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The subfamily Mutisioideae (Asteraceae).

Holocheilus Cass.

Bull. Sci. Soc. Philom. Paris: 73. 1818. TYPE: Perdicium brasiliense L. [= Holocheilus brasiliensis (L.) Cabrera]. 54 of Figs. 51-54.

Cleanthes D. Don, Trans. Linn. Soc. London 16: 194. 1830. TYPE: Cleanthes hieracioides D. Don [= Holocheilus hieracioides (D. Don) Cabrera].

Platycheilus Cass., Dict. sci. nat. 34: 212. 1825. TYPE: Platycheilus ochroleucus Cass. [= Holocheilus brasiliensis (L.) Cabrera].

Etymology. From Greek bolos, whole, all, entire, and cheilos, lip, because of the capitula with all the corollas bilabiate.

Herbs perennial, caulescent to acaulescent, scapose. Leaves pinnately veined, margin entire to runcinate, glabrous to pubescent; basal leaves rosulate; pseudopetiolate; blades elliptic, obovate, oblanceolate to spathulate, entire to pinnatisect; upper leaves alternate, reduced to bracts; sessile to clasping. Capitulescences monocephalous or loosely to densely corymbose; capitula pedunculate to subsessile, homogamous, discoid; receptacle epaleate; involucre one- to two-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae rostrate to slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles.

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

Habitat and Distribution. Genus of seven species distributed in southern Brazil, Paraguay, Uruguay, and northern Argentina.

Species list

1. Holocheilus brasiliensis (L.) Cabrera, Revista Mus. La Plata, Secc. Bot. 11 : 14. 1968. Basionym: Perdicium brasiliense L. TYPE: Brazil. s.d., Arduini s.n. (lectotype, designated by Katinas in Jarvis & Turland, 1998: 365, LINN 1003.4 not seen, LINN photo at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: Pdo. Tornquist, Abra de la Ventana, L. Pertusi 214 (LP).

2. Holocheilus fabrisii Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 9. 1968. TYPE: Argentina. Prov. Jujuy: Dept. Santa Barbara, cerro Centinela, H. A. Fabris et al. 5145 (holotype, LP!).

3. Holocheilus hieracioides (D. Don) Cabrera, Revista Mus. La Plata, Sect. Bot. 11: 7. 1968. Basionym: Cleanthes hieracioides D. Don. TYPE: Brazil: s.d., Sello s.n. (type, probably P). Additional specimen examined: Argentina. Prov. Salta: Dept. Capital, ruta 9 entre Castanares y Rio Vaqueros, V. Nunez 6 (MCNS).

4. Holocheilus illustris (Vell.) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 6. 1968. Basionym: Castra illustris Vell. TYPE: Tab. 81. T.8 (type, not localized). Additional specimen examined: Brazil. State Parana: Mun. Punta Grossa, est. Velha, rod. Do Cafe-Itaiacoca, G. Hatschbach 11593 (LP).

5. Holocheilus monocephalus Mondin, Napaea 11: 31. 1995. TYPE: Brazil. State Rio Grande do Sul: mun. Silo Jose dos Ausentes, Rocinha, C. A. Mondin et al. 982 (holotype, ICN not seen).

6. Holocheilus pinnatifidus (Less.) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 4. 1968. Basionym: Trixis pinnatifida Less. TYPE: Brazil: s.d., Sello s.n. (syntypes, probably P). Additional specimen examined: Brazil. State Sao Paulo: Bananal, serra da Bocaina-Sertao do rio Vermelho, A. C. Brade & A. P. Duarte 20126 (LP).

7. Holocheilus schulzii (Cabrera) Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 12. 1968. Basionym: Trixis schulzii Cabrera. TYPE: Argentina. Prov. Chaco: Colonia Benitez, A. Schulz 132 (holotype, LP!). Additional specimen examined: Argentina. Prov. Corrientes: estancia Garruchos, A. L. Cabrera 11840 (LP).

Literature. Cabrera (1936, 1968), Mondin (1995), Mondin and Vasques (2004), Katinas and Crisci (2008).

Jungia L. f., nom. cons.

Suppl. pl.: 58. 1781 (1782). TYPE: Jungia ferruginea L. f. 55 of Figs. 55-58.

Dumerilia Lag. ex DC., Ann. Mus. Natl. Hist. Nat. 19: 71. 1812, non Less., 1830. [= Jungia sect. Dumerilia (Lag. ex DC.) Harling, Acta Regiae Soc. Sci. Litt. Gothob., Bot. 4: 32. 1995]. TYPE: Dumerilia axillaris Lag. ex DC. [= Jungia axillaris (Lag. ex DC.) Spreng.].

Martrasia Lag., Amen. nat. Espan.: 36. 1811. TYPE: not designated, the genus was based on the four species M. aurieulata Lag., M. crenata Lag., M. pubescens Lag., and M. sericea Lag., all of them nomina nuda (Harling, 1995).

Rhinactina Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 1: 139. 1807, nom. nudum.

Tostimontia S. Diaz, Revista Acad. Colomb. Ci. Exact. 25: 9. 2001. TYPE: Tostimontia gunnerifolia S. Diaz (see Observation).

Trinacte Gaertn., Fruct. sem. pl.: 415. 1791. TYPE: based on Jungia ferruginea L.f.

Etymology. In honor of the German botanist Joachim Jungius (1587-1657), professor in Hamburg.

Herbs perennial, subshrubs, or shrubs, erect to trailing. Leaves alternate to rosulate, stipulate to exstipulate; petiolate, rarely sessile; blades elliptic to suborbicular-cordate, palmately veined, rarely pinnately veined, margin entire to lobate, glabrous to pubescent. Capitulescences corymbose to paniculate; capitula pedunculate to subsessile, homogamous, discoid or radiate; receptacle paleaceous, paleae thin, scarious, embracing the central florets; involucre one- to two-seriate. Florets dimorphic or isomorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae rostrate, slightly constricted to attenuate at the apex, glabrous to pubescent; pappus of plumose, capillary bristles, occasionally barbellate to ciliate.

[FIGURES 55-58 OMITTED]

Pollen Description. Jungia stuebelii (Hieron.) Crisci (Crisci, 1971). Pollen spheroidal, medium size, tricolporate, exine Trixis type, microechinate.

Habitat and Distribution. Neotropical genus of 28 species, ranging from southwestern Mexico to northern Argentina, including the Galapagos Islands.

Species list

1. Jungia axillaris (Lag. ex DC.) Spreng., Syst. veg. 4: 301. 1827. Basionym: Dumerilia axillaris Lag. ex DC. TYPE: In Peruvia, Chile et Panamaide, s. leg., ex herb. M. Lagasca s.n. (holotype, G-DC not seen). Additional specimen examined: Peru. s.d., C. Gay s.n. (LP).

2. Jungia beckii Harling, Novon 7: 246. 1997. TYPE: Bolivia. La Paz: Prov. Nor Yungas, ca. 4 km from Chuspipata towards Puente Villa, S. Beck 18672 (holotype, LPB not seen).

3. Jungia brachyphylla Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 124. 1953. TYPE: Colombia. Dept. Valle, cordillera al N de las Brisas, Carrizales, J. Cuatrecasas 22546 (holotype F, not seen).

4. Jungia calyculata Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 125. 1953. TYPE: Colombia. Dept. Magdalena: Sierra Nevada de Santa Marta, H. H. Smith 677 (isotype not seen, US not seen, US photo at LP!).

5. Jungia coarctata Hieron., Bot. Jahrb. Syst. 29: 81. 1900. TYPE: Ecuador. Prov. Pichincha: Lloa, A. Sodiro 184 (neotype, designated by Harling, 1991: 84, W not seen). Additional specimen examined: Peru. Dept. Amazonas: Prov. Chachapoyas, Puna-urcu, E-SE of Chachapoyas, J. J. Wurdack 690 (LP).

