Oriastrum pusillum Poepp. & Endl. TYPE: Chile. Andes, Cordillera de San Jago, s.d., s. leg. (type, not localized). Additional specimen examined: Chile. Prov. Cachapoal: Rancagua, cordillera de Codegua, E. Barros 9849 (LP).

34. Chaetanthera pulvinata (Phil.) Hauman, Physis (Buenos Aires) 3: 420. 1917. Basionym: Oriastrum pulvinatum Phil. TYPE: Chile. Prov. Coquimbo: Cordillera de Dona Ana, 1861, H. Volckmann s.n. (holotype, S not seen). Additional specimen examined: Argentina. Prov. San Juan: Valle del Cura, pirca de los Ingenieros, R. Pdrez Moreau 158 (LP).

35. Chaetantherapusilla (D. Don) Hook. & Am., Comp. Bot. Mag. 1: 106. 1835. Basionym: Tylloma pusillum D. Don. TYPE: Argentina. Prov. Mendoza: Andes de Mendoza, s.d., J. Gillies s.n. (type, probably GH). Additional specimen examined: Chile. Prov. Santiago: cordillera de Santiago, Jan 1872, Germain s.n. (LP).

36. Chaetanthera renifolia (J. Romy) Cabrera, Revista Mus. La Plata, Secc. Bot. 1 : 149. 1937. Basionym: Elachia renifolia J. Romy. TYPE: Chile. Prov. Santiago: La Polvadera, s.d., s. leg. (holotype, P? not seen, photo F n. 38089 at LP!). Additional specimen examined: Chile. Area Metropolitana: Cajdn de la Yerba Loca, Feb 1891, R. Philippi s.n. (LP).

37. Chaetanthera revoluta (Phil.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 106. 1937. Basionym: Egania revoluta Phil. TYPE: Chile. Prov. Antofagasta: entre Aguas Calientes y Socaire, 5 Feb 1885, F. Philippi s.n. (isotype, S not seen, S photo F n. 15979 at LP!). Additional specimen examined: Argentina. Prov. Salta: Dept. San Antonio de los Cobres, abra Alto Chorrillos, A. L. Cabrera 8874 (LP*).

38. Chaetanthera serrata Ruiz & Pav., Syst. veg. ft. peruv, chil.: 191. 1798. TYPE: Chile. Prov. Concepcion: Concepcion, s.d., H. Pavon s.n. (syntype, BM not seen, photo at LP!). Additional specimen examined: Chile. Prov. Concepci6n: Tome, Lomas de Cocholzue, Junge 1282 (LP).

39. Chaetanthera spathul(folia Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 141. 1937. TYPE: Chile. Prov. Coquimbo: Illapel, E1 Rodel, Jan 1888, Z. Vergara s.n. (isotype, S not seen). Additional specimen examined: Argentina. Prov. Mendoza: Cuchilla del Diablo, 15 Feb 1934, R. POrez Moreau s.n. (LP).

40. Chaetanthera sphaeroidalis (Reiche) Hicken, Darwiniana 1: 41. 1922. Basionym: Oriastrum sphaeroidale Reiche. TYPE: Chile. Prov. Atacama: desierto de Atacama, 1897, E. Delaigne s.n. (isotype, S not seen). Additional specimen examined: Chile. Prov. Huasco: Atacama, Vallenar, cordillera Laguna Chica, E. Werdermann 253 (LP).

41. Chaetanthera splendens (J. Romy) B. L. Rob., Proc. Amer. Acad. Arts 49: 514. 1913. Basionym: Elachia splendens J. Romy. TYPE: Chile. Prov. Santiago: colina Chimborango, etc., s.d., s. leg. (type, not localized). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Cabreria, C. Jiles 1567 (LP).

42. Chaetanthera stuebelii Hieron., Bot. Jahrb. Syst. 21: 368. 1896. TYPE: Bolivia. Sicasica, entre Tomarape y La Paz, A. Stuebel 15a (holotype, BD not seen). Additional specimen examined: Argentina. Prov. La Rioja: Sierra de Famatina, ladera este, A. Krapovickas & J. H. Hunziker 5332 (LP).

43. Chaetanthera tenella Less., Syn. gen. Compos.: 114. 1832. TYPE: Chile. Cachapual: Rancagua and Quillota, C. G. Bertero 162, 910 (probable syntypes, NY, NY photos at LP!).

44. Chaetanthera valdiviana Phil., Linnaea 28: 712. 1856. TYPE: Chile. Prov. Valdivia: San Juan, Jan 1855, R. Philippi s.n. (isotype S, not seen). Additional specimen examined: Chile. Prov. Valdivia: San Juan, R. Philippi 1368 (LP).

45. Chaetanthera villosa D. Don, Philos. Mag. Ann. Chem. 11: 391. 1832. TYPE: Argentina, Prov. Mendoza: El Planchon, s.d., J. Gillies s.n. (syntype, GH not seen). Additional specimen examined: Argentina. Prov. Rio Negro: Cerro Utne, C. C. Hosseus 435 (LP).

Literature. Cabrera (1937), Koster (1945), Ferreyra (1953), Cabrera (1954), Marticorena and Quezada (1974), Anderberg and Freire (1990), Arroyo et al. (2004), Davies (2006), Hershkovitz et al. (2006).

Chaptalia Vent., nom. cons.

Descr. pl. nouv. 7: pl. 61. 1802. TYPE: Chaptalia tomentosa Vent. 11 of Figs. 11-14. Leria DC., Ann. Mus. Natl. Hist. Nat. 19: 68. 1812, non Adans., 1763, nom. illeg. [= Chaptalia sect. Leria (DC.) Burkart, Darwiniana 6: 560. 1944]. TYPE: Leria nutans (L.) DC., based on Tussilago nutans L. [= Chaptalia nutans (L.) Pol.].

[FIGURES 11-14 OMITTED]

Lieberkuhna Cass., Dict. sci. nat. 26: 286. 1823. [= Chaptalia sect. Lieberkuhna (Cass.) Burkart, Darwiniana 6: 539. 1944]. TYPE: Lieberkuhna bracteata Cass., nom.illeg., based on Perdicium piloselloides Vahl [= Chaptalia piloselloides (Vahl) Baker].

Loxodon Cass., Dict. sci. nat. 27: 253. 1823. [= Chaptalia sect. Loxodon (Cass.) Burkart, Darwiniana 6: 534. 1944]. TYPE: Loxodon brevipes Cass., nora. illeg., based on Tussilago exscapa Pers. [= Chaptalia exscapa (Pers.) Baker].

Oxydon Less., Linnaea 5: 357. 1830. TYPE: Oxydon bicolor Willd. ex Less., nom. illeg., based on Chaptalia runcinata Kunth.

Thyrsanthema Neck. ex Kuntze, Revis. gen. pl. 1: 369. 1891, nom. rejic. (see obs. of Nesom, 1995: 154).

Etymology. In honor of the French chemist Jean Antoine Chaptal (1756-1832).

Herbs perennial, acaulescent, scapose. Leaves rosulate; sessile to pseudopetiolate; blades entire to lyrate, lanceolate, elliptical, obovate to spathulate, pinnately veined, margin entire to runcinate, glabrous to glabrescent above, densely pubescent beneath. Capitulescences monocephalous, on scapes; capitula heterogamous, disciform or radiate, chasmogamous or cleistogamous, erect or nodding; receptacle epaleate; involucre multiseriate. Florets trimorphic; marginal florets female, without staminodes, corolla true ray to bilabiate with minute inner lips, outer lip expanded or not; intermediate florets female, staminodes absent, corolla filiform irregularly tubular, true ray to bilabiate, shorter than the style; central florets bisexual, corolla bilabiate to tubular-bilabiate; anther apical appendages acute, tails smooth to papillose; style bifid, branches dorsally papillose. Cypselae rostrate to long-attenuate at the apex, glabrous to pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Chaptalia (86 of Figs. 86-91) (Parra & Marticorena, 1972; Salgado-Labouriau, 1982). Pollen subprolate to prolate, medium to large size, tricolporate, exine Mutisia type, microechinate.

Habitat and Distribution. Chaptalia has currently ca. 70 species distributed in America, from southern United States to Central Argentina, although the number of species could be reduced to 20 35 (see Observation).

Species list

1. Chaptalia albicans (Sw.) Vent. ex Steud, Nomencl. bot. 1: 344. 1840. Basionym: Tussilago albicans Sw. TYPE: Jamaica? s.d., P. Browne s.n. (lectotype, designated by Nesom, 1984: 398, LINN 953.16, microfiche 537 not seen, photo at LP!). Additional specimen examined: Bahamas. New Providence: near Nassau, Curtis 181 (MO).

2. Chaptalia angustata Urb., Symb. antill. 7: 432. 1912. TYPE: Dominican Republic. Santo Domingo: Constanza, H. von Tiirckheim 2908 (syntype, US!).

3. Chaptalia anisobasis S. F. Blake, J. Wash. Acad. Sci. 25: 324. 1935. TYPE: Colombia. Dept. Magdalena: Paramos of the Sierra Nevada de Santa Marta, about 30 mi. inland from Dibulla, W. Seifriz 484 (holotype, US!).

4. Chaptalia arechavaletae Hieron. ex Arechav., Anales Mus. Nac. Montevideo 2: 14. 1904. TYPE: Uruguay: s.d., J. Arechavaleta s.n. (holotype, B not seen, B photo F n. 15998 at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: Belgrano bajo, A. Burkart 1696 (SI).

5. Chaptalia azuensis Urb. & Ekman, Ark. Bot. 23: 95. 1930. TYPE: Dominican Republic. Prov. de Azua: Cordillera Central, Loma Nalga de Maco, E. L. Ekman H 6305 (isotype, US!).

6. Chaptalia callacallensis Cuatrec., Fontqueria 9: 5. 1985. TYPE: Peru. Dept. Amazonas: Prov. Chachapoyas, cerros Calla Calla, 18 km above Leimebamba, on road to Balsas, P. C. Hutchinson & J. K. Wright 5589 (type, US!).

7. Chaptalia chapadensis D. J. N. Hind, Kew Bull. 54: 993. 2000. TYPE: Brazil. State Bahia: Mun. Palmeiras, Pai Inacio, BR242W of Lencois at km 232, Mori & Boom 14388 (holotype, K not seen, K photo at LP!).

8. Chaptalia cipoensis Roque, Kew Bull. 60: 133. 2005. TYPE: Brazil. State Minas Gerais: Mun. Santana do Riacho, Serra do Cipo, Rodovia Belo Horizonte-Conceigao do Mato Dentro, km 132, Semir & Sazima CESC 4758 (holotype, UEC not seen).

9. Chaptalia comptonioides Britton & P. Wilson, Mem. Torrey Bot. Club 16: 117. 1920. TYPE: Cuba. Prov. Oriente: Ensenada Mora, Britton et al. 12937 (isotype, MO!).

10. Chaptalia cordata Hieron., Bot. Jahrb. Syst. 21: 370. 1896. TYPE: Ecuador. Prov. Carchi, Paramo del Angel, near El Voladero, G. Harling & Anderson 12104 (neotype, designated by Harling, 1991: GB not seen, photo in FI. Ecuador n. 42: 49, Fig. 7. 1991). Additional specimen examined: Peru. Dept. Amazonas: Prov. Bongara, along Yambrasbamba-Pomacocha trail, between Yanayacu and Pomacocha, J. J. Wurdack 1062 (LP).

11. Chaptalia cordifolia (Baker) Cabrera, FI. llustr. Catarinense 1: 67. 1973. Basionym: Trichocline cord(folia Baker. TYPE: Brazil. Rio de Janeiro, A. Glaziou 18320 (lectotype, designated by Cabrera 1973: 67, K not seen). Additional specimen examined: Brazil. State Santa Catarina: Campo dos Padres, Born Retiro, P. Reitz 2531 (LP).

12. Chaptalia crassiuscula Urb., Repert. Spec. Nov. Regni Veg. 21: 228. 1925. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, E. L. Ekman 9127 (holotype, S not seen, S photo at LP!).

13. Chaptalia crispata Urb. & Ekman, Ark. Bot. 23: 97. 1930. TYPE: Haiti. Massif de la Selle: Marigot, Rivi6re Chotard, E. L. Ekman H 8004a (type, probably in HAC).

14. Chaptalia dentata (L.) Cass., Dict. sci. nat. 26: 104. 1823. Basionym: Tussilago dentata L. TYPE: Lectotype, designated by Nesom, 1984: 397, the illustration in PI. Amer. 2: tab. 40, Fig. 77. 1755. Additional specimen examined: Bahamas. Andros Island: Conch Sound, J. Northrop & A. Northrop 400 (GH).

15. Chaptalia denticellata Urb. & Ekman, Ark. Bot. 23: 98. 1930. TYPE: Haiti. Massif de la Selle: Port au Prince, near Bassin-Laval, E. L. Ekman H 5493 (isotype, US!).

16. Chaptalia denticulata (Baker) Zardini, Darwiniana 19: 728. 1975. Basionym: Trichocline denticulata Baker. TYPE: Brazil. State Rio de Janeiro: ad fontem fluminis Soberbo, Glaziou 6031 (holotype, B not seen, B photo F n. 16015 at LP!). Additional specimen examined: Brazil. State Bahia: Pico das Almas, Rio de Contas, SE de Campo do Queiroz, R. M. Harley et al. 26584 (US).

17. Chaptalia dolichopoda Urb. & Ekman, Ark. Bot. 23: 99. 1930. TYPE: Haiti. Massif de la Selle: Croix-des-Bousquets, Badeau, inter Morne M6rillon and Morne Badeau, E. L. Ekman H 7796a (isotype, GH!).

18. Chaptalia eggersii Urb., Symb. antill. 3: 418. 1903. TYPE: Dominican Republic. Santo Domingo: Valle Nuevo, Eggers 2220 (isotype, SI!).