6. Jungia crenatifolia Harling, F1. Ecuador 42: 93. 1991. TYPE: Ecuador. Prov. Azuay, Paramo de Tinajillas, ca. 10 km S of Cumbe, G. Harling 5681 (holotype S, not seen).

7. Jungia ferruginea L. f., Suppl. Pl.: 390. 1781 (1782). TYPE: Sine loc., J. C. Mutis 70 (holotype, LINN n. 1046.1 not seen). Additional specimen examined: Colombia. Bogota: south side of ravine back of Parque Nacional, H. Schiefer 452 (LP).

8. Jungia fistulosa Hieron., Bot. Jahrb. Syst. 29: 83. 1900. TYPE: Ecuador. Prov. Cotopaxi: above Pilalo, G. Harling et al. 9003 (neotype, designated by Harling, 1991: 92, GB not seen).

9. Jungia floribunda Less., Linnaea 5: 38. 1830. TYPE: Brazil: 1926, Sello s.n. (lectotype, designated by Harling, 1995: 74, K not seen). Additional specimen examined: Paraguay. Dept. La Cordillera: camino Caacupe a Tobaty, A. Schinini 4683 (LP).

10. Jungia glandulifera Harling, Acta Regiae Soc. Sci. Litt. Gothob., Bot. 4: 86. 1995. TYPE: Ecuador. Prov. Loja: Cariamanga, along road to Gonzanama, G. Harling 6014 (holotype, S not seen).

11. Jungia gracilis Harling, Novon 2: 18. 1992. TYPE: Peru. Dept. Amazonas: Rio Utcubamba valley, 31 km along road S of Tingo, L. R. M. King & L. E. Bishop 9278 (holotype, US not seen, US photo at LP!). Additional specimen examined: Brazil. Brasilia, DF: Reserva Ecologica do IBGE, B. A. S. Pereira 594 (LP).

12. Jungia hirsuta Cuatrec., Brittonia 8: 48. 1954. TYPE: Ecuador. Prov. Chimborazo: canyon of Rio Chanchan, near Huigra, W. H. Camp 3011 (isotype, NY not seen, NY photo at LP!).

13. Jungia karstenii Cuatrec., Brittonia 8: 186. 1956. TYPE: Colombia. Dept. Magdalena: Sierra Nevada de Santa Marta, San Miguel, s.d., Karsten s.n. (holotype, P not seen).

14. Jungia mitis Benoist, Bull. Soc. Bot. France 84: 634. 1938. TYPE: Ecuador. Prov. Pichincha: Valle de Lloa, slopes of Ungui, Benoist 2677 (holotype, P not seen).

15. Jungia ovata Harling, Acta Regiae Soc. Sci. Litt. Gothob., Bot. 4: 116. 1995. TYPE: Ecuador. Prov. Napo: road Quito-Baeza, 13 km E of Papallacta, Bohlin & Bohlin 676 (holotype, GB not seen).

16. Jungia paniculata (DC.) A. Gray, Proc. Amer. Acad. Arts 5: 145. 1861. Basionym: Dumerilia paniculata DC. TYPE: Peru: s.d., J. de Jussieu s.n., herb. Jussieu 9524 (holotype, P not seen). Additional specimen examined: Ecuador. Chimborazo: 9.7 km S of Chunchi, T. Stuessy & G. Nesom 5935a (LP).

17. Jungia pauciflora Rusby, Bull. New York Bot. Gard. 4: 401. 1907. TYPE: Bolivia. Dept. Cochabamba: vicinity of Cochabamba, 1891, M. Bang 2048 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Salta: Dept. Capital, San Lorenzo, L. Novara 339 (MCNS).

18. Jungia polita Griseb., Abh. Konigl. Ges. Wiss. Gottingen 24: 215. 1879. TYPE: Bolivia. Dept. Tarija: between Tarija and Santa Ana, P. G. Lorentz & G. H. E. W. Hieronymus 952 (holotype, GOET, not seen). Additional specimen examined: Argentina. Prov. Cordoba: Dept. Minas, C. Sayago 3145 (LP).

19. Jungia pringlei Greenm., Publ. Field Columbian Mus., Bot. Ser. 2: 286. 1907. TYPE: Mexico. State Michoacan: near Uruapan, C. G. Pringle 10357 (isotype, US not seen, US photo at LP!).

20. Jungia rugosa Less., Linnaea 5: 36. 1830. TYPE:, Herb. Willdenow 15992 (lectotype, designated by Harling, 1995: 52, B not seen). Additional specimen examined: Peru. Cuzco: Prov. Paucastambo, quebrada de Llulluchayoc, C. Vargas C. 4309 (LP).

21. Jungia schuerae Harling, Novon 2: 16. 1992. TYPE: Peru. Dept. Lima: Prov. Huarochiri, Matucana, E. Asplund 11101 (holotype, S not seen). Additional specimen examined: Peru. Prov. Yungay: Dept. Ancash, arriba de Yungay, A. Lopez M. et al. 7407 (LP).

22. Jungia sellowii Less., Syn. gen. Compos.: 416. 1832. TYPE: Sine loc., s.d., Sello s.n. [47] (neotype, designated by Harling, 1995: 79, P not seen). Additional specimen examined: Brazil. State Santa Catarina: Morro de Campo Alegre, Sao Francisco do Sul, P. Reitz & R. M. Klein 10958 (LP).

23. Jungia sordida J. Kost., Blumea 5: 683. 1945. TYPE: Bolivia. Abra de la Senda, T. Herzog 1839 (lectotype, designated by Harling, 1995: 64, L not seen). Additional specimen examined: Argentina. Prov. Jujuy: Laguna Yala, A. L. Cabrera & M. Marchionni 12968 (LP).

24. Jungia spectabilis D. Don, Trans. Linn. Soc. London 16: 227. 1830. TYPE: Ecuador. Prov. Guayas: Guayaquila, s.d., Tafalla s.n. (lectotype, designated by Harling, 1995: 83, BM not seen). Additional specimen examined: Ecuador. Prov. Manabi: near Jipijapa, O. Haught 3449 (LP).

25. Jungia stuebelii (Hieron.) Crisci, Bol. Soc. Argent. Bot. 13: 341. 1971. Basionym: Leucheria stuebelii Hieron. TYPE: Peru. Dept. Cajamarca: Prov. Cajamarca, Quinuamayo, entre Encafiada y Celendin, A. Sagastegui 7385 (neotype, designated by Harling, 1995: 29, LP!).

26. Jungia vitocensis Cuatrec., Anales Ci. Univ. Madrid 4: 230. 1935. TYPE: Peru. Dept. Junin: Prov. Tarma, camino de Vitoc a los altos de Palca, J. Isern 462 (holotype, MA not seen). Additional specimen examined: Peru. Prov. Contumazfi: Dept. Cajamarca, vicinity of Guzmando, A. Sagastegui 2901 (LP).

27. Jungia weberbaueri Cerrate, Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, Bot. 4: 22. 1951. TYPE: Peru. Dept. Huanuco: Prov. Pachitea, near Chaglla, A. Weberbauer 6701 (isotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cuzco: Prov. Calea, Manto, Marin 2406 (LP).

28. Jungia woodii D. J. N. Hind, Kew Bull. 59:311. 2004. TYPE: Bolivia. La Paz: Murillo, ca. 15-20 km E of summit on descent into Zongo valley, Wood 15029 (holotype, K not seen).

Observation. The monospecific genus Tostimontia (Diaz-Piedrahita, 2001) from Sierra de Santa Marta in Colombia was distinguished from Jungia by the cespitose habit, peltate leaves with foliaceous bracts, and solitary capitula. All these characters, however, are found in Jungia and thus Tostimontia becomes a synonym of Jungia.