19. Chaptalia ekmanii Urb., Repert. Spec. Nov. Regni Veg. 21: 227. 1925. TYPE: Cuba. Prov. Pinar del Rio: Pinar de Cajalbana, E. L. Ekman 17340 (holotype, S not seen, photo LP!).

20. Chaptalia estribensis G. L. Nesom, Phytologia 78: 160. 1995. TYPE: Mexico. State Hidalgo: Municipio Tenango de Doria, 8 to 11 km SW of Tenango de Doria, D. E. Breedlove & Almeda 59571 (holotype, CAS not seen). Additional specimen examined: Mexico. State Hidalgo: El Cirio, al E de Tenango de Doria, Municipio Tenango de Doria, Torres C. & Garcia 823 7 (MEXU).

21. Chaptalia exscapa (Pers.) Baker, F1. bras. 6: 379. 1884. Basionym: Tussilago exscapa Pers. TYPE: Uruguay. Montevideo, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Avellaneda, Wilde, A. Burkart 2191 (SI).

22. Chaptalia flavicans Urb. & Ekman, Ark. Bot. 23: 101. 1930. TYPE: Haiti. Massif de la Selle: Morne des Commissaires ad Grand Gosier, E. L. Ekman H 6903 (holotype, US!).

23. Chaptalia graminifolia (Dusen) Cabrera, F1. Ilustr. Catarinense 1: 60. 1973. Basionym: Chaptalia piloselloides (Vahl) Baker var. graminifolia Dusen. TYPE: Brazil. State Parana: entre Capao Grande y Villa Velha, P. Dusen 4062 (syntype, B not seen, B photo F n. 16003 at LP!). Additional specimen examined: Brazil. State Minas Gerais: Serra do Caparao, Pico Luez Inacio, Brade 16890 (LP).

24. Chaptalia hermogenis M. D. Moraes, Novon 8: 173. 1998. TYPE: Brazil. State Sao Paulo: Eldorado, Nucleo Caverna do Diablo, Parque Estadual de Jacupiranga, M. D. Moraes & F. Arzolla 367 (isotype, LP!).

25. Chaptalia hidalgoensis L. Cabrera & G. L. Nesom, Sida 20: 1363. 2003. TYPE: Mexico. State Hidalgo: Mun. Zimapan, Verdosas, V. M. Huerta 1352 (holotype, IEB not seen).

26. Chaptalia hololeuca Greene, Leafl. Bot. Observ. Crit. 1: 192. 1906. TYPE: Mexico. State Coahuila: and vicinity, Saltillo, E. Palmer 192 (holotype, US!). Additional specimen examined: Mexico. State Coahuila: Saltillo, Palmer 528 (GH).

27. Chaptalia ignota Burkart, Physis (Buenos Aires) 11: 102. 1932. TYPE: Argentina. Prov. Buenos Aires: San Isidro, A. Burkart 2153 (isotype, LP!).

28. Chaptalia incana Cuatrec., Proc. Biol. Soc. Wash. 74: 24. 1961. TYPE: Colombia. Dept. Magdalena: Sierra Nevada de Santa Marta, Hoya del Rio Donachui, Meollaca (or Meuyaca), J. Cuatrecasas & R. Romero Castaneda 24438 (holotype, US!).

29. Chaptalia integerrima (Vell.) Burkart, Darwiniana 6: 576. 1944. Basionym: Tussilago integerrima Vell. TYPE: Lectotype, selected by Burkart, 1944: 576, the illustration at F1. flumin icon. 8: tab. 140. 1831 (1827). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Bolivar, Pirovano, A. Burkart 7064 (SI).

30. Chaptalia isernina Cuatrec., Anales Ci. Univ. Madrid 4: 212. 1935. TYPE: Peru. Prov. Chanchamayo: Chanchamayo, J. Isern 42 (holotype, F!).

31. Chaptalia latipes Urb. & Ekman, Ark. Bot. 23: 99. 1930. TYPE: Haiti. Massif du Nord: Gros-Morne, Belanse, E. L. Ekman H 4921 (holotype, S not seen, S photo at LP!).

32. Chaptalia leiocarpa (DC.) Urb., Symb. antill. 8: 747. 1921. Basionym: Leria leiocarpa DC. TYPE: Cuba. s.d., R. de la Sagra s.n. (isotype, F!).

33. Chaptalia leptophylla Urb., Repert. Spec. Nov. Regni Veg. 21: 229. 1925. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, Rio Jimbambay, E. L. Ekman 9559 (isotype, SI!).

34. Chaptalia lyratifolia Burkart, Darwiniana 6: 527. 1944. TYPE: Mexico. State Nuevo Leon: Sierra Madre, above Monterrey, C. G. Pringle 10217 (isotype, SI!).

35. Chaptalia madrensis G. L. Nesom, Phytologia 78: 158. 1995. TYPE: Mexico. Nuevo Leon: ca. 35 km ENE of Dr. Arroyo, NNW of Cerro Pena Nevada, G. Nesom 4758A (holotype, US!).

36. Chaptalia malcabalensis Cuatrec., Ciencia (Mexico) 24: 123. 1965. TYPE: Peru. Dept. Amazonas: Prov. Chachapoyas, cerro Malcabal (Cerro Tumbe), 3 6 km SW of Molino-pampa, J. J. Wurdack 141 (holotype, US!).

37. Chaptalia mandonii Sch. Bip. ex Burkart, Darwiniana 6: 551. 1944. TYPE: Bolivia. Larecaja, en la vecindad de Sorata, monte Chilica, Mandon 11 (isotype, GOET not seen). Additional specimen examined: Argentina. Prov. Catamarca: Dept. Andalgalfi, Cuesta de Mina Capillitas, A. L. Cabrera et al. 24738 (LP).

38. Chaptalia media (Griseb.) Urb., Symb. antill. 3: 419. 1903. Basionym: Leria media Griseb. TYPE: Cuba. Cuba oriental C. Wright 332 (isotype, MO!).

39. Chaptalia membranacea Urb., Symb. antill. 3: 418. 1903. TYPE: Dominican Republic. Santo Domingo: Puerto Plata, Loma Isabel, Eggers 1582 (isotype, SI!).

40. Chaptalia meridensis S. F. Blake, Contr. U. S. Natl. Herb. 20: 540. 1924. TYPE: Venezuela. Merida: Paramo del Molino, A. Jahn 954 (holotype, US!).

41. Chaptalia modesta Burkart, Physis (Buenos Aires) 11: 100. 1932. TYPE: Argentina. Prov. Salta: Pampa Grande, Jan 1897, C. L. Spegazzini s.n. (holotype, LP!).

42. Chaptalia montana Britton, Bull. Torrey Bot. Club 50: 51. 1923. TYPE: Cuba. Prov. Oriente: Sierra Maestra, Fr. Leon 10802 (type, probably in HAC).

43. Chaptalia mornicola Urb. & Ekman, Ark. Bot. 23: 97. 1930. TYPE: Haiti. Massif de la Selle: Nouvelle Touraine, Chapelle Faure, E. L. Ekman H 1413 (holotype, S not seen, S photo at LP!).

44. Chaptalia nipensis Urb., Repert. Spec. Nov. Regni Veg. 21: 229. 1925. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, E. L. Ekman 3094 (holotype, S not seen, S photo at LP!).

45. Chaptalia nutans (L.) Pol., Linnaea 41: 582. 1877. Basionym: Tussilago nutans L. TYPE: America: s.d., s. leg. (lectotype, designated by Nesom, 1995: 162, LINN n. 995.5 not seen, photo at LP!). Additional specimen examined: Brazil. State Silo Paulo: Campos de Butanta, Joly 145AB3 (LP*).

46. Chaptalia oblonga D. Don, Trans. Linn. Soc. London 16: 240. 1830. TYPE: Peru. s.d., H. Ruiz & J. A. Pavon s.n. (holotype, MA not seen, MA photo at LP!).

47. Chaptalia obovata C. Wright, Anales Acad. Ci. Med. Habana 6: 212. 1869. TYPE: Cuba. s.d., C. Wright 3617 (isotype, US!).

48. Chaptalia ovalis D. Don, Trans. Linn. Soc. London 16: 241. 1830. TYPE: Peru. s.d., H. Pavon s.n. (holotype, MA not seen, MA photo at LP!).

49. Chaptalia paramensis Cuatrec., Proc. Biol. Soc. Wash. 74: 25. 1961. TYPE: Colombia. Dept. Magdalena: Sierra de Perija, between Cerro Venado and Cerro Avion, J. Cuatrecasas & R. Romero Castaneda 25117 (holotype, US!).

50. Chaptalia piloselloides (Vahl) Baker, F1. bras. 6: 378. 1884. Basionym: Perdicium piloselloides Vahl. TYPE: Uruguay. Montevideo, s.d., Commerson s.n. (type, not localized). Additional specimen examined: Argentina. Prov. Buenos Aires: Partido Avellaneda, Sarandi, A. Burkart 2208 (SI).

51. Chaptalia primulacea Greene, Leafl. Bot. Observ. Crit. 1: 195. 1906. TYPE: Dominican Republic. Santo Domingo, C. Wright et al. 261 (isotype, US!).

52. Chaptalia pringlei Greene, Leafl. Bot. Observ. Crit. 1: 192. 1906. TYPE: Mexico. State Oaxaca: above Dominguillo, C. G. Pringle 5796 (lectotype, designated by Nesom, 1995: 161, US!). Additional specimen examined: Mexico. State Oaxaca: ca. 4 km W of Tamazulapan on road to Chilapa de Diaz, G. Nesom 4405 (MO).

53. Chaptaliapumila (Sw.) Urb., Symb. antill. 3: 420. 1903. Basionym: Tussilago pumila Sw. TYPE: Jamaica. s.d., A. Swartz s.n. (holotype, M!).

54. Chaptalia rocana Britton & P. Wilson, Mere. Torrey Bot. Club 16: 118. 1920. TYPE: Cuba. Mtns. of southern Santa Clara, Rio Caracusey, Banao mtns., Leon & Roca 7904 (type, probably in HAC).

55. Chaptalia rotundifolia D. Don, Trans. Linn. Soc. London 16: 242. 1830. TYPE: Peru. Pillao, s.d., H. Ruiz & J. A. Pavon s.n. (holotype, MA not seen).

56. Chaptalia runcinata Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 5. 1818. TYPE: Colombia: F. Humboldt & A. Bonpland 2031 (lectotype, designated by Nesom, 1995: 168, P not seen, P photo F n. 38167 at LP!). Additional specimen examined: Argentina. Prov. Buenos Aims: Partido La Plata, alrededores de La Plata, Elizalde, A. L. Cabrera 907 (SI).

57. Chaptalia shaferi Britton & P. Wilson, Mem. Torrey Bot. Club 16:118. 1920. TYPE: Cuba. Prov. Oriente: valley of Rio Yamaniguey, J. A. Shafer 4203 (holotype, GH!).

58. Chaptalia similis R. E. Fr., Nova Acta Regiae Soc. Sci. Upsal. 1: 95. 1905. TYPE: Argentina. Prov. Jujuy: Santa Catalina, Kurtz 11460 (syntype, BAF not seen). Additional specimen examined: Argentina. Prov. Catamarca: Dept. Andalgala, cuesta de Mina Capillitas, A. L. Cabrera et al. 24738 (LP).

59. Chaptalia sinuata (Less.) Baker, F1. bras. 6: 378. 1884. Basionym: Leria nutans (L.) DC. var. sinuata Less. TYPE: Uruguay. Montevideo, Sellow 1413, 2212 (syntypes, B not seen, B photo F n. 16006 at LP!). Additional specimen examined: Argentina. Prov. Misiones: Dept. Candelaria, entre Candelaria y Santa Ana, 20 Nov 1896, N. Alboffs.n. (LP).

60. Chaptalia spathulata (D. Don) Hemsl., Biol. Cent.-Amer., Bot. 2: 255. 1881-1882. Basionym: Leria spathulata D. Don. TYPE: Mexico. State Guerrero: Mazatlan, M. Sesse & J. M. Mocino 2028 (holotype, MA not seen).

61. Chaptalia stenocephala (Griseb.) Urb., Symb. antill. 3: 420. 1903. Basionym: Leria stenocephala Griseb. TYPE: Cuba. s.d., C. Wright 2874 (isotype, MO!).

62. Chaptalia stuebelii Hieron., Bot. Jahrb. Syst. 21: 371. 1896. TYPE: Ecuador. Prov. Pichincha: Cerro Pichincha, Holmgren 297 (neotype, designated by Harling, 1991: 53, S not seen). Additional specimen examined: Ecuador. Prov. Azuay: valley of the Rio Paute, between Paute and Cuenca, Parroquia Jose Victor Izquierdo at "Pupazche," Camp 2573 (US).

63. Chaptalia texana Greene, Leafl. Bot. Observ. Crit. 1: 191. 1906. TYPE: United States. State Texas: W Texas, G. C. Neally 297 (holotype, US!). Additional specimen examined: Mexico. State Baja California: La Champagna, Sierra de Las Palmas, S of Sta. Rosalia, Gentry & Fox 11768 (MEXU).

64. Chaptalia tomentosa Vent., Descr. pl. nouv. 7: tab. 61. 1802. TYPE: Walter, Carol. Bosc (holotype G-DC, microfiche 1238, frame 15!). Additional specimen examined: United States. State Alabama: Spring Hill, Graves 1950 (MO).

65. Chaptalia transiliens G. L. Nesom, Rhodora 86: 127. 1984. TYPE: Mexico. Nuevo Leon: ca. 35 km ENE of Dr. Arroyo, NNW of Cerro Pena Nevada, G. Nesom 4759 (holotype, US!).

66. Chaptalia turquinensis Borhidi & O. Muniz, Acta Bot. Acad. Sci. Hung. 17: 35. 1971. TYPE: Cuba. Prov. Oriente: Sierra Maestra, Pico Turquino, Paso de los Angustios, V. Samek 815 (type, probably in HAC).