Literature. Harling (1995, 1997), Hind (2004).

Leucheria Lag.

Amen. nat. Espan.: 32. 1811. TYPE: Leucheria hieracioides Cass. 56 of Figs. 55-58. Bertolonia DC. ex Cass., Opusc. phytol. 2: 153. 1826, nom. nudum, a name attributed to de Candolle by Cassini, but apparently never published by de Candolle (Crisci, 1976b).

Cassiopea D. Don, Trans. Linn. Soc. London 16: 215. 1830, a name frequently listed as a synonym of Leucheria, but it was actually used only as a sectional name by Don and by de Candolle (Crisci, 1974a).

Clybatis Phil., Anales Univ. Chile 41: 742. 1872. TYPE: Clybatis volkmanni Phil. [= Leucheria nutans (J. Remy) Reiche].

Chabraea DC., Ann. Mus. Natl. Hist. Nat. 19: 65, 71. 1812. TYPE: Chabraea purpurea (Vahl) DC., based on Perdicium purpureum Vahl [= Leucheria purpurea (Vahl) Hook. & Am.].

Eizaguirrea J. Remy, Fl. chil. 3: 401. 1849. TYPE: Eizaguirrea candollei J. Remy [= Leucheria floribunda DC.].

Frageria Delile ex Steud., Nomencl. bot. 1: 645. 1840, nom. nudum.

Lasiorrhiza Lag., Amen. nat. Espan.: 32. 1811, pro parte, on the basis of two species: Perdicium purpureum (Vahl) Hook. & Am. and Perdicium brasiliense L. [= Holocheilus brasiliensis (L.) Cabrera] (Crisci, 1976b).

Leucaeria DC., Ann. Mus. Natl. Hist. Nat. 19: 66. 1812, an orthographic variant of the name Leucheria.

Leuchaeria Less., Linnaea 5: 10. 1830, an orthographic variant of the name Leucheria.

Leuceria D. Don, Trans. Linn. Soc. London 16: 212. 1830, an orthographic variant of the name Leucheria.

Leukeria Endl., Erich. bot.: 249. 1841, an orthographic variant of the name Leucheria.

Maclovia DC. ex Steud., Nomencl. bot. 1: 338. 1840, nom. nudum.

Mimela Phil., Anales Univ. Chile 27: 336. 1865. TYPE: Mimela pedicularifolia Phil. [= Leucheria scrobiculata D. Don].

Ptilurus D. Don, Trans. Linn. Soc. London 16: 218. 1830. TYPE: Ptilurus daucifolius D. Don [= Leucheria daucifolia (D. Don) Crisci].

Etymology. From the Greek leukon, white, and erion, wool, because of the pubescence of these plants.

Herbs annual or perennial, caulescent to acaulescent and scapose, woolly to silky. Leaves pubescent; basal leaves rosulate to sub-rosulate; petiolate; blades linear to ovate, pinnately veined, margin entire to partite; upper leaves numerous, similar to the basal ones but reduced, or lacking; sessile. Capitulescences monocephalous, or corymbose to paniculate; capitula pedunculate, homogamous, discoid or radiate; involucre one- to five-seriate; receptacle epaleate to paleate, paleae embracing the marginal florets, with the open part facing either the center of the capitulum or the outer part of the capitulum. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex, pubescent; pappus of scabrid to plumose, capillary bristles.

Pollen Description. Leucheria (112 of Figs. 112-116) (Crisci 1974a, 1976b). Pollen spheroidal, small or medium size, tricolporate, exine Oxyphyllum type (see also Parra & Marticorena, 1972; Markgraf & D'Antoni, 1978; Wingenroth & Heusser, 1983).

Habitat and Distribution. Genus of 47 South American species distributed in the Andean region from Peru to the Islas Malvinas (Falkland Islands) and in Patagonia.

Species list

1. Leucheria achillaeifolia Hook. & Am., Comp. Bot. Mag. 2: 43. 1836. TYPE: Argentina. Prov. Santa Cruz: Port Desire, s.d., C Darwin 391 (type, probably K). Additional specimen examined: Argentina. Prov. Neuquen: Sierra Auca Mahuida, H. A. Fabris 914 (LP).

2. Leucheria amoena Phil., Linnaea 28: 714. 1856. TYPE: Chile. Prov. Linares: cordillera de Linares, Jan 1857, R. Germain s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquen: Andacollo, Arroyo Guaraco, A. L. Cabrera 11152 (LP).

3. Leucheria apiifolia Phil., Anales Univ. Chile 87:111. 1894. TYPE: Chile. Prov. Curico: s.d., Vidal s.n. (holotype, SGO not seen, SGO photo at LP!). Additional specimen examined: Chile. Prov. Curico: Dept. Curico, a orillas de la Laguna de Teno, C. Marticorena & O. Matthei 892 (LP).

4. Leucheria bridgesii Hook. & Am., Comp. Bot. Mag. 1: 36. 1835. TYPE: Cordillera, Bridges 486 (holotype, P not seen, P photo in LP!). Additional specimen examined: Chile. Prov. Santiago: Perez Caldera, C. Marticorena & O. Matthei 664 (LP).

5. Leucheria candidissima D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Chile. Sine loc., s.d., J. Gillie,; s.n. (isotype, GH not seen, GH photo at LP!). Additional specimen examined: Argentina. Prov. Neuquen: Andacollo, Cordillera del Viento, Cerro Corona, M. Jereb 17 (LP).

6. Leucheria cerberoana J. Remy, Fl. chil. 3: 386. 1847. TYPE: Chile. Coquimbo, s.d., C. Gay s.n. (type, probably P). Additional specimen examined: Chile. Prov. Coquimbo: carretera Panamericana, a 3 km al N de Puerto Oscuro, C. Marticorena et al. 1423 (LP).

7. Leucheria coerulescens J. Remy, F1. chil. 3: 382. 1849. TYPE: Chile: s.d., C. Gay s.n. (isotype, P not seen). Additional specimen examined: Argentina. Prov. Rio Negro: Isla Victoria, A. L. Cabrera 11503 (LP).

8. Leucheria congesta D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. San Juan: Dept. Calingasta, Cerro Castafio, H. A. Fabris & J. M. Marchionni 2339 (LP).

9. Leucheria cumingii Hook. & Am., Comp. Bot. Mag. l: 36. 1835. TYPE: Chile. Prov. Coquimbo: s.d., Cuming 906 (holotype, W not seen, W photo F n. 3286 at LP!). Additional specimen examined: Chile. Prov. Coquimbo: Punta Teatino, La Serena, Behn 8571 (LP).

10. Leucheria daucifolia (D. Don) Crisci, Darwiniana 20: 52. 1976. Basionym: Ptilurus daucifolius D. Don. TYPE: not localized. Additional specimen examined: Peru. Dept. Puno: Prov. Carabaya, Fauchinta, Allinccapacc, C. Vargas C. 7179 (LP).

11. Leucheria diemii Cabrera, Bol. Soc. Argent. Bot. 11: 288. 1969. TYPE: Argentina. Prov. Neuquen: Parque Nacional Nahuel Huapi, S del Cerro Fia, W. Diem 3311 (holotype, LP!). Additional specimen examined: Argentina. Prov. Neuquen: Parque Nacional Nahuel Huapi, Cerro Mesa, W. Diem 3602 (LP).

12. Leucheria eriocephala Speg., Anales Soc. Ci. Argent. 53: 31. 1902. TYPE: Argentina. Prov. Santa Cruz: Karr-aik, Lago Argentino, 1898, C. Ameghino s.n. (holotype, LP!).

13. Leucheria floribunda DC., Icon. sel. pl. 4: 88. 1839. TYPE: In excelsiis Andibus Chilensium legit cl. Gay (type, not localized). Additional specimens examined: Chile. Prov. Aconcagua: Valle Alista, al N de Rio Blanco, O. Zollner 3784 (LP); Cordillera de Santiago, Grandjot 3353 (GH*).

14. Leucheria garciana J. Remy, Fl. chil. 3: 381. 1849. TYPE: Chile. Prov. Colchagua: Colchagua, C. Gay 314 (holotype, P not seen, P photo F n. 38113 at LP!). Additional specimen examined: Chile. Prov. Curico: a orillas de la Laguna de Teno, C. Marticorena & O. Matthei 887 (LP).

15. Leucheria gayana (J. Remy) Reiche, Fl. chil. 4: 429. 1905. Basionym: Chabraea gayana J. Remy. TYPE: Chile: s.d., C. Gay s.n. (isotype, GH not seen, GH photo F n. 16045 at LP!). Additional specimen examined: Argentina. Prov. Mendoza: Tunuyan, Valle del Alto Tunuyan, A. Ruiz Leal 2063 (LP).

16. Leucheria gilliesii Hook. & Am., Comp. Bot. Mag. 1: 35. 1835. TYPE: Sine loc., s.d., s. leg. (type, probably K). Additional specimen examined: Argentina. Prov. Mendoza: San Rafael, Valle Hermoso, cerca del Refugio, Lagiglia 668 (LP).

17. Leucheria glabriuscula (Phil.) Reiche, Fl. chil. 4: 414. 1905. Basionym: Chabraea glabriuscula Phil. TYPE: Chile. Prov. Santiago: Cordillera de las Aranas, Dec 1854, Germain s.n. (isotype, LP!).

18. Leucheria glacialis (Poepp. ex Less.) Reiche, Anales Univ. Chile 116: 203. 1905. Basionym: Lasiorrhiza glacialis Poepp. ex Less. TYPE: Chile. Antuco, E. F. Poeppig 217 (holotype, P not seen, P photo F n. 20733 at LP!). Additional specimen examined: Chile. Prov. Talca: entre Laguna del Maule y Paso Pehuenches, M. Ricardi et al. 961 (LP).

19. Leucheria glandulosa D. Don, Trans. Linn. Soc. London 16: 217. 1830. TYPE: Chile. Coquimbo, s.d., Caldcleugh s.n. (type, probably G). Additional specimen examined: Chile. Prov. Valparaiso: Cuesta de Zapata, A. L. Cabrera 11445 (LP).

20. Leucheria graui Katinas, M. C. Telleria & Crisci, Novon 18: 366. 2008. TYPE: Chile. Prov. Maule, Laguna del Maule, J. Grau 2893 (holotype, LP!).

21. Leucheria hahnii Franch., Miss. Sci. Cape Horn, Bot. 5: 349. 1889. TYPE: Argentina. Prov. Tierra del Fuego: Canal del Beagle, Hahn 89 (holotype, P not seen, P photo F n. 38114 at LP!). Additional specimen examined: Argentina. Prov. Santa Cruz: Cerro Cazador, M. Gentili 417 (LP).

22. Leucheria hieracioides Cass., Dict. sci. nat. 55: 392. 1828. TYPE: Chile. s.d., d'Urville s.n. (holotype, P not seen). Additional specimen examined: Chile. Prov. Santiago, Dept. Santiago, Cuesta de Chacabuco, C. Marticorena & E. Weldt 615 (LP).

23. Leucheria integrifolia (Phil.) Crisci, Darwiniana 16: 628. 1971. Basionym: Chabraea integrifolia Phil. TYPE: Chile. Prov. Linares: Andes de Linares, s.d., R. Germain s.n. (holotype, SGO not seen). Additional specimen examined: Chile. Prov. Linares: precordillera, O. Zollner 2461 (LP).

24. Leucheria landbeckii (Phil.) Reiche, Fl. chil. 4: 423. 1905. Basionym: Chabraea landbeckii Phil. TYPE: Chile. Prov. Santiago: Cordillera de Las Aranas, 1861, Landbeck s.n. (holotype, LP!). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Carlos, Real de los Trece, Estancia Lloncha, A. Ruiz Leal 7154 (LP).

25. Leucheria leontopodioides (Kuntze) K. Schum., Just's Bot. Jahresber. 26: 378. 1900. Basionym: Lasiorrhiza leontopodioides Kuntze. TYPE: Argentina. Patagonia, s.d., Moreno & Tonini 149 (type, not localized). Additional specimen examined: Argentina. Prov. Santa Cruz: Cerro Calafate, H. Sleumer 1163 (LP).

26. Leucheria lithospermifolia (Less.) Reiche, Anales Univ. Chile 116: 208. 1905. Basionym: Lasiorrhiza lithospermifolia Less. TYPE: Chile. Prov. Bio Bio: Sierra Velluda, Andes de Antuco, E. E Poeppig 215 (holotype, P not seen). Additional specimen examined: Chile. Prov. Bio Bio: Laguna Laja, O. Zollner 5526 (LP).

27. Leucheria magna Phil., Anales Univ. Chile 87: 108. 1894. TYPE: Chile. Prov. Aysen: Valle del Rio Palena, s.d., F. Delfin s.n. (holotype, SGO, not seen, SGO photo at LP!). Additional specimen examined: Chile. Prov. Aysen: Aysen, M. Bruzzone 149 (LP).

28. Leucheria menana J. Remy, Fl. chil. 3: 387. 1849. TYPE: Chile. Prov. Coquimbo: penascos maritimos de Coquimbo, C. Gay 916 (holotype, P not seen, P photo F n. 16053 at LP!). Additional specimen examined: Chile. Prov. Coquimbo: La Serena, E. Barros 2653 (LP).

29. Leucheria millefolium Dusen & Skottsb., Kongl. Svenska Vetensk. Acad. Handl. 56: 333. 1916. TYPE: Argentina. Prov. Chubut: region del Lago General Pico, Valle Frias, Hogberg 12 (isolectotype designated by Crisci, 1976b: 48, BAF not seen). Additional specimen examined: Argentina. Prov. Rio Negro: Cerro Anecon, Ferruglio 54 (LP).

30. Leucheria multiflora Phil., Anales Univ. Chile 41: 743. 1872. TYPE: Chile. Prov. Aconcagua: Los Molles, Philippi 1126 (isotype, LP!). Additional specimen examined: Chile. Prov. Aconcagua: Quintero, El Durazno, Junge 6846 (LP).

31. Leucheria nutans (J. Remy) Reiche, Anales Univ. Chile 116: 208. 1905. Basionym: Chabraea nutans J. Remy. TYPE: Chile. Prov. Concepcion: cordillera, C. Gay 386 (holotype, P not seen). Additional specimen examined: Argentina. Prov. Rio Negro: Cerro Catedral, U. Eskuche 02-0 (LP).

32. Leucheria oligocephala J. Remy, Fl. chil. 3: 383. 1849. TYPE: Chile: s.d., C. Gay s.n. (isotype, P not seen). Additional specimen examined: Chile. Prov. Santiago: Cerros de Penalolen, Mahu 997 (LP).

33. Leucheria paniculata Poepp. ex Less., Syn. gen. Compos.: 403. 1832. TYPE: Chile. Prov. Bio Bio: Cordillera de Antuco, E. F. Poeppig 218 (isotype, NY, NY photo at LP!).

34. Leucheria papillosa Cabrera, Revista Sudamer. Bot. 3: 58. 1936. TYPE: Argentina. Prov. Rio Negro: Cerro Lopez, A. L. Cabrera & M. M. Job 339 (holotype, LP!). Additional specimen examined: Argentina. Prov. Rio Negro: region del Lago Nahuel Huapi, Paso de las Nubes, A. L. Cabrera 5905 (LP).