67. Chaptalia undulata Urb. & Ekman, Ark. Bot. 23: 100. 1930. TYPE: Haiti. Massif du Nord: St. Louis du Nord, Morne Baron, s. leg. H 4696 (type, probably in HAC).

68. Chaptalia vegaensis Urb. & Ekman, Ark. Bot. 23: 96. 1930. TYPE: Dominican Republic. Santo Domingo: Prov. La Vega, Bonao, Loma Peguera, E. L. Ekman H 11480 (holotype, S not seen, S photo at LP!).

Observation. A revision in progress by L. Katinas shows that many of the species under the name Chaptalia (e.g., C. anisobasis, C. estribensis, C. meridensis) deviate from the generic concept mainly by the presence of staminodes. Thus, they should be excluded from the genus and moved to Gerbera. Other species, such as many of those of the Antilles (e.g., C. crispata, C. dolichopoda, C. membranacea), are variants of one morphologically heterogeneous species and their names should be treated as synonyms. As a result, the probable number of species of the genus will be 20-35 (Katinas, 2004a).

Literature. Burkart (1944), Nesom (1995).

Chucoa Cabrera

Bol. Soc. Argent. Bot. 6: 40. 1955. TYPE: Chucoa ilicifolia Cabrera. 12 of Figs. 11-14. Weberbaueriella Ferreyra, Bol. Soc. Peruana Bot. 5: 2. 1955. TYPE: Weberbaueriella johnstoniana Ferreyra [= Chucoa ilicifolia Cabrera].

Etymology. From the type locality in the province of Santiago Chuco, Peru.

Shrubs. Leaves alternate; sessile to subsessile; blades lanceolate to obovate, pinnately veined, margin denticulate to spinose, lanuginous to glabrous above, lanose beneath. Capitulescences monocephalous or two- to three-headed, terminal; capimla pedunculate, homogamous or occasionally heterogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, rarely the marginal ones female with staminodes, corolla actinomorphic, tubular-funnelform to subcampanulate, shallowly to deeply five-lobed; anther apical appendages acute, tails smooth to papillose, filaments smooth to papillose at the base; style bifid to bilobed, branches dorsally papillose. Cypselae glabrous to subglabrous; pappus of scabrid bristles, all capillary and equal in length or the innermost longer and somewhat flattened at the base.

Pollen Description. Chucoa ilicifolia (87 of Figs. 86-91). Pollen subprolate to prolate, elliptic in equatorial view, subtriangular in polar view, large size, P x E = (70-85 x 72-69) [micro]m, tricolporate, colpi long, with margins and psilate membrane, mesoapermrate. Exine Mutisia type, microechinate, 10-13.5 [micro]m thick at the equator, slender at the poles, ectosexine compact. Ratio ectosexine/endosexine: 1: 1. Nexine 2-2.5 [micro]m thick.

Habitat and Distribution. Genus of two species endemic to rocky dry slopes of northwestern Peru.

Species list

1. Chucoa iliciJblia Cabrera, Bol. Soc. Argent. Bot. 6: 42. 1955. TYPE: Peru. Dept. La Libertad: Prov. Santiago Chuco, Angasmarca-Tulpo, A. Lopez Miranda 1090 (holotype, LP!*).

2. Chucoa lanceolata (H. Beltran & Ferreyra) G. Sancho, S. E. Freire & Katinas, Taxon 54: 86. 2005. Basionym: Gochnatia lanceolata H. Beltran & Ferreyra. TYPE: Peru. Dept. Arequipa: Prov. Castilla, Andagua, Carolina 1 (holotype, USM not seen; isotype, US!).

Observation. There are some problems in trying to sort out priority between the names Chucoa and Weberbaueriella since both names were published in the same year, 1955, and date of priority has not been conclusively established. After first published, Weberbaueriella has not appeared in any other modern literature and Chucoa is the name most commonly used for the taxon. If Weberbauerella Ulbrich, a genus of Fabaceae established in 1906, and Weberbaueriella Ferreyra were ruled out to be treated as homonyms, it would establish Chucoa as unquestionably the correct name for the Asteraceae genus (Brummitt, 2005). The current recommendation of the General Committee, following the opinion of the Committee for Spermatophyta, is that Weberbauerella (Fabaceae) and Weberbaueriella (Asteraceae) are treated as confusable names (Barrie, 2006).

Literature. Ferreyra (1995), Sancho et al. (2005).

Cnicothamnus Griseb.

Abh. Konigl. Ges. Wiss. Gottingen 19: 196. 1874. TYPE: Cnicothamnus lorentzii

Griseb. 13 of Figs. 11-14. Lefrovia Franch., J. Bot. (Morot) 2: 377. 1888. TYPE: Lefrovia rhaponticoides Franch. [= Cnicothamnus lorentzii Griseb.].

Etymology. From the Greek knecos, yellow, yellowish, a name applied to Carthamus tinctorius with deep orange flowers, and thamnos, shrub, by the resemblance of the flower color of both species.

Shrubs to small trees. Leaves alternate; short-petiolate; blades wide-elliptic to ovate, pinnately veined, margin denticulate to dentate, glabrous above and floccosepubescent beneath. Capitulescences monocephalous, terminal; capitula sessile to short-pedunculate, homogamous, radiate; receptacle epaleate; involucre multiseriate, six- to ten-seriate, phyllaries acute or with showy, orbicular to spatulate, laciniate apical appendages. Florets dimorphic, bisexual; marginal florets with corolla subbilabiate (3+1 corolla lips), outer lip expanded; central florets with corolla tubular-funnelform, deeply five-lobed; anther apical appendages apiculate, tails smooth; style bilobed, branches dorsally smooth. Cypselae densely villose; pappus of scabrid bristles, half capillary and short and half somewhat paleaceous, clavate at the apex and longer.

Pollen Description. Cnicothamnus lorentzii (88 of Figs. 86-91). Pollen subprolate to prolate, elliptic in equatorial view, circular in polar view, large size, P x E - (66-76x48-54) [micro]m. Tricolporate, colpi long, with margin and scabrate membrane, mesoaperture diffuse? Exine Mutisia type, echinate, 8-11 [micro]m thick, spines of 2-2.5 [micro]m length. Ratio ectosexine/endosexine: 1.5:1; 1:1. Nexine ca. 2 [micro]m thick. Note: pollen similar to that of type II of Gochnatia.

Habitat and Distribution. Genus of two species from the subtropical forests of Bolivia and northwestern Argentina.

Species list

1. Cnicothamnus azafran (Cabrera) Cabrera, Notas Mus. La Plata, Bot. 9: 256. 1944. Basionym: Cnicothamnus lorentzii Griseb. var. azafran Cabrera. TYPE: Argentina. Prov. Salta: Rio Maiz Gordo, A. Ragonese 334 (holotype, LP!).

2. Cnicothamnus lorentzii Griseb., Abh. Konigl. Ges. Wiss. Gottingen 19: 197. 1874. TYPE: Argentina. Prov. Tucuman, P. G. Lorentz 182 (isotype, CORD not seen). Additional specimens examined: Argentina. Prov. Jujuy: Ledesma, Sierra de Calilegua, S. Venturi 5310 (LP); Prov. Salta, 50 km SE from Salta city, S. Maldonado 408 (LP*).

Literature. Cabrera (1978), Freire (1996), Freire et al. (2002).

Cyclolepis D. Don

Philos. Mag. Ann. Chem. 11: 392. 1832. TYPE: Cyclolepis genistoides D. Don. 14 of Figs. 11-14.

Cyclopis Gillies, Arch. Bot. (Paris) 2: 468. 1833, an ortographic variant of the name Cyclolepis.

Etymology. From the Greek kyclos, circle, ring, and lepis, scales, by the imbricate disposition of the phyllaries.

Shrubs branched, gynodioecious, short branches modified in thorns. Leaves alternate, caducous; short-petiolate; blades elliptic, pinnately veined, margin entire, pubescent on both surfaces. Capitulescences racemose to spicate; capitula subsessile, homogamous, discoid; receptacle epaleate; involucre three- to six-seriate. Florets isomorphic in bisexual capitula, corolla actinomorphic, tubular-funnelform, deeply five-lobed, lobes coiled; anther apical appendages apiculate, tails slightly papillose; style bifid, branches dorsally smooth; female capitula with florets without staminodes, corolla tubular-filiform, shallowly five-lobed, tube long, up to 2/3 of corolla length, lobes straight. Cypselae villose; pappus of scabrid bristles, half capillary and short and half somewhat paleaceous, plumose at the apex and longer.

Pollen Description (Telleria & Forcone, 2002) (89 of Figs. 86-91). Pollen spheroidal to subprolate, elliptic to spheroidal in equatorial view, circular in polar view, medium to large size, tricolporate, exine Mutisia type, microechinate.

Habitat and Distribution. Monotypic genus that grows in salty soils from Paraguay to northern Patagonia in Argentina.

Species list

1. Cyclolepis genistoides D. Don, Philos. Mag. Ann. Chem. 11 : 392. 1832. TYPE: Sine loc., s. d., J. Gillies s.n. (holotype, probably K not seen). Additional specimens examined: Argentina. Prov. Salta: Quebrada de las Conchas, A. L. Cabrera 3782 (LP); Prov. Chubut: Trelew, A. Forcone 459 (CORD*).

Literature. Cabrera (1951, 1971b), Freire et al. (2002).

Dinoseris Griseb.

Symb. ft. argent.: 213. 1879. TYPE: Dinoseris salicifolia Griseb. 15 of Figs. 15-18.

Etymology. From the Greek deinos, terrible, powerful, mighty, and seris, chicory, and lettuce.

Shrubs to little trees. Leaves opposite; petiolate; blades elliptic to lanceolate, pinnately veined, margin entire to serrulate, glabrous. Capitulescences monocephalous or two- to three-headed, terminal; capitula pedunculate, homogamous, ligulate; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla ligulate, limb shallowly five-dentate; anther apical appendages acute, tails papillose; style bifid, branches dorsally papillose. Cypselae truncate to slightly attenuate at the apex, glabrous; pappus of scabrid, capillary bristles.

[FIGURES 15-18 OMITTED]

Pollen Description (Telleria & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate.

Habitat and Distribution. Monotypic genus that grows on the slopes in the subtropical forests of Bolivia and northern Argentina.

Species list

1. Dinoseris salicifolia Griseb., Symb. ft. argent.: 214. 1879. TYPE: Argentina. Prov. Tucuman: Sierra de Tucuman, Suciliaco, P G. Lorentz & G. H. E. W. Hieronymus 862 (isosyntype, CORD not seen). Additional specimen examined: Argentina. Prov. Salta: Campo Quijano, July 1954, Corte s.n. (LP).

Observation. Some authors (e.g., Ariza Espinar, 1973; Bremer, 1994) consider the only species of Dinoseris (D. salicifolia) a species of Hyaloseris. However, other authors (e.g., Cabrera, 1978; Katinas, 1996a) prefer to maintain both genera as separate. Some characters that distinguish both taxa are: brachyblasts and three- to six-flowered capitula in Hyaloseris versus lack of brachyblasts and capitula with more than 20 florets in Dinoseris.

Literature. Ariza Espinar (1973), Cabrera (1978), Katinas (1996a).

Duidaea S. F. Blake

Bull. Torrey Bot. Club 58: 496. 1931. TYPE: Duidaea tatei S. F. Blake. 16 of Figs. 15-18.

Etymology. Named for Mount Duida (state Amazonas, Venezuela), which towers above Rio Orinoco at La Esmeralda.

Shrubs. Leaves alternate, clustered at the stem apex; sessile to subsessile, the petioles, when present, slightly clasping; blades linear to oblanceolate, subcoriaceous to coriaceous, one-veined, margin entire, revolute, glabrous to densely pubescent beneath. Capitulescences monocephalous, rarely several heads per branch, either axillar or seemingly terminal; capitula short- to longpedunculate, pendulous, homogamous, bilabiate; receptacle epaleate; involucre multiseriate, phyllaries reddish. Florets isomorphic, bisexual, corolla bilabiate; anthers commonly light purple, generally exserted, apical appendages acuminate, tails papillose; style deep purple, bifid, branches dorsally smooth. Cypselae glabrous to densely pubescent; pappus of scabrid bristles, the outermost shorter.

Pollen Description. Duidaea marahuacensis (90 of Figs. 86-91). Pollen spheroidalsubprolate, spheroidal or elliptic in equatorial view, circular in polar view, large size, P x E = (62-72 x 52-64) [micro]m. Tricolporate, colpi long, with margin and scabrate membrane. Exine Gongylolepis type, 11-14 [micro]m thick in the whole grain, spines 2.5 [micro]m length. Ratio ectosexine/endosexine among spines: 1:3; 1:4. SEM: tectum very perforate. Note: nexine scarcely distinguishable. Pollen similar to that of Glossarion rhodanthum and Quelchia bracteata (see also Carlquist, 1957b; Telleria, 2008).

Habitat and Distribution. Duidaea is endemic to two isolated tepuis in Amazonas, Venezuela and contains four species.

Species list

1. Duidaea marahuacensis Steyerm., Brittonia 30: 50. 1978. TYPE: Additional specimens examined: Venezuela. Terr. Fed. Amazonas: Cerro Marahuaca, S. S. Tillet et al. 752.348 (holotype, VEN!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dtpo. Atabapo, Cerro Marahuaca, R. Liesner 24644 (MO, US*).

2. Duidaea pinifolia S. F. Blake, Bull. Torrey Bot. Club 58: 499. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Summit of Mt. Duida, G. H. H. Tate 530 (isotype, US!).

3. Duidaea rubriceps S. F. Blake, Bull. Torrey Bot. Club 58: 500. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Mt. Duida: Brocchinia Hills, G. H. H. Tate 589 (isotype, US!).

4. Duidaea tatei S. F. Blake. Bull, Torrey Bot. Club. 58: 498. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Mr. Duida: Gorge of Carlo Negro, Savanna Hills, G. H. H. Tate 814 (isotype, US!).

Literature. Maguire et al. (1957a), Aristeguieta (1964), Pruski (1997).

Eriachaenium Sch. Bip.

Flora 38: 120. 1855. TYPE: Eriachaenium magellanicum Sch. Bip. 17 of Figs. 15-18.

Etymology. From the Greek erion, wool, and from the Latin achaenium, a type of fruit, by the villose fruits.

Herbs perennial, dwarf, with stout, oblique to vertical rhizomes, stems prostrate to ascending. Leaves alternate; sessile, clasping; blades oblanceolate, pinnately veined, margin entire to undulate-dentate, glabrous to subglabrous above, tomentose beneath. Capitulescences monocephalous, axillar; capitula pedunculate, heterogamous, disciform; receptacle epaleate; involucre uniseriate. Florets dimorphic; marginal florets female with staminodes, corolla tubular-funnelform, deeply fourlobed; central florets bisexual or male with a rudimentary ovary, corolla tubular-funnelform, deeply five-lobed; anther apical appendages rounded to acute, basally constricted and demarcated from the thecae, basally auriculate with tails very short, smooth to slightly papillose; style bilobed, dorsally papillose. Cypselae truncate at the apex, shaggy, dimorphic, the marginal cypselae conspicuously bigger than the central ones; pappus absent.

Pollen Description (91 of Figs. 86-91). Pollen spheroidal to prolate, spheroidal or elliptic in equatorial view, circular in polar view, medium size, P x E = (30-36 x 24-30) [micro]m. Tricolporate, colpi long with thin margin and microgranulate membrane, mesoaperture diffuse. Exine Mutisia type, microechinate, 2-6 [micro]m thick, slightly slender at the poles. Ratio ectosexine/endosexine: 1:1.5; 1:2. Nexine 1.5 [micro]m thick. SEM: tectum punctate. Note: grains with pyramidal or rectangular shape and grains biaperturate were observed. Some grains have the endosexine with compact aspect, as occurs in Adenocaulon chilense (Mutisieae). Pollen similar to that of Adenocaulon and Artemisia verlotorum (Anthemideae).

Habitat and Distribution. Monotypic genus endemic to Patagonia in Argentina and Chile.

Species list

1. Eriachaenium magellanicum Sch. Bip., Flora 38: 121. 1855. TYPE: Chile. Prov. Magallanes: Oazy Harbour, Lechler 1256 (type, probably SGO). Additional specimens examined: Argentina. Prov. Santa Cruz: Jan 1903, C. L. Spegazzini s.n. (LP*); Rio Gallegos, O'Donell 4091 (LP*); id., Estancia Guer Aike, H. Sleumer 908 (LP*).

Literature. Cabrera (1971b), Robinson and Brettell (1973), Moore (1983), Katinas (2000).

Eurydochus Maguire & Wurdack

Bol. Soc. Venez. Ci. Nat. 20: 57. 1958. TYPE: Eurydochus bracteatus Maguire &

Wurdack. 18 of Figs. 15-18.

Etymology. Derived from the Greek eurys, broad, large, and dochos, a receptacle, dechomai, to receive, in reference to the broad receptacles.

Trees or shrubs. Leaves alternate, often clustered apically; petiolate, the petioles clasping; blades elliptic to oblanceolate, chartaceous to sub-coriaceous, pinnately veined, margin entire, glabrous. Capitulescences monocephalous, heads subterminal; capitula pedunculate, homogamous, bilabiate, six- to many-flowered; receptacle epaleate; involucre multiseriate, hemispherical, phyllaries subequal, the outer phyllaries commonly decurrent onto the peduncle. Florets isomorphic, bisexual, corolla bilabiate, outer lip expanded; anthers brown to purple, exserted, apical appendages acute, slightly coiled at the apex, tails papillose, connate; style rose-colored, bifid, branches dorsally smooth. Cypselae puberulent; pappus of fragile, scabrid, capillary bristles, sometimes individually deciduous, the outermost shorter.

Pollen Description. Pollen prolate, elliptic in equatorial view, circular in polar view, large size, P x E = (72-80 x 54-60) [micro]m. Tricolporate, long colpi with microgranulate membrane. Exine Wunderlichia type, 8-10 [micro]m thick in the whole grain; spines 3 [micro]m length. Ratio ectosexine/endosexine among spines: 1:1 ; 1.5:1; 1:1.5. Nexine 1.5 [micro]m thick. Note: pollen grains similar to those of Salcedoa mirabaliarum F. Jimenez Rodr. & Katinas and Wunderlichia azulensis and W. mirabilis Riedel ex Baker (see also Jimenez Rodriguez et al., 2004; Telleria, 2008).

Habitat and Distribution. Eurydochus is a poorly known monotypic genus known only from two collections in the western Guayana Highland. It occurs on Cerro de la Neblina on the border of Venezuela and Brazil and on nearby Serra Pirapucu in Brazil.

Species list

1. Eurydochus bracteatus Maguire & Wurdack, Bol. Soc. Venez. Ci. Nat. 20: 57. 1958. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 42186 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Rio Yatua, Canon Grande, B. Maguire et al. 42238 (US*).

Literature. Maguire and Wurdack (1958), Aristeguieta (1964), Pruski (1991, 1997).

Gerbera L., nom. cons.

Sp. pl.: 885. 1753. TYPE: Arnica gerbera L. [= Gerbera linnaei Cass. (type cons.)]. 19 of Figs. 19-22.

Aphyllocaulon Lag., Amen. nat. Espan: 38. 1811, species name not designated.

Atasites Neck., Elem. bot. 1: 7. 1790, nom. illeg.

Berniera DC., Prodr. 7: 18. 1838, nom. illeg. TYPE: Berniera nepalensis DC.

[= Gerbera maxima (D. Don) Beauverd].

Epiclinastrum Bojer ex DC., Prodr. 7: 18. 1838. TYPE: Epiclinastrum candollei

Bojer ex DC. [= Gerbera piloselloides (L.) Cass.].

Gerberia Cass., Bull. Sci. Soc. Philom. Paris: 34. 1817, an orthographic variant of the name Gerbera.

Lasiopus Cass., Bull. Sci. Soc. Philom. Paris: 152. 1817, non D. Don, 1836.

[= Gerbera sect. Lasiopus (Cass.) Sch. Bip., Flora 27: 780. 1844]. TYPE: Lasiopus ambiguus Cass. [= Gerbera ambigua (Cass.) Sch. Bip.].

Leptica E. Mey. ex DC., Prodr. 7: 16. 1838. TYPE: Leptica ciliata E. Mey ex DC.

[= Gerbera piloselloides (L.) Cass.].

Oreoseris Wall. ex DC., Prodr. 7: 17. 1838. TYPE: Oreoseris lanuginosa Wall. ex

DC. [= Gerbera gossypina (Royle) Beauverd].

Etymology. In honor of the eighteenth century German naturalist Traugott Gerber (d. 1743), physician, traveler, and plant collector.

Herbs perennial, acaulescent, scapose. Leaves rosulate; sessile, petiolate to pseudopetiolate; blades linear-oblanceolate to orbicular, pinnately veined, margin entire to deeply pinnatisect, densely pubescent on both surfaces or only beneath. Capitulescences monocephalous, on scapes; capitula heterogamous, radiate or rarely disciform; receptacle epaleate; involucre multiseriate. Florets dimorphic to trimorphic; marginal florets female generally with filiform to well developed staminodes, corolla bilabiate with an expanded outer lip, rarely minute; intermediate florets (when present) female with or without staminodes, corolla bilabiate, rarely filiformbilabiate, as long as the style or longer; central florets bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bilobed to bifid, dorsally papillose. Cypselae rostrate to long-attenuate at the apex, glabrous to pubescent; pappus of scabrid, capillary bristles.

[FIGURES 19-22 OMITTED]

Pollen Description. Gerbera hieracioides (Kunth) Zardini (92 of Figs. 92-94) (Telleria & Katinas, 2004). Pollen subprolate to spheroidal, large size, tricolporate, exine Mutisia type, slightly microechinate (see also Southworth, 1983; Lin et al., 2005).

Habitat and Distribution. Genus of 29 species distributed in Africa, southern Asia, and America.

Species list

1. Gerbera ambigua (Cass.) Sch. Bip., Flora 27: 780. 1844. Basionym: Lasiopus ambiguus Cass. TYPE: Africa. South Africa: Cape Sonnerat, s.d., s. leg. (holotype, P not seen). Additional specimen examined: Africa. South Africa: Natal, NW Pietermaritzburg, N side of Old Howick road, E of Ling road, T. Mc Farland 128 (WIS).

2. Gerbera aurantiaca Sch. Bip., Flora 27: 781. 1844. TYPE: Africa. South Africa: Natal, Pietennaritzburg, Krauss 388 (holotype, P? not seen; isotype, BM not seen). Additional specimen examined: Africa. South Africa: Natal, Old Howick road, E of Link road, about 4 km NW of Pietermaritzburg business district, T. Mc Farland 265 (WIS).

3. Gerbera bojeri (DC.) Sch. Bip., Flora 27: 780. 1844. Basionym: Lasiopus bojeri DC. TYPE: Africa. Madagascar: s.d., Bojer s.n. (holotype, G-DC not seen). Additional specimen examined: Africa. Madagascar: vicinity of Station Forestiere Antsampandrano, T. Croat 29094 (MO).

4. Gerbera cordata (Thunb.) Less., Linnaea 5: 297. 1830. Basionym: Arnica cordata Thunb. TYPE: Africa. South Africa: Cape province, Capb. Spei, s.d., s. leg. herb. Thunberg 19988 (paratype, UPS not seen). Additional specimen examined: Africa. South Africa: W Cape province, Keurbooms River Forest Res., Willosoman, Dahlstrand 9528 (MO).

5. Gerbera crocea (L.) Kuntze, Revis. gen. pl. 3: 149. 1898. Basionym: Arnica crocea L. TYPE: Sine loc., s.d., s. leg. (holotype, LINN 1001, n. 7 not seen). Additional specimens examined: Africa. South Africa Cape, Hermanus, F. Rogers 26453 (US); Vogelgat, W of base camp, L Williams 3704 (WIS).

6. Gerbera delavayi Franch., J. Bot. (Morot) 2: 68. 1888. TYPE: China. Yunnan: Choui-tsin-yn, J. M. Delavay 1918 (isotype, GH!).

7. Gerbera diversifolia Humbert, Mem. Soc. Linn. Normandie 25: 151, 307. 1923. TYPE: Africa. Madagascar: Tampoketsa d'Analamaitso, haut bassin du Bemarivo, Perrier de la Bathie 3230 (holotype, P not seen, P photo at LP!).

8. Gerbera elliptica Humbert, Mem. Soc. Linn. Normandie 25: 154, 307. 1923. TYPE: Africa. Madagascar: Perrier de la Bathie 3190 (lectotype, designated by Hansen, 1985b: 452, P not seen, P photo at LP!). Additional specimen examined: Africa. Madagascar: Massif de l'Andringitra, Iratsy, vallees de la Riambava et de la Antsifotra, H. Humbert 3654 (GH).

9. Gerbera emirnensis Baker, J. Linn. Soc., Bot. 22: 498. 1887. TYPE: Africa. Madagascar: Central Madagascar, Baron 3457 (lectotype, designated by Hansen, 1985b: 452, K not seen, K photo at LP!).

10. Gerbera galpinii Klatt, Bull. Herb. Boissier 4: 844. 1896. TYPE: Africa. South Africa: Transvaal, Barberton, Umotali valley, E. E. Galpin 1132 (isotype, K not seen, K photo at LP!).

11. Gerbera gossypina (Royle) Beauverd, Bull. Soc. Bot. Geneve 2: 40. 1910. Basionym: Chaptalia gossypina Royle. TYPE: It is found at Mussooree, and similar elevations throughout Gurhual and Simore (type not localized). Additional specimen examined: India. Landour, Mussoorie, April 1929, F. Cleland s.n. (US).

12. Gerbera henryi Dunn, J. Linn. Soc., Bot. 35: 511. 1903. TYPE: China. Yunnan: Mentzge, A. Herny 9111 (isotype, US not seen, US photo at LP!).

13. Gerbera hieracioides (Kunth) Zardini, Bol. Soc. Argent. Bot. 16: 105. 1974. Basionym: Onoseris hieracioides Kunth. TYPE: Ecuador. Prov. Chimborazo: Alausi, Sitzan, s.d., s. leg. (holotype, P not seen). Additional specimen examined: Peru. Dept. Cajamarca: Prov. Contumasa, Ditstrito Guzmango, Herilla, A. Sagastegui 106 (LP*).

14. Gerbera hintonii (Bullock) Katinas, Novon 8: 383. 1998. Basionym: Chaptalia hintoni Bullock. TYPE: Mexico. Mexico: District Temascaltepec, Nanchititla, G. B. Hinton 3098 (isotype, MO!).

15. Gerbera hypochaeridioides Baker, J. Linn. Soc., Bot. 20: 193. 1883. TYPE: Africa. Madagascar: Central Madagascar, Baron 2088 (lectotype, designated by Hansen, 1985b: 452, K not seen, K photo at LP!).

16. Gerberajamesonii Bolus ex Adlam, Gard. Chron. 3: 775. 1888. TYPE: Africa. South Africa: Natal, s.d., Adlam s.n. (lectotype, designated by Hansen, 1985a: 21, SAM not seen). Additional specimens examined: Africa. South Africa: Transvaal, 2 Oct 1925, E. M. Young s.n. (WIS); Hong Kong, Chung Chi College, S. Y. Hu 6385 (US).

17. Gerbera leandrii Humbert, Mem. Inst. Sci. Madagascar, Ser. B, Biol. Veg. 6: 197. 1955. TYPE: Sine loc., H. Humbert 23599 (holotype, P not seen, P photo at LP!).