35. Leucheria polyclados (J. Remy) Reiche, Fl. chil. 4: 415. 1905. Basionym: Chabraea polyclados J. Remy. TYPE: Without data (holotype, P 17, 1/70 not seen). Additional specimen examined: Chile. Prov. Atacama: cercanias de Plaza, A. Krapovickas & J. Hunziker 5792 (LP).

36. Leucheria pteropogon (Griseb.) Cabrera, Darwiniana 9: 62. 1949. Basionym: Senecio pteropogon Griseb. TYPE: Argentina. Prov. Salta: alrededores del Nevado del Castillo, P. G. Lorentz & G. H. E. W. Hieronymus 101 (isotype, LP!). Additional specimen examined: Argentina. Prov. Catamarca: faldeo S de las Cumbres de la Sierra de Granadillas, Quebrada de los Potrerillos, H. Sleumer & Vervoorst 2633 (LP).

37. Leucheria purpurea (Vahl) Hook. & Am., Comp. Bot. Mag. 2: 43. 1836. Basionym: Perdicium purpureum Vahl. TYPE: Argentina. Prov. Tierra del Fuego: Magallanes, s.d., D. Commerson s.n. (lectotype, designated by Crisci, 1976b: 43, P not seen). Additional specimen examined: Argentina. Prov. Santa Cruz: Esperanza, H. Sleumer 1085 (LP).

38. Leucheria rosea Poepp. ex Less., Syn. gen. Compos.: 402. 1832. TYPE: Chile. Puente Vizcachas-La Guardia, s.d., E. F. Poeppig 574 (holotype, P not seen, P photo F n. 20732 at LP!). Additional specimen examined: Chile. Prov. Santiago: Com. Las Condes, Cerro Naranjo, A. L. Cabrera 483 (LP).

39. Leucheria runcinata D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. Mendoza: entre Las Cuevas y Cristo Redentor, H. A. Fabris 1254 (LP).

40. Leucheria salina (J. Remy) Hieron., Bol. Acad. Nac. Ci. 4: 50. 1881. Basionym: Chabraea salina J. Remy. TYPE: Chile: s.d., C. Gay s.n. (isotype, GH not seen, GH photo F n. 16058 at LP!). Additional specimen examined: Argentina. Prov. La Rioja: Dept. General Sarmiento, Quebrada La Hedionda, J. H. Hunziker 2207 (LP).

41. Leucheria scrobiculata D. Don, Philos. Mag. Ann. Chem. 11: 389. 1832. TYPE: Sine loc., s.d., J. Gillies s.n. (type, probably K). Additional specimen examined: Argentina. Prov. Mendoza: Arroyo de los Leones, A. Ruiz Leal 11743 (LP).

42. Leucheria senecioides Hook. & Am., Bot. Beechey Voy. 1: 28. 1830. TYPE: Chile. Concepcion, s.d., s. leg. (type, probably K). Additional specimen examined: Chile. Prov. Concepcion: Fundo Valencia, G. Looser 3991 (LP).

43. Leucheria suaveolens (d'Urv.) Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 538. 1897. Basionym: Perdicium suaveolens d'Urv. TYPE: not localized. Additional specimen examined: Argentina. Islas Malvinas (Falkland Islands): East Falkland, Mt. Tumbledown, W of Port Stanley, D. M. Moore 519 (LP).

44. Leucheria tenuis Less., Syn. gen. Compos.: 401. 1832. TYPE: Chile. Prov. O'Higgins: Rancagua, Monte La Leona, C. Bertero 159 (holotype, P not seen, P photo at LP!). Additional specimen examined: Chile. Prov. Aconcagua: Tabaco, O. Zollner s.n. (LP).

45. Leucheria thermarum (Phil.) Phil., Verh. Deutsch. Wiss. Verein Santiago de Chile 2: 203. 1892. Basionym: Chabraea thermarum Phil. TYPE: Chile. Prov. Nuble: Banos de Chillan, s.d., Philippi s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Rio Negro: Dept. Bariloche, Cerro Otto, J. V. Crisci 533 (LP).

46. Leucheria tomentosa (Less.) Crisci, Darwiniana 20: 114. 1976. Basionym: Lasiorrhiza tomentosa Less. TYPE: Chile. Prov. Valparaiso: Valparaiso, E. F. Poeppig 217 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Aconcagua: Ligua, E. Barros 7435 (LP).

47. Leucheria viscida (Bertero ex Colla) Crisci, Darwiniana 20: 67. 1976. Basionym: Chabraea viscida Bertero ex Colla. TYPE: Chile. Prov. O'Higgins: Rancagua, Tagua Tagua, C. Bertero 160 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Curico: Cerro Condell, E. Barros 2619 (LP).

Literature. Crisci (1976b), Katinas et al. (2008a, b)

Leunisia Phil.

Anales Univ. Chile 23: 383. 1863; Linnaea 33: 120. 1864. TYPE: Leunisia laeta Phil. 57 of Figs. 55-58.

Etymology. In honor of the German clergyman and botanist Johannis Leunis (1802-1873).

Shrubs. Leaves alternate; sessile; blades lanceolate to oblong, pinnately veined, margin entire to dentate, pubescent. Capitulescences monocephalous, terminal; capitula short-pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails papillose; style bifid, branches apically rounded, crowned by collector hairs. Cypselae attenuate at the apex, pubescent; pappus paleaceous setaceous, scabrid bristles, unequally fused at the base.

Pollen Description (Parra & Marticorena, 1972; Crisci, 1974a). Pollen prolate, medium or large size, tricolporate, exine Trixis type.

Habitat and Distribution. Monotypic genus endemic to the Andean region of central Chile.

Species list

1. Leunisia laeta Phil., Anales Univ. Chile 23: 383. 1863. TYPE: Chile. Prov. Coquimbo: Huatalame, s.d., H. Volkmann s.n. (holotype, G not seen, G photo F n. 28865 at LP!). Additional specimen examined: Chile Prov. Aconcagua: Cordon Caquis, Cerro Chache, O. Zollner 429 (LP).

Literature. Reiche (1905).

Lophopappus Rusby

Bull. Torrey Bot. Club 21: 487. 1894, emend. Cabrera, Bol. Soc. Argent. Bot. 5: 45. 1953. TYPE: Lophopappus foliosus Rusby. 58 of Figs. 55-58.

Etymology. From the Greek lophos, crest, and pappos, apical tuft of bristles on the fruit, with reference to the pappus bristles plumose at the apex.

Shrubs, stems with brachyblasts and macroblasts. Leaves alternate; sessile; blades linear to elliptic, pinnately veined, margin entire to spinose-dentate, glabrous to pubescent. Capitulescences monocephalous or in loose to dense apical cymes; capitula short-pedunculate, homogamous, discoid or disciform; receptacle epaleate; involucre multiseriate. Florets isomorphic or dimorphic, bisexual, corollas all or partly tubular-funnelform, five-lobed, or all or partly bilabiate; anther apical appendages acute, tails papillose; style bifid, branches rounded at the apex, dorsally papillose. Cypselae attenuate to slightly constricted at the apex, pubescent; pappus of scabrid, capillary bristles, plumose at the apex.

Pollen Description. Lophopappus (113 of Figs. 112-116) (Telleria et al., 2003). Pollen subprolate to prolate-spheroidal, medium to large size, tricolporate, exine Trixis type, Proustia type, or both types in a single specimen, with polar caps only in L. berberidifolius Cabrera and L. peruvianus Cabrera (see also Parra & Marticorena, 1972; Crisci, 1974a).

Habitat and Distribution. Genus of six species distributed in the Andes from Peru to Argentina.

Species list

1. Lophopappus berberidifolius Cabrera, Bol. Soc. Argent. Bot. 5: 48. 1953. TYPE: Peru. Dept. Huancavelica: Sachahuacta, 7 km al SO de Conaica, O. Tovar 961 (holotype, LP!).