18. Gerbera linnaei Cass., Dict. sci. nat. 18: 460. 1821, nom. novum. Basionym: Arnica gerbera L. TYPE: Sine loc., s.d., s. leg. (lectotype, designated by Hansen, 1985a: 9, L1NN 1001 not seen). Additional specimen examined: Africa, South Africa: C. Peninsula, Kogelfontein, T. Salter 8532B (MO).

19. Gerbera maxima (D. Don) Beauverd, Bull. Soc. Bot. Geneve 2: 44. 1910. Basionym: Chaptalia maxima D. Don. TYPE: Nepal, Narainhetty, Hamilton & Wallich s.n. (isosyntypes, K not seen).

20. Gerbera nivea (Wall. ex DC.) Sch. Bip., Flora 27: 780. 1844. Basionym: Oreoseris nivea Wall. ex DC. TYPE: Nepal. Gossin-Than, Wallich 3143/253 (lectotype, designated by Hansen, 1988: 66, K not seen). Additional specimen examined: China. Yunnan: between Li-Kang and Joung-Ning, J. Rock 17333 (US).

21. Gerbera parva N. E. Br., Bull. Misc. Inform. Kew: 27. 1895. TYPE: Africa. South Africa: Natal, Drakensberg, near Bushman's river, Evans 57 (lectotype, designated by Hansen, 1985a: 16, K not seen, K photo at LP!).

22. Gerbera perrieri Humbert, Mem. Soc. Linn. Normandie 25: 151, 307. 1923. TYPE: Sine loc., Perrier de la Bathie 3219 (lectotype, designated by Hansen, 1985b: 452, P not seen, P photo at LP!).

23. Gerbera petasitifolia Humbert, Mem. Soc. Linn. Normandie 25: 152, 307. 1923. TYPE: Sine loc., Perrier de la Bathie 2877 (lectotype, designated by Hansen, 1985b: 452, P not seen, P photo at LP!).

24. Gerbera piloselloides (L.) Cass., Dict. sci. nat. 18: 461. 1821. Basionym: Arnica piloselloides L. TYPE: Sine loc., s.d., Burmann s.n., herb. Oldenland (holotype?, G-DC not seen). Additional specimen examined: Africa. Cameroon: Northwest province, near Verkovi village, Mt. Oku, D. Thomas & H. McLeod 6046 (MO).

25. Gerbera raphanifolia Franch., J. Bot. (Morot) 2: 67. 1888. TYPE: China. Yunnan: Mo-che-tsin, J. M. Delavay 1922 (isotype, GH!).

26. Gerbera serrata (Thunb.) Druce, Bot. Exch. Club Brit. Isles Rep.: 624. 1917. Basionym: Arnica serrata Thunb. TYPE: Sine loc., s.d., s. leg. herb. Thunberg 20009 (holotype, UPS not seen). Additional specimen examined: Africa. South Africa: Cape, Knysne Distr., Afgunst Flats, Fourcade 5309 (MO).

27. Gerbera tomentosa DC., Prodr. 7: 16. 1838. TYPE: Cap. Bonae-Spei, s.d., s. leg. (type, not localized). Additional specimens examined: Africa. South Africa: Cape Province, s.d., Zagler s.n. (MO); Kirstenbosch, Newlands, Wemmers Hoek, Dist. Paarl, R. Compton 745 (US).

28. Gerbera viridifolia (DC.) Sch. Bip., Flora 27: 780. 1844. Basionym: Lasiopus viridifolius DC. vat. hirsutus DC. TYPE: Africa. South Africa: Keiskama and Buffelriver, Drege 5110 (lectotype, designated by Hansen, 1985a: 24, G-DC not seen). Additional specimen examined: Africa. South Africa: Natal, Greytown, Aug 1934, J. Wylie s.n. (WIS).

29. Gerbera wrightii Harv., F1. cap. 3: 521. 1865. TYPE: Africa. South Africa: Cape peninsula, near Simonstown, C. Wright 343 (holotype, TCD not seen).

Observation. There are many cultivars of Gerbera, an ornamental genus grown widely for the cut-flower market. An example is Gerbera hybrida, a functional diploid carrying chromosomes or chromosomal fragments from other Gerbera species such as G. jamesonii and G. viridifolia (Teeri et al., 2006).

Literature. Zardini (1974), Hansen (1985a,b, 1988), Katinas (1998).

Glossarion Maguire & Wurdack

Mem. New York Bot. Gard. 9: 390. 1957, emend. Pruski, Brittonia 41: 39. 1989. TYPE: Glossarion rhodanthum Maguire & Wurdack. 20 of Figs. 19 22. Guaicaia Maguire, Mere. New York Bot. Gard. 17: 437. 1967. TYPE: Guaicaia bilabiata Maguire [= Glossarion bilabiatum (Maguire) Pruski].

Etymology. From the Greek diminutive of glossa, tongue, in reference to the ligulate corollas that are occasionally found in the genus, these corollas being "diminutive" bilabiate corollas lacking the inner lips.

Shrubs. Leaves alternate; petiolate, the petioles clasping; blades narrowly lanceolate to elliptic, coriaceous, pinnately veined, margin entire, abaxially buff-tomentose. Capitulescences monocephalous, heads terminal or sometimes axillar; capitula pedunculate, homogamous, isomorphic or dimorphic; receptacle epaleate; involucre, multiseriate. Florets bisexual, isomorphic capitula with all ligulate or all bilabiate corollas; dimorphic capitula with ligulate and bilabiate corollas; ligulate corolla five-dentate; bilabiate corollas, with the inner lip either deeply two- to three-cleft; anthers generally exserted, apical appendages acute, tails slightly papillose, connate with the tails of adjacent stamens; style red, bifid, branches dorsally smooth. Cypselae scarcely glandular or glabrous; pappus of scabrid bristles, somewhat flattened at the apex, the outermost shorter.

Pollen Description. Glossarion rhodanthum. Pollen prolate, elliptic in equatorial view, circular in polar view, large size P x E = ca. (74-80x56-60) [micro]m. Tricolporate, long colpi with psilate membrane. Exine Gongylolepis type, 12-16 gm thick in the whole grain, spines 2.5 [micro]m length. Ratio ectosexine/ endosexine: ca. 1:1. Nexine ca. 2.5 gm thick. Note: pollen is similar to that of Duidaea marahuacensis and Quelchia bracteata (see also Carlquist, 1957b; Barroso & Maguire, 1973; Telleria, 2008).

Habitat and Distribution. Glossarion is a genus of two species endemic to Cerro de la Neblina in the Guayana Shield of southern Venezuela.

Species list

1. Glossarion bilabiatum (Maguire) Pruski, Brittonia 41: 39. 1989. Basionym: Guaicaia bilabiata Maguire. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, below Pico Phelps, B. Maguire et al. 60475 (holotype, NY!).

2. Glossarion rhodanthum Maguire & Wurdack, Mem. New York Bot. Gard. 9: 390. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 37027 (holotype, US!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Rio Yatua, ca. 2 km N Cumbe camp, B. Maguire et al. 37149 (US*).

Observation. Guaicaia was described as being most closely related to Glossarion, differing from it mainly by capitula having only bilabiate florets. However, capitula of Glossarion frequently contain not only ligulate florets, but also bilabiate ones. Therefore, Glossarion has been redefined by Pruski (1989a) to include Guaicaia.

Literature. Maguire et al. (1957a), Aristeguieta (1964), Maguire (1967), Pruski (1989a, 1991, 1997).

Gochnatia Kunth

in Humb., Bonpl. & Kunth Nov. gen. sp. 4 (folio ed.): 15. 1818. TYPE: Gochnatia vernonioides Kunth. 21 of Figs. 19 22.

Anastraphia D. Don, Trans. Linn. Soc. London 16: 295. 1830. TYPE: Anastraphia ilicifolia D. Don [= Gochnatia sagraeana Jervis & Alain].

Leucomeris D. Don, Prodr. fl. nepal.: 169. 1825. [= Gochnatia sect. Leucomeris (D. Don) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 128. 1971]. TYPE: Leucomeris spectabilis D. Don [= Gochnatia spectabilis (D. Don) Less.].

Pentaphorus D. Don, Trans. Linn. Soc. London 16: 296. 1830. [= Gochnatia sect. Pentaphorus (D. Don) DC., Prodr. 7: 24. 1838]. TYPE: Pentaphorus foliolosus D. Don [= Gochnatia foliolosa (D. Don) D. Don ex Hook. & Am.].

Richterago Kuntze, Revis. gen. pl. 1: 360. 1891. TYPE: Seris discoidea Less. [= Gochnatia discoidea (Less.) Cabrera].

Seris Less., Linnaea 5: 253. 1830, non Willd, 1807, nom. illeg.

Etymology. In honor of Dr. Caroli Gochnat, a physician from Strasburg who studied the Cichorioideae ca. 1808.

Shrubs or subshrubs, rarely trees, monoecious, gynodioecious, or polygamous. Leaves alternate; petiolate; blades ovate, elliptic to linear-elliptic, pinnately to three-veined, margin entire, serrulate to strongly dentate-spinose, subglabrous to glabrous above, densely tomentose beneath. Capitulescences monocephalous, corymbose, racemose, paniculate to glomerulose cymes; capitula pedunculate to subsessile, homogamous or less commonly heterogamous, discoid or disciform; receptacle epaleate; involucre multiseriate, occasionally the phyllaries decurrent onto the peduncle. Florets isomorphic, less commonly sub-dimorphic, bisexual or female with staminodes, corolla actinomorphic, tubular-funnelform, deeply five-lobed, or marginal corollas tubular-bilabiate with tube short (1/3 of corolla length), two incisions deeper than the others; anther apical appendages apiculate, tails smooth to papillose; style bilobed, branches dorsally smooth. Cypselae villose; pappus of scabrid, capillary bristles occasionally somewhat flattened, equal to unequal in length.

Pollen Description. Gochnatia (93 of Figs. 92-94). Pollen spheroidal to prolate, circular in polar view, medium or large size. Tricolporate, colpi long with scabrate membrane, with or without mesoaperture (see also Parra & Marticorena, 1972; Roque & Silvestre-Capelato, 2001; Lin et al., 2005). On the basis of the pollen size and the exine thickness, two pollen types were recognized: type I and type II. Type I: pollen medium size, exine Mutisia type, microechinate, 2 6 [micro]m thick. Found in G. argentina (Cabrera) Cabrera, G. barrosoae Cabrera (93A of Figs. 92-94), G. blanchetiana (DC.) Cabrera, G. hatschbachii Cabrera, G. haumaniana Cabrera, G. polymorpha (Less.) Cabrera (93B, C of Figs. 92-94), G. pulchra Cabrera, G. rusbyana Cabrera, G. sordida (Less.) Cabrera, and G. velutina (Bong.) Cabrera. Gochnatia barrosoae (93A of Figs. 92-94). Pollen spheroidal-subprolate, size P x E = (3944x30-34) [micro]m. Exine 5-6 [micro]m thick, microechinate. Nexine 1.5 [micro]m thick. Gochnatia blanchetiana. Pollen subprolate, medium size, P x E = (30-38x26-32) [micro]m. Exine 2-4 [micro]m thick, scabrate. Nexine ca. 1 [micro]m thick. Gochnatia haumaniana. Pollen spheroidalsubprolate, size P x E = (32-36x25-29) [micro]m. Mesoaperturate. Exine 2-4 [micro]m thick, microechinate. Nexine ca. 1 [micro]m thick. Type lI: pollen large or medium size, exine Wunderlichia or Mutisia type, 6-10 [micro]m thick. Found in G. amplexfolia, G. angustifolia G. Sancho, S. E. Freire & Katinas, G. buchii, G. glutinosa, G. magna, G. palosanto Cabrera, G. rotundifolia Less., G. shaferi, and G. vernonioides Kunth. Gochnatia amplexifolia. Pollen prolate, medium to large size, P x E = (47-51 x 32-36) [micro]m. Mesoaperturate. Exine Wunderlichia type, 6-7 [micro]m thick, echinate, spines 2.5 [micro]m length. Nexine ca. 1.5 [micro]m thick. Gochnatia angustifolia (Sancho et al., 2005). Pollen subprolate, large size, exine Mutisia type, microechinate. Gochnatia buchii (93E of Figs. 92-94). Pollen subprolate-prolate, large size, P x E = (54-58x38-47) [micro]m. Mesoaperturate. Exine Wunderlichia type, 8-10 [micro]m thick, echinate, spines 2.5 [micro]m length. Nexine 2 [micro]m thick. Gochnatia glutinosa. Pollen subprolate, medium size, P x E = ca. (44-49x36-37) [micro]m. Exine Wunderlichia type, ca. 6 [micro]m thick, microechinate. Nexine ca. 1.5 [micro]m thick. Gochnatia magna. Pollen subprolate-prolate, medium to large size, P x E = (45-52x33-38) [micro]m. Exine Wunderlichia type, 8-9.5 [micro]m thick, echinate, spines 2.5 [micro]m length. Nexine ca. 2 [micro]m thick. Gochnatia palosanto. Pollen subprolate, medium to large size, P x E = (49-56x37-44) [micro]m. Exine Wunderlichia type, 8 [micro]m thick, slightly thickened at the poles, microechinate. Nexine ca. 1.5 [micro]m thick. Gochnatia rotundifolia. Pollen prolate, large size, P x E = ca. (56x38) [micro]m. Exine Wunderlichia type, ca. 8 [micro]m thick, echiuate, spines ca. 1.5 [micro]m length. Nexine ca. 2 [micro]m thick. Gochnatia shaferi (93D of Figs. 92-94). Pollen prolate, large size, P x E = (55-6l x 41-47) [micro]m. Exine Wunderlichia type, 8-9 [micro]m thick, microechinate. Nexine 1.5-2 [micro]m thick. Gochnatia vernonioides. Pollen prolate, medium to large size, P x E = (48-62 x 35-43) [micro]m. Exine Wunderlichia type, 7-9 [micro]m thick, microechinate. Nexine 1.5 [micro]m thick.

Habitat and Distribution. Genus of 70 species most from Mexico to Argentina, two species from Asia (China, India, Nepal, Pakistan, and southeastern Asia).

Species list

1. Gochnatia amplexifolia (Gardner) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 155. 1971. Basionym: Seris amplexifolia Gardner. TYPE: Brazil. State Minas Geraes: Diamond District, elevated grassy tracts, G. Gardner 4787 (lectotype, designated by Roque & Pirani, 2001: 1157, BM not seen). Additional specimen examined: Brazil. State Minas Geraes: Santa Ana do Riacho, Serra do Cipo, G. Hatschbach 35312 (LP*).

2. Gochnatia angustifolia G. Sancho, S. E. Freire & Katinas, Taxon 54: 89. 2005. TYPE: Bolivia. Potosi: Jose M. Linares Lizarazu, Jatum Palmar, Torrico & Peca 336 (holotype, LPB!).

3. Gochnatia arborescens Brandegee, Zoe 5: 163. 1903. TYPE: Mexico. State Baja California: Canon de Santa Maria, El Juste, 1 March 1898, C. A. Purpus s.n. (holotype, UC not seen). Additional specimen examined: Mexico. State Baja California: between San Antonio and Port of Bahia de los Muertos, Wiggins 5632 (US).

4. Gochnatia arequipensis Sandwith, Bull. Misc. Inform. Kew: 100. 1934. TYPE: Peru. Arequipa: Volcan de Arequipa, Lechler 2751 (holotype, K not seen). Additional specimen examined: Peru. Arequipa: Monte Chiwata, Eyerdam & Beetle 22120 (LP).

5. Gochnatia argentina (Cabrera) Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. Basionym: Moquinia argentina Cabrera. TYPE: Argentina. Prov. Chaco: Colonia Benitez, Schulz 173 (holotype, LP!). Additional specimen examined: Argentina. Prov. Tucuman: Campo Alegre, Cruz Alta, S. Venturi 703 (LP).

6. Gochnatia argyrea (Dusen ex Malme) Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. Basionym: Moquinia argyrea Dusen ex Malme. TYPE: Brazil. State Parana: Capao Grande, Vila Velha, P. K. H. Dusen 4035 (lectotype, designated by Cabrera, 1971: 93, S!).

7. Gochnatia attenuata (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 8. 1960. Basionym: Anastraphia attenuata Britton. TYPE: Cuba. Prov. Oriente: Serra Nipe, J. A. Shafer 3113 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: Sierra Nipe, E. L. Ekman 19174 (LP).

8. Gochnatia barrosoae Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. TYPE: Brazil. State Minas Geraes: Ituitaba, facenda do fundao, Macedo 1138 (holotype, LP!). Additional specimens examined: Brazil. State Silo Paulo, Mathes 3 (LP*); id., Mogi-Guacu, Mantovani 903 (LP*).

9. Gochnatia blanchetiana (DC.) Cabrera, Notas Mus. La Plata, Bot. 15: 42. 1950. Basionym: Baccharis blanchetiana DC. TYPE: Brazil. State Bahia: Serra Jacobina, Blanchet 2569 (holotype, G-DC!). Additional specimen examined: Brazil. State Minas Gerais, Serra do Espinaco, Irwin et al. 28329 (SP*).

10. Gochnatia boliviana S. F. Blake, Contr. U. S. Natl. Herb. 22: 651. 1924. TYPE: Bolivia. Santa Cruz: Sierra de Santa Cruz, May 1892, C. E. O. Kuntze s.n. (holotype, US!). Additional specimen examined: Bolivia. Dept. Santa Cruz: Florida, A. L. Cabrera & Gutierrez 33804 (LP).

11. Gochnatia buchii (Urb.) J. Jimenez Alm., Phytologia 8: 327. 1962. Basionym: Anastraphia buchii Urb. TYPE: Haiti. Prope Poste Coudau, W. Buch 1017 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Dominican Republic. Falda del Morro: Montecristo, Jimenez 1598 (LP); Montecristo, Jimenez 3613 (LP*).

12. Gochnatia calcicola (Britton) Jervis & Alain, Candollea 17: 120. 1960. Basionym: Anastraphia calcicola Britton. TYPE: Cuba. Prov. Oriente: Guantanamo, L. Britton 2042 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: Guantanamo, E. L. Ekman 15764 (S).

13. Gochnatia cardenasii S. F. Blake, J. Wash. Acad. Sci. 25: 324. 1935. TYPE: Bolivia. Potosi: Quechisla, M. Cardenas 326 (holotype, US!). Additional specimen examined: Argentina. Prov. Jujuy: Tafna, Cordo & Ferrer 88-B-17 (S1).

14. Gochnatia cordata Less., Linnaea 5: 263. 1830. TYPE: Brazil: sine loc, s. d., Sellow s.n. (holotype, B not seen, B photo F n. 15855 at LP!). Additional specimen examined: Argentina. Prov. Entre Rios: Federacion, A. Burkart & Crespo 23169 (LP).

15. Gochnatia cowellii (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 8. 1960. Basionym: Anastraphia cowellii Britton. TYPE: Cuba. Prov. Santa Clara: Palm Barren, Britton & Cowell 10183 (holotype, NY!).

16. Gochnatia crassifolia (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia crassifolia Britton. TYPE: Cuba. Prov. Oriente: Rio Yamaniguey, J. A. Shafer 4261 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: Baracoa, E. L. Ekman 4023 (S).

17. Gochnatia cubensis (Carabia) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia cubensis Carabia. TYPE: Cuba. Prov. Oriente: Sierra de Nipe, J. P. Carabia 3788 (holotype, NY not seen, NY photo at LP!).

18. Gochnatia curviflora (Griseb.) O. Hoffm., Nat. Pflanzenfam. 5: 337. 1893. Basionym: Moquinia curviflora Griseb. TYPE: Bolivia. Tarija: Cuesta del Tambo, P. G. Lorentz & G. Hieronymus 879 (isotype, CORD not seen). Additional specimen examined: Bolivia. Tarija, Fiebrig 2838 (LP).

19. Gochnatia decora (Kurz) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 131. 1971. Basionym: Leucomeris decora Kurz. TYPE: Burma. Pegu, Yomah, S. Kurz 2241 (isotype, G not seen). Additional specimen examined: Myanmar. Bhamo: Lweji, Maung Mya 5309 (LP).

20. Gochnatia densicephala (Cabrera) G. Sancho, Novon 9: 559. 1999. Basionym: Gochnatia paniculata (Less.) Cabrera var. densicephala Cabrera. TYPE: Brazil. State Rio de Janeiro, A. E M. Glaziou 11072 (holotype, K!).

21. Gochnatia discoidea (Less.) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 153. 1971. Basionym: Seris discoidea Less. TYPE: Brazil. Serra da Gacheira, Sellow 794, 994 [lectotype (sic), designated by Roque & Pirani, 2001: 1158, not seen]. Additional specimen examined: Brazil. State Bahia: Igregia Velha, Blanchet 3345 (LP).

22. Gochnatia discolor Baker, F1. bras. 6: 350. 1884. TYPE: Brazil. State Minas Geraes, 1840, Claussen s.n. (isotype, G!).

23. Gochnatia ekmanii (Urb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia ekmanii Urb. TYPE: Cuba. Prov. Pinar del Rio: Sierra de los Organos, E. L. Ekman 13865 (isotype, NY not seen, NY photo at LP!).

24. Gochnatia elliptica (Leon) Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia elliptica Leon. TYPE: Cuba. Prov. Oriente: Ovando, Maisi, Fr. Leon 17580 (isotype, NY not seen, NY photo at LP!).

25. Gochnatia enneantha (S. F. Blake) Alain, Bull. Torrey Bot. Club 92: 304. 1965. Basionym: Anastraphia enneantha S. F. Blake. TYPE: Dominican Republic. Lajama, Samana Peninsula, W. L. Abbott 2287 (holotype, US!). Additional specimen examined: Dominican Republic. Cordillera Central: Samana, Los Haitises, E. L. Ekman H-15498 (S).

26. Gochnatia floribunda Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 125. 1971. TYPE: Brazil. State Minas Geraes: Cerro do Frio, G. Gardner 4806 (isotype, G!).

27. Gochnatia foliolosa (D. Don) D. Don ex Hook. & Am., Comp. Bot. Mag. 1: 108. 1835. Basionym: Pentaphorus foliolosus D. Don. TYPE: Chile: sine loc., s. d., Caldeleugh s.n. (types, probably BM, K), Additional specimen examined: Chile. Prov. Santiago: Santiago, Cerro Renca, A. L. Cabrera 3451 (LP).

28. Gochnatia gardneri (Baker) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia gardneri Baker. TYPE: Brazil. State Goyaz: Capella da Passe, G. Gardner 4183 (lectotype, designated by Cabrera, 1971: 109, K!).

29. Gochnatia glutinosa (D. Don) D. Don ex Hook. & Am., Comp. Bot. Mag. 1: 108. 1835. Basionym: Pentaphorus glutinosus D. Don. TYPE: Argentina. Prov. Mendoza: near San Isidro, s.d., J. Gillies s.n. (type, probably K). Additional specimens examined: Argentina. Prov. Tucuman: Taft, H. A. Fabris 1343 (LP); Prov. Salta: Trancas, Cuesta del Anca, Jan 1897, C. L. Spegazzini s.n. (LP*).

30. Gochnatia gomezii (Leon) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia gomezii Leon. TYPE: Cuba. Prov. Oriente: Cerro de Miraflores, Fr. Leon & Gomez 28876 (holotype, HAC not seen). Additional specimen examined: Cuba. Prov. Oriente: Moa, Cerro Miraflores, Marie-Victorin et al. 21591 (GH).

31. Gochnatia hatschbachii Cabrera, Bol. Mus. Bot. Munic. 15: 3. 1974. TYPE: Brazil. State Minas Geraes: Jaboticatubas, Serra do Cipo, G. Hatschbach 29951 (holotype, LP!).

32. Gochnatia haumaniana Cabrera, Notas Mus. La Plata, Bot. 15: 44. 1950. TYPE: Paraguay. Pedro Juan Caballero: Sierra de Amambay, T. Rojas 6575 (holotype LP!). Additional specimen examined: Paraguay. Pedro Juan Caballero: Sierra de Amambay, T. Rojas 10391 (K*).

33. Gochnatia hiriartiana Medrano, Villasenor & Medina, Novon 14: 434. 2004. TYPE: Mexico. Hidalgo: Meztitlan, Gonzalez-Medrano, G. G. Hernandez & G. Rodriguez 17920 (holotype, MEXU not seen).

34. Gochnatia hypoleuca (DC.) A. Gray, Proc. Amer. Acad. Arts 19: 57. 1883. Basionym: Moquinia hypoleuca DC. TYPE: Mexico. State Nueva Leon: Monterrey, Berlandier 1391 (isotype, US!). Additional specimen examined: Mexico. State Hidalgo: Canada del Vaquero, Gonzalez Quintero 3215 (LP).

35. Gochnatia ilicifolia Less., Linnaea 5: 261. 1830. TYPE: America (Guiane?), sine loc., s.d., Richard s.n. (holotype, B not seen). Additional specimen examined: Bahamas Islands. Andros Island: Coppice, Small & Carter 8526 (K).

36. Gochnatia intertexta (C. Wright ex Griseb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia inertexta C. Wright ex Griseb. TYPE: Cuba. Bahia Onda, prope San Marcos, C. Wright 2877 (isotype, BM not seen). Additional specimen examined: Cuba. Prov. Pinar del Rio: Cajalbana, Alain A-1680 (NY).

37. Gochnatia magna M. C. Johnst. ex Cabrera, Revista Mus. La Plata, Secc. Bot. 12:147. 1971. TYPE: Mexico. State Tamaulipas: half mile E from Nuevo Morelos, J. Graham & M. C. Johnston 4485 (holotype, MEXU not seen). Additional specimen examined: Mexico. San Luis Potosi, A. Cronquist 11277 (NY).

38. Gochnatia maisiana (Leon) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia maisiana Leon. TYPE: Cuba. Prov. Oriente: Mesa del Chivo, Fr. Leon 18243 (isotype, NY not seen, NY photo at LP!).

39. Gochnatia mantuensis (C. Wright ex Griseb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia mantuensis C. Wright ex Griseb. TYPE: Cuba. Nueva Filipina, prope Mantua, C. Wright 2876 (isotype, BM not seen). Additional specimen examined: Cuba. Prov. Pinar del Rio: Guane, Shafer 11208 (LP).

40. Gochnatia microcephala (Griseb.) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia microcephala Griseb. TYPE: Cuba. Ex Catae, 1860 64, C. Wright s.n. (isotype, NY not seen). Additional specimen examined: Cuba. Boca Guantanamo a Mantua, Bro. Hioram 4874 (NY).

41. Gochnatia mollisima (Malme) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia mollissima Maline. TYPE: Brazil. Rio Grande do Sul, G. A. Malme 648 (holotype, S!).

42. Gochnatia montana (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia montana Britton. TYPE: Cuba. Prov. Pinar del Rio: Sierra Caliente, J. A. Shafer 13781 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Guane, E. L. Ekman 18725 (S).

43. Gochnatia obtusifolia (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia obtusifolia Britton. TYPE: Cuba. Prov. Oriente: camp La Gloria, J. A. Shafer 8189 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Southern Baracoa region, Fr. Leon 11963 (NY).

44. Gochnatia oligantha (Urb.) R. A. Howard, J. Arnold Arbor. 42: 137. 1961. Basionym: Anastraphia oligantha Urb. TYPE: Dominican Republic. Santo Domingo, 1871, C. Wright et al. 248 (isotype, NY!). Additional specimen examined: Dominican Republic. Montecristo, N of Villa Isabel, Jimenez 3614 (US).