2. Lophopappus blakei Cabrera, Bol. Soc. Argent. Bot. 5: 48. 1953, nom. novum. Basionym: Proustia cuneata S. F. Blake. TYPE: Peru. Dept. Cuzco: Ollantaytambo, O. F. Cook & G. B. Gilbert 538 (holotype, US not seen, US photo at LP!). Additional specimen examined: Peru. Dept. Cuzco: desvio km 82, Estancia Piscacucho, A. L. Cabrera & H. A. Fabris 13462 (LP).

3. Lophopappus cuneatus R. E. Fr., Ark. Bot. 5: 29. 1906. TYPE: Bolivia. Quebrada Honda, pr. Tarija, R. E. Fries 1045 (isotype, US not seen, US photo at LP!). Additional specimen examined: Argentina. Jujuy: Dept. Cochinoca, Abra Pampa, Cerro Huancar, A. L. Cabrera 15425 (LP).

4. Lophopappus foliosus Rusby, Bull. Torrey Bot. Club 21: 487. 1894. TYPE: Bolivia. Dept. La Paz: vicinity of La Paz, M. Bang 66 (isosyntype, US not seen, US photo at LP!). Additional specimen examined: Bolivia. Dept. La Paz: Valle de la Luna, A. L. Cabrera 2570 (LP).

5. Lophopappus peruvianus Cabrera, Bol. Soc. Argent. Bot. 5: 50. 1953. TYPE: Peru. Dept. Ancash: Prov. Bolognesi, cerca de Aquia, E. Cerrate 1552 (holotype, LP!).

6. Lophopappus tarapacanus (Phil.) Cabrera, Revista Mus. La Plata, Secc. Bot. 12: 157. 1971. Basionym: Gochnatia tarapacana Phil. TYPE: Chile. Prov. Tarapaca: Sotoca, s.d., Philippi s.n. (type, probably SGO). Additional specimen examined: Chile. Prov. Tarapaca: a+- 10 km volviendo de Tranque de Caritaya, M. Ricardi & C. Marticorena 25604 (LP*).

Observation. Lophopappus has been also placed in the subtribes Mutisiinae and Gochnatiinae (Cabrera, 1961), because of the tubular florets and rounded, dorsally papillose style branches. The type of exine stratification in pollen of Lophopappus (Crisci, 1974a; Telleria et al., 2003) clearly indicates that this genus belongs to the subtribe Nassauviinae (= tribe Nassauvieae). Style and corolla features link Lophopappus to the genus Proustia, and some authors (Ferreyra, 1995) consider that Lophopappus should be merged with Proustia.

Literature. Cabrera (1953a, 1971b).

Macrachaenium Hook. f

Fl. antarct. 2: 321. 1847. TYPE: Macrachaenium gracile Hook. f. 59 of Figs. 59-62.

[FIGURES 59-62 OMITTED]

Etymology. From the Greek makros, big, and achaena or achaenium, the typical fruit of Asteraceae, in reference to the size of the cypselae.

Herbs perennial, acaulescent, with one or more scapes. Leaves pinnately veined, pubescent; basal leaves rosulate, base inflated and vaginate; pseudopetiolate; blades broadly elliptic, margin runcinate to pinnately partite; upper leaves similar to the basal ones but reduced, or lacking; sessile to decurrent. Capitulescences mono cephalous, on scapes; capitula homogamous, discoid or radiate; receptacle epaleate; involucre two- to three-seriate. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages truncate to acute, tails smooth; style bifid, branches rounded at the apex, dorsally papillose. Cypselae attenuate at the apex, glabrous; pappus of plumose, capillary bristles.

Pollen Description. Pollen subprolate, elliptic in equatorial view, circular in polar view, large size, P x E = (55 - 63 x 42 - 49) [micro]m. Tricolporate, colpi long with psilate membrane, mesoaperturate. Exine Trixis type, 6-8 [micro]m at the equator slightly slender at the poles, with polar elevations. Nexine 1.2-2 [micro]m thick. (see also Parra & Marticorena, 1972; Crisci, 1974a; Telleria et al., 2003).

Habitat and Distribution. Monotypic genus with two varieties endemic to Nothofagus forests in southern Argentina and Chile.

Species list

Macrachaenium gracile Hook. f., Fl. antarct. 2: 321. 1847. TYPE: Argentina. Prov. Rio Negro: Dept. Bariloche, Lago Nahuel Huapi, Laguna Frias, cerro Riggi, A. L. Cabrera 6048 (holotype of M. gracile var. radiatum, LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: R. Goodall 283 (LP).

Observation. Macrachaenium has sometimes been placed in the subtribe Mutisiinae (Cabrera, 1961) because of its rounded, dorsally papillose style branches. The type of exine stratification in the pollen of Macrachaenium (Crisci, 1974a; Telleria et al., 2003) clearly indicates the position of this genus within the Nassauviinae (= tribe Nassauvieae).

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

Marticorenia Crisci

J. Arnold Arbor. 55: 38. 1974. TYPE: Leuceriafoliosa Phil. [= Marticorenia foliosa (Phil.) Crisci]. 60 of Figs. 59-62.

Etymology. In honor of the Chilean botanist Clodomiro Marticorena (b. 1929) who made important contributions to the knowledge of the flora of Chile.

Shrubs. Leaves alternate; sessile, decurrent to clasping; blades ovate-elliptic, pinnately veined, margin lobulate, pubescent. Capitulescences a dichotomous cyme; capitula pedunculate, homogamous, discoid; receptacle paleaceous, paleae embracing all the florets; involucre one- to two-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex, pubescent; pappus of plumose, capillary bristles.

Pollen Description. Pollen subprolate to prolate, medium size, tricolporate, exine Trixis type (see Crisci, 1974b).

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

Species list

1. Marticorenia foliosa (Phil.) Crisci, J. Arnold Arbor. 55: 40. 1974. Basionym: Leuceria foliosa Phil. TYPE: Chile. Prov. Santiago: Las Aranas, Cajon del Mapocho, R. Philippi 1132 (neotype, designated by Crisci, 1974b: 42, LP!). Additional specimen examined: Chile. Prov. Aconcagua, Valle Castro, cerca de Rio Blanco, O. Zollner 2994 (LP).

Literature. Crisci (1974b).

Moscharia Ruiz & Pav., nom. cons.

Fl. peruv, prodr.: 91. 1794, non Forssk., 1775. TYPE: Moscharia pinnatifida Ruiz & Pay. 61 of Figs. 59-62.

Moscaria Pers., Syn. pl. 2: 379. 1807, an orthographic variant of the name Moscharia.

Moschifera Molina, Sag. stor. nat. Chili: 294. 1810, an orthographic variant of the name Moscharia.

Mosigia Spreng., Syst. veg. 3: 366. 1828, nom. superfluum.

Gastrocarpha D. Don, Brit. Fl. Gard. 3: 229. 1827-1829. TYPE: Gastrocarpha runcinata D. Don [= Moscharia pinnatifida Ruiz & Pav.].

Etymology. From the Greek moschos, musky, because of the strong musky fragrance of the plant.

Herbs annual. Leaves pinnately veined, glabrous to pubescent; basal leaves subrosulate; petiolate; blades oblanceolate, pinnately parted; upper leaves petiolate to amplexicaul; blades lanceolate, margin entire to lobed. Capitulescences a capituliform pseudocephalium; capitula subsessile, homogamous, discoid; outer capitula of the pseudocephalia with two florets, the involucre of two phyllaries, the outer phyllary keeled, embracing the two florets, the inner phyllary not embracing and placed between the two florets and interpreted as a palea; central capitulum one- to five-flowered, involucre uniseriate; receptacle paleate or epaleate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, pubescent; pappus of very short, crown-like, scabrid bristles, narrowly paleaceous, or pappus absent in the outer cypselae.