45. Gochnatia oligocephala (Gardner) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia oligocephala Gardner. TYPE: Brazil. State Ceara: Serra de Araripe, G. Gardner 2422 (holotype, BM not seen). Additional specimen examined: Brazil. State Bahia: Rio Branco, C. H. Curran 284 (NY).

46. Gochnatia orbiculata (Malme) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Moquinia orbiculata Malme. TYPE: Brazil. State Parana: Vilha Velha, P. Dusen 9437 (lectotype, designated by Cabrera 1971: 105, S!). Additional specimen examined: Brazil. State Rio Grande do Sul, Tweedie 998 (K).

47. Gochnatia palosanto Cabrera, Notas Mus. La Plata, Bot. 1: 62. 1935. TYPE: Argentina. Prov. Tucuman: Trancas, Vipos, C. Schreiter 85 (holotype, LP!). Additional specimen examined: Argentina. Prov. Salta: Dept. Candelaria, Sierra de la Candelaria, S. Venturi 9793 (LP*).

48. Gochnatia paniculata (Less.) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Spadonia paniculata Less. TYPE: Brazil. Sao Paulo: Casa Branca, Loefgren 16994 (neotype, designated by Sancho, 1999: 557, US!).

49. Gochnatia parvifolia (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia parvifolia Britton. TYPE: Cuba. Prov. Oriente: Barren Savannas, Shafer 2938 (holotype, NY!).

50. Gochnatia patazina Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 28. 1971. TYPE: Peru. La Libertad: Pataz, between Hualillas and Tayabamba, A. Lopez & A. Sagdstegui 3409 (holotype, LP!).

51. Gochnatia pauciflosculosa (C. Wright ex Hitchc.) Jervis ex Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 48. 1971. Basionym: Anastraphia pauciflosculosa C. Wright ex Hitchc. TYPE: Bahamas Islands. Inagua: 12 April 1890, A. S. Hitchcock s.n. (lectotype, designated by Cabrera, 1971: 50, MO not seen). Additional specimen examined: Bahamas Islands. Fortune Island, Eggers 3866 (K).

52. Gochnatia picardae (Urb.) J. Jimenez Alto., Phytologia 8: 327. 1962. Basionym: Anastraphia picardae Urb. TYPE: Haiti. Prope Cadets, Picarda 1073 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Haiti. Massif de la Selle: Croix-des Bauquets, gorge of Grande-Riviere de cul de sac, E. L. Ekman H-5385 (K).

53. Gochnatia polymorpha (Less.) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Spadonia polymorpha Less. TYPE: Brazil: s.d., Sellow s. n., (holotype, B not seen). Additional specimens examined: Brazil. State Bahia: Jacobina, Igreja Velha, Blanchet 3251 (F); State Rio de Janeiro: Tijuca, A. E M. Glaziou 3039 (LP*).

54. Gochnatia pulchra Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 106. 1971. TYPE: Brazil. Sao Paulo: Ityrapina, 28 April 1923, Gehrt s.n. (holotype, LP!).

55. Gochnatia purpusii Brandegee, Zoe 5: 240. 1906. TYPE: Mexico. State Puebla: Tehuacan, C. A. Purpus 1164 (holotype, UC not seen). Additional specimen examined: Mexico. Tehuacan, Purpus 4248 (NY).

56. Gochnatia ramboi Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 98. 1971. TYPE: Brazil. State Rio Grande do Sul: Palmeira, B. Rambo 51961 (holotype, LP!).

57. Gochnatia recurva (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. Basionym: Anastraphia recurva Britton. TYPE: Cuba. Prov. Oriente: Serpentine hills near mouth of Rio Yamaniguey, J. A. ShaJer 4257 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Cuba. Prov. Oriente: between Moa and Punta Anden, Leon 20946 (LP).

58. Gochnatia rotundifolia Less., Syn. gen. Compos.: 102. 1832. TYPE: Brazil: sine loc., s.d., Sellow s.n. (holotype, B not seen). Additional specimens examined: Brazil. State Sao Paulo: Capital, Jabaquara, O. Handro 157 (LP); id., Ypiranga, Brade 6265 (LP*).

59. Gochnatia rusbyana Cabrera, Notas Mus. La Plata, Bot. 15: 41. 1950. TYPE: Bolivia. Yungas, M. Bang 2252 (isotypes, G!, LP!, MO!).

60. Gochnatia sagraeana Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 9. 1960. TYPE: Cuba. Habana, Jibacoa, Fr. Leon et al. 19708 (type, probably HAC). Additional specimen examined: Cuba. Habana, Fr. Leon 7094 (NY).

61. Gochnatia shaferi (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18: 10. 1960. Basionym: Anastraphia shaferi Britton. TYPE: Cuba. Prov. Oriente: Rio Naranja, J. A. Shafer 3865 (isotype, US!). Additional specimens examined: Cuba. Prov. Oriente: La Caridad, Lopez Figueiras 1738 (NY); id., Sierra de Nipe, Victorin & Clemente 22031 (LP, NY*).

62. Gochnatia smithii B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 32: 50. 1896. TYPE: Mexico. State Oaxaca: Cuicatlan, L. C. Smith 363 (lectotype, designated by Cabrera, 1971:140, K not seen). Additional specimen examined: Mexico. State Oaxaca: Cuesta de Coyula, Conzatti 4135 (US).

63. Gochnatia sordida (Less.) Cabrera, Notas Mus. La Plata, Bot. 15: 43. 1950. Basionym: Spadonia polymorpha Less. var. sordida Less. TYPE: Brazil. s.d., Sello s.n. (isotype, K!).

64. Gochnatia spectabilis (D. Don) Less., Linnaea 5: 261. 1830. Basionym: Leucomeris spectabilis D. Don. TYPE: Nepal. Sirinagur, 1821, Wallich s.n. (isotype, G not seen). Additional specimen examined: India. Uttar Pradesh: Rajpur, Dehra Dun District, Galrola 32 (LP).

65. Gochnatia suffrutescens Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 150. 1971. TYPE: Brazil. State Minas Geraes: Sao Joao del Rey, Mello Barreto 4744 (holotype, F not seen).

66. Gochnatia tortuensis (Urb.) J. Jimenez Alto., Phytologia 8: 328. 1962. Basionym: Anastraphia tortuensis Urb. TYPE: Haiti: La Tortue Island, E. L. Ekman H-4313 (isotype, US!). Additional specimen examined: Haiti. Presquile du Nord-Ouest: Port-de Paix, E. L. Ekman H-3553 (S).

67. Gochnatia vargasii Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 19. 1971. TYPE: Peru. Apurimac: Abancay, C. Vargas 16317 (holotype, LP!).

68. Gochnatia velutina (Bong.) Cabrera, Notas Mus. La Plata, Bot. 15: 44. 1950. Basionym: Moquinia velutina Bong. TYPE: Brazil. State Sao Paulo: s.d., Langsdorff s.n. (holotype, LE not seen, LE photo at LP!). Additional specimen examined: Brazil. State Parana: Ponta Grossa, Vila Velha, R. M. Klein & S. B. Smith 14885 (LP).

69. Gochnatia vernonioides Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 16. 1818. TYPE: Peru. Dept. Amazonas: s.d., F. Humboldt & A. Bonpland s.n. (isotype, K not seen). Additional specimens examined: Peru. Dept. Amazonas: Prov. Chachapoyas, Tingo, R. Ferreyra 7097 (LP); Prov. Chachapoyas, Leimebamba, A. Lopez et al. 4364 (LP*).

70. Gochnatia wilsonii (Britton) Jervis & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 18:10. 1960. Basionym: Anastraphia wilsonii Britton. TYPE: Cuba. Santa Clara: Rio Tobaya, Trinidad, N. L. Britton et al. 5573 (holotype, NY not seen, NY photo at LP!).

Observation. See Observations in Actinoseris. The Asian species of Gochnatia belonging to the sect. Leucomeris (G. decora and G. spectabilis) as well as those belonging to the sect. Pentaphorus (G. foliolosa and G. glutinosa) have been lately regarded as independent genera from Gochnatia (i.e., Leucomeris and Pentaphorus respectively; Hind, 2007; Panero & Funk, 2008).

Literature. Jervis (1954), Cabrera (1971a, 1974), Sancho (1996, 1999, 2000), Freire et al. (2002), Gonzalez-Medrano et al. (2004), Sancho et al. (2005).

Gongylolepis R. H. Schomb.

Linnaea 20: 759. 1847. TYPE: Gongylolepis benthamiana R. H. Schomb. 22 of Figs. 19-22.

Cardonaea Aristeg., Maguire & Steyerm., Mere. New York Bot. Gard. 23: 892. 1972. TYPE: Cardonaea jauaensis Aristeg., Maguire & Steyerm. [= Gongylolepis jauaensis (Aristeg., Maguire & Steyerm.) V. M. Badillo].

Neocaldasia Cuatrec. Revista Acad. Colomb. Ci. Exact. 6: 16. 1944. TYPE: Neocaldasia colombiana Cuatrec. [= Gongylolepis colombiana (Cuatrec.) Cuatrec.].

Etymology. From the Greek gongylos, round, and lepis, scale, in reference to the rounded phyllary apices.

Shrubs or trees, less commonly subshrubs. Leaves alternate, often clustered at the apex; sessile to petiolate, petioles, when present, commonly clasping; blades oblanceolate, lanceolate to obovate, coriaceous, venation pinnate, sub-palmate or sub-parallel, margin entire, sometimes revolute, glabrous to abaxially pubescent. Capitulescences monocephalous or cymose; capitula sessile to pedunculate, peduncles sometimes with leaf-like bracts, homogamous, bilabiate, six- to many-flowered; receptacle epaleate; involucre multiseriate, campanulate to hemispherical, phyllaries graduate. Florets isomorphic, bisexual, corolla bilabiate; anthers purple-brown, mostly exserted, apical appendages acuminate, tails slightly papillose, often connate; style white to tinged with purple, bifid, branches dorsally smooth. Cypselae commonly glabrous; pappus of scabrid, persistent capillary bristles.

Pollen Description. Gongylolepis (94 of Figs. 92-94). Pollen spheroidal to subprolate, spheroidal or elliptic in equatorial view, circular in polar view, medium or large size. Tricolporate, colpi long or short with margin and psilate membrane. Exine Gongylolepis type, uniformly thickened. Note: pollen very similar to that of Achnopogon (see also Carlquist, 1957b; Barroso & Maguire, 1973). Gongylolepis benthamiana. Pollen subprolate, elliptic in equatorial view, large size, P x E = (72-94x61-78) [micro]m. Colpi 47-60 [micro]m length. Exine 10-11 [micro]m thick in the whole grain, spines 2.5-4 [micro]m length. Nexine 2-4 [micro]m thick. Gongylolepis huachamacari (94 of Figs. 92-94). Pollen spheroidal-subprolate, spheroidal in equatorial view, large size p x E = (74-88 x 70-80) [micro]m. Long colpi. Exine 7-10 [micro]m thick, spines 5-6 [micro]m length. Nexine ca. 3 [micro]m thick. SEM: tectum very perforate with striate aspect, apertural membrane scabrate (see also Telleria, 2008).

Habitat and Distribution. The genus contains 14 species, 13 of which are endemic to the Guayana region of Colombia, Venezuela, Guyana, extending southward toward Manaus, Brazil. However, one species, Gongylolepis colombiana (Cuatrec.) Cuatrec., occurs in the northern Andes of Colombia and Venezuela.

Species list

1. Gongylolepis benthamiana R. H. Schomb., Linnaea 20: 760. 1847. TYPE: Guyana: Schomburgk 11 (holotype, K not seen). Additional specimen examined: Venezuela. State Bolivar: Distr. Roscio, 13 km al N de Santa Elena de Vaiven, J. A. Steyermark & R. Liesner 127536 (MO*).

2. Gongylolepis bracteata Maguire, Mere. New York Bot. Gard. 8: 156. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Sipapo, B. Maguire & L. Politi 27604 (isotype, US).

3. Gongylolepis colombiana (Cuatrec.) Cuatrec., Fieldiana, Bot. 27: 51. 1950. Basionym: Neocaldasia colombiana Cuatrec. TYPE: Colombia. Norte de Santander, J. Cuatrecasas 13435 (isotype, US!). Additional specimen examined: Venezuela. State Tachira, H. van der Werff & F. Ortega 8077 (MO).

4. Gongylolepis cortesii (S. Diaz) Pruski & S. Diaz, FI. Venez. Guayana 3: 285. 1997. Basionym: Eutydochus cortesii S. Diaz. TYPE: Colombia. Guinia: Serrania del Naquen, camino de Carlo Rico al Campamento 15, Cortes et al. 238 (isotype, US!).

5. Gongylolepis erioclada S. F. Blake, Bull. Torrey Bot. Club 58: 495. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Duida, G. H. H. Tate 758 (holotype, US!).

6. Gongylolepis fruticosa Maguire, Steyerm. & Wurdack, Mere. New York Bot. Gard. 9: 437. 1957. TYPE: Venezuela. State Bolivar: cerro Chimanta, J. A. Steyermark & J. J. Wurdack 1136 (holotype, NY!).

7. Gongylolepis glaberrima S. F. Blake, Bull. Torrey Bot. Club 58: 494. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Mt. Duida, G. H. H. Tate 617 (holotype, NY!).

8. Gongylolepis huachamacari Maguire, Mere. New York Bot. Gard. 8: 159. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Huachamacari, B. Maguire et al. 30135 (holotype, NY!). Additional specimens examined: Venezuela. Dept. Rio Negro: Cerro Aracamuni, 1[degrees] 26' N, 65[degrees] 47' W, R. Liesner & F. Delascio 21960 (MO*); Terr. Fed. Amazonas, Cerro Huachamacari, B. Maguire et al. 30080 (MO*).

9. Gongylolepis jauaensis (Aristeg., Steyerm. & Wurdack) V. M. Badillo, Ernstia 46: 8. 1988. Basionym: Cardonaea jauaensis Aristeg., Steyerm. & Wurdack. TYPE: Venezuela. State Bolivar: Meseta de Jaua, Cerro Jaua, J. A. Steyermark 97945 (holotype, VEN!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dept. Atabapo, cerro Marahuaca, below Salto Los Homos, R. Liesner 25112 (MO).

10. Gongylolepis martiana (Baker) Steyerm. & Cuatrec., Fieldiana, Bot. 28: 640. 1953. Basionym: Stifftia martiana Baker. TYPE: Colombia. Terr. Fed. Amazonas: Araracuara, s.d., C. Martius s.n. (holotype, M not seen). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dept. Atures, 10 km al S del Rio Guayapo, O. Huber 3293 (MO).

11. Gongylolepis oblanceolata Pruski, Ann. Missouri Bot. Gard. 76: 994. 1989. TYPE: Brazil. State Amazonas: Barcelos, Plato da Serra Araca, M. G. Silva et al. 7169 (holotype, INPA!). Additional specimen examined: Brazil. State Amazonas: Municipio de Barcelos, plato da Serra Araca, I. L. Amaral 1557 (US).

12. Gongylolepis paniculatu Maguire & K. D. Phelps, Mem. New York Bot. Gard. 8: 157. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Guanay, B. Maguire et al. 31655 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Cerro Yavi, K. Phelps & C. Hitchcock 8 (MO).

13. Gongylolepis paruana Maguire, Mere. New York Bot. Gard. 8: 159. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Serrania Paru, Rio Paru, Cano Asisa, Rio Ventuari, R. S. Cowan & J. J. Wurdack 31239 (holotype, NY!).

14. Gongylolepis pedunculatu Maguire, Mere. New York Bot. Gard. 8: 156. 1953. TYPE: Venezuela. Amazonas: Serrania Paru, Rio Paru, Carlo Asisa, Rio Ventuari, R. S. Cowan & J. J. Wurdack 31131 (holotype, NY!).

Literature. Maguire et al. (1957a), Aristeguieta (1964), Steyermark et al. (1972), Badillo (1988), Pruski (1989b, 1991, 1997), Diaz-Piedrahita and Velez-Nauer (1993).

Gypothamnium Phil.

FI. atacam.: 27. 1860. TYPE: Gypothamnium pinifolium Phil. 23 of Figs. 23-26. Etymology. From the Greek gyps, gypos, vulture, and thamnos, shrub.

Shrubs. Leaves alternate, densely arranged in the upper branches; sessile; blades linear, margin entire, one-veined, glabrous. Capitulescences monocephalous, terminal; capitula sessile, heterogamous, radiate; receptacle epaleate; involucre multiseriate. Florets dimorphic; marginal florets female, with well-developed staminodes, corolla bilabiate, outer lip expanded; central florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages reddish, truncate, tails papillose; style bilobed, branches dorsally papillose. Cypselae villose; pappus of scabrid bristles, the innermost flattened, ciliate at the apex, the outermost shorter, not ciliate, rigid.

Pollen Description (Telleria & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate (see also Parra & Marticorena, 1972).

Habitat and Distribution. Monotypic genus endemic to northern Chile.

Species list

1. Gypothamnium pinifolium Phil., Fl. atacam.: 27. 1860. TYPE: Chile. Prov. Antofagasta: cerca de las Animas, Cachinal de la Costa, Taltal, en el valle de Guanillo mas arriba de Paposo, etc., R. Philippi 2349, s.n. (syntypes, SGO not seen, SGO photos at LP!). Additional specimen examined: Chile. Prov. Antofagasta: Taltal, Francke 1044 (LP).

Literature. Cabrera (1951).

Hecastocleis A. Gray

Proc. Amer. Acad. Arts 17: 220. 1882. TYPE: Hecastocleis shockleyi A. Gray. 24 of Figs. 23-26.

Etymology. From the Greek hekastos, each, and kleis, lock, key, to shut up, which refers to each floret enclosed by the phyllaries of the involucre.

Shrubs. Leaves alternate; sessile; blades linear-elliptic to oblanceolate, pinnately veined, margin spiny, glabrous above, subglabrous beneath. Capitulescence a pseudocephalium, several-headed, surrounded by spiny wide-ovate bracts; capitula sessile, one-flowered; receptacle epaleate; involucre multiseriate, cylindrical, phyllaries coriaceous with spiny apex. Florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages subacute, tails smooth; style bilobed, branches dorsally smooth. Cypselae glabrous; pappus a ring of short, laciniate scales.

[FIGURES 23-26 OMITTED]

Pollen Description (Telleria & Katinas, 2005) (95 of Figs. 95-100). Pollen spheroidal, medium size, tricolpate, exine Mutisia type, scabrate.

Habitat and Distribution. Monotypic genus endemic to dry, rocky slopes in western United States.

Species list

1. Hecastocleis shocklevi A. Gray, Proc. Amer. Acad. Arts 17: 221. 1882. TYPE: United States. State Nevada: Esmeralda County, Candelaria, W. H. Schockley 55 (isotype, US not seen, US photo at LP!). Additional specimens examined: United States. State Nevada: 5 mi. of Basalt, on Mt. Montgomery-Tonopah highway, Mineral-Esmeralda Co. line, Train 3973 (LP*); State California, Inyo Co., Eureka valley to Fish Lake valley, Alexander & Kellogg 5301 (US*); Death valley region, Leadfield, Gilman 1144 (US*).

Literature. Simpson (1978), Panero and Funk (2002).

Hyalis D. Don ex Hook. & Arn.

Comp. Bot. Mag. 1: 108. 1835. TYPE: Hyalis argentea D. Don ex Hook. & Am. 25 of Figs. 23-26.

Etymology. From the Greek hyaleos, of glass, transparent, probably because of the light color of the leaves or the shiny, scarious involucral bracts.

Subshrubs rhizomatous, single to few-branched stems. Leaves alternate; sessile; blades oblong to linear-elliptic, margin entire to serrulate, three-veined, glabrous to pubescent. Capitulescences densely corymbose; capitula short-pedunculate, homogamous, radiate, five- to six-flowered; receptacle epaleate; involucre three- to six-seriate. Florets dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded; central floret 1, tubular-funnelform corolla, deeply five-lobed; anther apical appendages apiculate, tails papillose; style bifid, branches dorsally smooth. Cypselae villose; pappus of scabrid to smooth, capillary bristles, unequal in length.

Pollen Description. Hyalis argentea (Telleria & Forcone, 2002). Pollen prolatesubprolate, medium to large size, tricolporate, exine Mutisia type, microechinate.

Habitat and Distribution. Genus of two species that grow in sandy areas and in forests of Schinopsis Engl. (Anacardiaceae) and Copernicia Mart. (Arecaceae) of northern Argentina, Bolivia, and Paraguay.

Species list

Hyalis argentea D. Don ex Hook. & Am., Comp. Bot. Mag. 1: 108. 1835. TYPE: Argentina. Prov. Mendoza: Mendoza, s.d., J. Gillies s.n. Prov. San Luis: Rio Cuarto, s.d., J. Gillies s.n.. Prov. Cordoba: Villa del Rio Cuarto, pampas of Cordoba, s.d., J. Gillies s.n. Prov. Buenos Aires: salt plains of Bahia Blanca, north Patagonia, s.d., Tweedie s.n. (syntypes, probably K). Additional specimen examined: Argentina. Prov. Mendoza: Tupungato, A. Ruiz Leal 3701 (LP).

2. Hyalis lancifolia Baker, F1. bras. 6: 368. 1882. TYPE: Argentina. Prov. Salta: s.d., Pearce s.n. (type, not localized). Additional specimen examined: Argentina. Prov. Chaco: San Fernando, A. L. Cabrera 4083 (LP).

Literature. Cabrera (1951, 1978), Freire et al. (2002).

Hyaloseris Griseb.

Symb. ft. argent.: 212. 1879. TYPE: Gochnatia cinerea Griseb. [= Hyaloseris cinerea (Griseb.) Griseb.]. 26 of Figs. 23-26.

Etymology. From the Greek hyaleos, of glass, and seris, chicory, lettuce, probably because of the shiny, scarious involucral bracts.

Shrubs or little trees, with brachyblasts and macroblasts. Leaves alternate to opposite, clustered in the brachyblasts; short-petiolate; blades linear-oblong, elliptic to ovate, pinnately veined, margin entire to dentate, glabrous to pubescent. Capitulescences monocephalous or two- to four-headed, racemose to spicate; capitula sessile to pedunculate, homogamous, discoid or ligulate; receptacle epaleate; involucre multiseriate. Florets isomorphic or with transitional corollas in the same capitulum, bisexual, ligulate, shallowly five-dentate limb, occasionally some segments partially connate, sub-bilabiate (4+1 corolla lips), rarely bilabiate; anther apical appendages acute, tails papillose; style bifid, branches dorsally papillose. Cypselae glabrous to pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Hyaloseris cinerea (96 of Figs. 95-100) (Telleria & Katinas, 2004). Pollen subprolate, large size, tricolporate, exine Mutisia type, microechinate.

Habitat and Distribution. Genus of seven species of dry and rocky soils in Bolivia and Argentina.

Species list

1. Hyaloseris andrade-limae Cristobal & Cabrera, Hickenia 1: 255. 1982. TYPE: Argentina. Prov. Santiago del Estero: Dept. Guasayan, Sierra de Guasayan, ruta 64, 70 km de Santiago del Estero, A. Krapovickas et al. 37415 (isotype, LP!).

2. Hyaloseris boliviensis J. Kost., Blumea 5: 668. 1945. TYPE: Bolivia. Pulquina, T. Herzog 1798 (isotype, LP!). Additional specimen examined: Bolivia. Dept. Cochabamba: Prov. Totora, Higuerita, Steinbach 3877 (LIL, not seen).

3. Hyaloseris camataquiensis Hieron. ex J. Kost., Blumea 5: 667. 1945. TYPE: Bolivia. Catamaqui, Fiebrig 3067 (isotype, S not seen). Additional specimen examined. Bolivia. Quechisla, M. Cardenas 11 (LP).

4. Hyaloseris cinerea (Griseb.) Griseb., Symb. ft. argent.: 213. 1879. Basionym: Gochnatia cinerea Griseb. TYPE: Argentina. Prov. Catamarca: Fuerte de Andalgala, P. G. Lorentz 690 (isotype, CORD not seen). Additional specimen examined: Argentina, Prov. Cordoba, entre Taninga y Chancani, Sierra de Pocho, A. Ragonese 9335 (LP*).

5. Hyaloseris longicephala B. L. Turner, Phytologia 59:317. 1986. TYPE: Bolivia. Chuquisaca: Prov. Oropeza, an der strasse yon Sucre Richtung Cochabamba, vor Surima, T. Feuerer & N. Hohne 45762 (holotype, TEX not seen).

6. Hyaloseris quadriflora J. Kost., Blumea 5: 667. 1945. TYPE: Bolivia. Cochabamba: Cerro de San Pedro, T. Herzog 2461 (isotypes, CONC!, LP!).

7. Hyaloseris rubicunda Griseb., Symb. ft. argent.: 213. 1879. TYPE: Argentina. Prov. Catamarca: Dept. Belen, Portezuelo, entre Cortadera y Cienaga, Sehickendantz 284 (lectotype, designated by Ariza Espinar, 1973:205 as "isotype", CORD not seen). Additional specimen examined: Argentina. Prov. Salta: alrededores de Cachi, Jan 1897, C. L. Spegazzini s.n. (LP).

Literature. Ariza Espinar (1973), Cristobal and Cabrera (1982), Turner (1986).

Ianthopappus Roque & D. J. N. Hind

Novon 11: 97. 2001. TYPE: Gochnatia corymbosa Less. [= Ianthopappus corymbosus (Less.) Roque & D. J. N. Hind]. 27 of Figs. 27-30.

Etymology. From the Latin ianthum, purple, and pappus, apical tuft of bristles on the fruit, referring to the color of the pappus.

Subshrubs unbranched. Leaves alternate; short-petiolate; blades wide-obovate to orbicular, three-veined, margin entire, glabrous above, densely white-tomentose beneath. Capitulescences corymbose, two- to 18-headed; capitula long-pedunculate, heterogamous, radiate; receptacle epaleate; involucre multiseriate, three- to six-seriate. Florets dimorphic; marginal florets female with staminodes, corollas bilabiate, outer lip expanded; central florets bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed; anther apical appendages apiculate, tails papillose; style bilobed, branches dorsally smooth. Cypselae villose; pappus purplish, scabrid bristles, the innermost longer and plumose at the apex.

Pollen Description. lanthopappus corymbosus (97 of Figs. 95-100) (Roque & Silvestre Capelato, 2001). Pollen prolate, large size, tricolporate, exine Mutisia type, microechinate. Pollen close to that of type II of Gochnatia.

Habitat and Distribution. Monotypic genus from eastern Argentina, southeastern Brazil, and northwestern Uruguay in "campestre" vegetation with predominance of steppe.

Species list

1. Ianthopappus corymbosus (Less.) Roque & D. J. N. Hind, Novon 11: 97. 2001. Basionym: Gochnatia corymbosa Less. TYPE: Brazil. Sello 3479 (neotype, designated by Cabrera, 1970: 52, K not seen). Additional specimens examined: Argentina. Prov. Corrientes: Paso Troncon, Palacios-Cuezzo 2304 (LP); Uruguay. Dept. Artigas: Rio Cuareim, J. Chebataroff 3015 (LP*).

[FIGURES 27-30 OMITTED]

Literature. Cabrera (1970), Roque and Hind (2001), Freire et al. (2002).

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 2
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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