Pollen Description. Moscharia (Parra & Marticorena, 1972). Pollen spheroidalprolate, medium size, tricolporate, exine Oxyphyllum type (see also Crisci, 1974a). Habitat and Distribution. Genus of two species endemic to the biogeographic region of Central Chile.

Species list

1. Moscharia pinnatifida Ruiz & Pav., Syst. veg. fl. peruv, chil. 1: 186. 1798. TYPE: Habitat in Regni Chilensis, aridis arenosis et petrosis locis ad Aconcaguae, Quillotae et Rancaguae provincias, et copiose versus S. Jacobi Chilensis urbem (type, not localized). Additional specimen examined: Chile. Region Metropolitana: Reserva Nacional Rio Clarillo, M. Bonifacino 1321 (LP).

2. Moscharia solbrigii Crisci, Contr. Gray Herb. 205: 172. 1974. TYPE: Chile. Prov. Coquimbo: Dept. Ovalle, south end Fray Jorge forest, J. L. Morrison & C. R. Worth 16441 (holotype, GH not seen). Additional specimen examined: Chile. Prov. Coquimbo: Dept. Ovalle, Parque Nacional Fray Jorge, C. Marticorena et al. 521 (LP).

Literature. Crisci (1974c), Hellwig (1985), Katinas and Crisci (2000), Katinas et al. (2008a).

Nassauvia Comm. ex Juss.

Gen. pl.: 175. 1789. TYPE: Nassauvia magellanica J. F. Gmel. 62 of Figs. 59-62.

Acanthophyllum Hook. & Am., Comp. Bot. Mag.: 37. 1835. TYPE: Acanthophyllum axillare (Lag. ex Lindl.) Hook. & Am., based on Triptilion axillare Lag. ex Lindl. [= Nassauvia axillaris (Lag. ex Lindl.) D. Don].

Caloptilium Lag., Amen. nat. Espan.: 34. 1811. [= Nassauvia sect. Caloptilium (Lag.) Benth., in Benth. & Hook., Gen. p1. 2: 503. 1873]. TYPE: Caloptilium lagascae (D. Don) Hook. & Am., based on Sphaerocephalus lagascae D. Don [= Nassauvia lagascae (D. Don) F. Meigen var. lagascae].

Mastigophorus Cass., Dict. sci. nat. 34 222. 1825. [= Nassauvia sect. Mastigophorus (Cass.) DC., Prodr. 7: 50. 1838]. TYPE: Mastigophorus gaudichaudii Cass. [= Nassauvia gaudichaudii (Cass.) Cass. ex Gaudich.].

Nassovia Batsch, Tab. affin, regni veg. 251:1802, an orthographic variant of the name Nassauvia.

Panargyrum DC., Ann. Mus. Natl. Hist. Nat. 19: 9. 1812, an orthographic variant of the name Panargyrus established by de Candolle and used by most researchers, including Cabrera (1982).

Panargyrus Lag., Amen. nat. Espan.: 33. 1811. [= Nassauvia sect. Panargyrum (Lag.) Wedd., Chlor. andina 1: 52. 1855]. TYPE: Panargyrum lagascae DC. [= Nassauvia darwinii (Hook. & Am.) O. Hoffm. & Dusen].

Pentanthus Less., Syn. gen. Compos.: 398. 1832. TYPE: Pentanthus aculeatus Less. [= Nassauvia aculeata (Less.) Poepp. & Endl. var. aculeata].

Piptostemma Spach, Hist. nat. veg. 10: 34. 1841, a name attributed to Don by Spach and considered a synonym of Nassauvia, but never published by Don. The name was retaken by de Candolle as a section of Nassauvia (Crisci, 1974a; Cabrera, 1982).

Portalesia Meyen, Reise Chile l: 316. 1834. TYPE: Portalesia procumbens Meyen [= Nassauvia lagascae (D. Don) F. Meigen vat. lagascae].

Sphaerocephalus Lag. ex DC., Ann. Mus. Natl. Hist. Nat. 19: 67. 1812. TYPE: Sphaerocephalus lagascae D. Don. [= Nassauvia lagascae (D. Don) F. Meigen var. lagascae].

Strongyloma DC., Prodr. 7: 52. 1838. [= Nassauvia subgen. Strongyloma (DC.) Cabrera, Darwiniana 24: 364. 1982]. TYPE: Strongyloma axillare (Lag. ex Lindl.) DC., based on Triptilion axillare Lag. ex Lindl. [= Nassauvia axillaris (Lag. ex Lindl.) D. Don].

Strongylomopsis Speg., Comun. Mus. Nac. Buenos Aires 1: 135. 1899. TYPE: Strongylomopsis fuegiana Speg. [= Nassauvia fuegiana (Speg.) Cabrera].

Triachne Cass., Bull. Sci. Soc. Philom. Paris: 11. 1818. TYPE: Triachne pygmaea Cass. [= Nassauvia pygmaea (Cass.) Hook. var. pygmaea].

Trianthus Hook. f., F1. antarct. 2: 320. 1847. TYPE: Trianthus ulicinus Hook. f. [= Nassauvia ulicina (Hook. f.) Macloskie].

Etymology. In honor of the prince of Nassau, who took part in the trip of Bougainville around the world in 1766.

Herbs perennial, subshrubs or shrubs, sometimes cushion-like, frequently with brachyblasts and macroblasts. Leaves imbricate; sessile to clasping; blades linearsubulate to ovate, several parallel veins reduced in number in the linear blades, margin entire to partite, spiny to unarmed, sometimes revolute, glabrous to pubescent. Capitulescences monocephalous, racemose, spicate, to glomerulose or pseudocephalium; capitula sessile to pedunculate, homogamous, discoid or radiate, five-flowered, occasionally one- to four-flowered; receptacle epaleate; involucre two-seriate. Florets isomorphic, bisexual, corolla bilabiate, outer lip expanded or not; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, glabrous to pubescent; pappus of bristles filiform to narrowly paleaceous, laciniate to plumose, rarely crown-like.

Pollen Description. Nassauvia (Parra & Marticorena, 1972; Crisci, 1974a). Pollen spheroidal, prolate to suboblate, small to medium size, tricolporate, exine Oxyphyllum type (see also Markgraf & D'Antoni, 1978; Wingenroth & Heusser, 1983).

Habitat and Distribution. Genus of 38 South American species distributed in the Andes and Patagonia, from southern Bolivia to the Islas Malvinas (Falkland Islands).

Species list

1. Nassauvia aculeata (Less.) Poepp. & Endl., Nov. gen. sp. pl. 1: 12. 1835. Basionym: Pentanthus aculeatus Less. TYPE: Chile. Andes de Antuco, Sierra Velluda, E. F. Poeppig 227 (lectotype, designated by Cabrera, 1982: 357, P not seen). Additional specimen examined: Argentina. Prov. Neuquen: Pino Hachado, A. L. Cabrera & J. V. Crisci 19113 (LP).

2. Nassauvia ameghinoi Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 542. 1897. TYPE: Argentina. Prov. Santa Cruz: San Jorge, Feb 1896, C. Ameghino s.n. (holotype, LP!).

3. Nassauvia argentea Phil., Anales Univ. Chile 87: 82. 1894. TYPE: Chile. Prov. Valdivia: Huahim, Jan 1887, O. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Argentina. Prov. Neuquen: San Martin de los Andes, Cerro Chapelco, A. L. Cabrera 19722 (LP).

4. Nassauvia argyrophylla Speg. ex Cabrera, Notas Mus. La Plata, Bot. 4:160 1939. TYPE: Argentina. Prov. Chubut: Rio Corcovado, 1/15 March 1901, N. Illin s.n. (holotype, LP!).

5. Nassauvia axillaris (Lag. ex Lindl.) D. Don, Philos. Mag. Ann. Chem. 11 : 390. 1832. Basionym: Triptilion axillare Lag. ex Lindl. TYPE: Chile, s.d., s. leg. (type, not localized). Additional specimen examined: Argentina. Prov. Mendoza: Dept. San Carlos, Quebrada del Paso de la Cruz de Piedra, A. Ruiz Leal 11765 (LP).

6. Nassauvia chubutensis Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 615. 1897. TYPE: Argentina. Prov. Chubut: Teka-choique, 1889, C. Moyano s.n. (holotype, LP!).

7. Nassauvia coronipappa Arroyo & Martic., Brittonia 40:332.1988. TYPE: Chile. Ultima Esperanza: Cerro Donoso, Rio de las Chinas valley, M. T. K. Arroyo et al. 87314 (holotype, CONC!).

8. Nassauvia cumingii Hook. & Am., Comp. Bot. Mag. 1: 37. 1835. TYPE: Cordillera, s.d., H. Cuming 237 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Mendoza: Las Heras, Las Cuevas, refugio militar General Lamadrid, O. Boelcke et al. 9818 (LP).

9. Nassauvia darwinii (Hook. & Am.) O. Hoffm. & Dusen, Wiss. Erg. Schwed. Exp. Magellansl. 3: 112. 1900. Basionym: Panargyrum darwinii Hook. & Am. TYPE: Argentina. Prov. Santa Cruz: Port Desire, C. Darwin 313 (lectotype, designated by Cabrera, 1982:351, K not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Punta Maria flat, between Viamonte and Rio Grande, R. N. Goodall 316 (LP).

10. Nassauvia dentata Griseb., Abh. Konigl. Ges. Wiss. Gottingen 6: 125. 1854. TYPE: Sine loc., R. Philippi 125 (isotypes, K not seen, P not seen). Additional specimen examined: Argentina. Prov. Neuquen: Dept. Lacar, Hua-Hum, Cerro Malo, J. H. Hunziker 6996 (LP).

11. Nassauvia digitata Wedd., Chlor. andina 1: 48. 1855. TYPE: Chile: s.d., C. Gay s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Neuquen: Dept. Norquin, Termas de Copahue, A. L. Cabrera 6195 (LP).

12. Nassauvia dusenii O. Hoffm., Rep. Princeton Univ. Exp. Patagonia, Botany 8: 879. 1906. TYPE: Chile. Patagonia occidentalis, Rio Aysen, P. K. Dusen 584 (isotype, B not seen, B photo F n. 16032 at LP!). Additional specimen examined: Argentina. Prov. Chubut: Dept. Futaleufu, Esquel, La Hoya, A. L. Cabrera et al. 23132 (LP).

13. Nassauvia fuegiana (Speg.) Cabrera, Physis (Buenos Aires) 10: 285. 1931. Basionym: Strongylomopsis fuegiana Speg. TYPE: Argentina. Prov. Tierra del Fuego: Golfo San Sebastian, Jan 1896, O. Mauri s.n. (holotype, LP!).

14. Nassauvia gaudichaudii (Cass.) Cass. ex Gaudich., Ann. Sci. Nat. (Paris) 5: 103. 1825. Basionym: Mastigophorus gaudichaudii Cass. TYPE: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), April 1825, C. Gaudichaud s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), East Falkland, Eliza Cove, ca. 1 1/2 miles S of Port Stanley, D. M. Moore 559 (LP).

15. Nassauvia glomerata (D. Don) Wedd., Chlor. andina 1: 53. 1855. Basionym: Panargyrum glomeratum D. Don. TYPE: Chile. Andes of Chile, Valle de los Ciegos, s.d., J. Gillies s.n. (lectotype, designated by Cabrera, 1982:361, K not seen). Additional specimen examined: Argentina. Prov. Mendoza: Malalhue, Alto Valle del Atuel, Quebrada del Arroyo Niehd, A. Ruiz Leal & F. Roig 15677 (LP).

16. Nassauvia glomerulosa (Lag. ex Lindl.) D. Don, Philos. Mag. Ann. Chem. 11: 390. 1832. Basionym: Triptilion glomerulosum Lag. ex Lindl. TYPE: Argentina. Cordillera del Portillo, 1813, M. Nee s.n. (isotype, G not seen, G photo F n. 8235 at LP!). Additional specimen examined: Argentina. Prov. Chubut: Tecka, Estancia La Blanche, A. Soriano 2216 (LP).

17. Nassauvia hillii Cabrera, Notas Mus. La Plata, Bot. 22: 161. 1939. TYPE: Argentina. Prov. Neuquan: Dept. Alumine, Pulmari, cerca de Piedra Gaucha, H. F. Comber 356 (isotype, LP!).

18. Nassauvia juniperina Skottsb., Kongl. Svenska Vetensk. Acad. Handl. 56: 328. 1916. TYPE: not localized. Additional specimen examined: Argentina. Prov. Chubut: a 35 km de Esquel, A. Ruiz Leal 23964 (LP).

19. Nassauvia lagascae (D. Don) F. Meigen, Bot. Jahrb. Syst. 18: 445. 1894. Basionym: Sphaerocephalus lagascae D. Don. TYPE: Chile. Andes of Chile, ascent to Los Peuquenes, J. Gillies 33 (holotype, E not seen). Additional specimen examined: Argentina. Prov. Mendoza: San Rafael, Distrito Sosneao, Alto Valle del Atuel, Laguna Atuel, A. Ruiz Leal 16882 (LP).

20. Nassauvia latissima Skottsb., Kongl. Svenska Vetensk. Acad. Handl. 56: 330. 1916. TYPE: Argentina. Prov. Tierra del Fuego: Isla de los Estados, C. Skottsberg 273 (isotype, SI not seen, SI photo at LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Dept. Bahia Tetis, Estancia Moat, D. M. Moore 1649 (LP).

21. Nassauvia looseri Cabrera, Darwiniana 24: 363. 1982, nom. novum. Basionym: Panargyrum heterophyllum Phil. TYPE: Chile. Cordillera de Santiago, Las Aranas, Jan 1860, G. Landbeck s.n. (isotype, LP!). Additional specimen examined: Chile. Prov. Santiago: Refugio de La Parva, a 4 km de Farellones, Cordillera de Las Condes, G. Looser 5755 (LP).

22. Nassauvia maeviae Cabrera, Darwiniana 24: 367. 1982. TYPE: Argentina. Prov. Santa Cruz: Dept. Guer Aike, laguna La Leona, 51[degrees]31'' S, 69[degrees]47" W, 7 Dec 1975, M. N. Correa s.n. (holotype, SI!). Additional specimen examined: Argentina. Prov. Santa Cruz: Dept. Guer Aike, a 14 km desvio de ruta nacional 3 hacia la Estancia Guakenen Aike, L. Katinas 24 (LP).

23. Nassauvia magellaniea J. F. Gruel., Syst. nat.: 1281. 179l. TYPE: Chile or Argentina. Magallania, s.d., D. Commerson s.n. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Usuahia, Laguna Azul, M. N. Correa & R. L. Perez Moreau 1920 (LP).

24. Nassauvia pentacaenoides Speg., Anales Soc. Ci. Argent. 53: 34. 1902. TYPE: Argentina. Prov. Santa Cruz: Karr-aik, Lago Argentino, March 1898, C. Ameghino s.n. (holotype, LP!).

25. Nassauvia pinnigera D. Don, Philos. Mag. Ann. Chem. 11: 390. 1832. TYPE: Argentina. Prov. Mendoza: Andes of Mendoza, ascent to Planchon, J. Gillies 36 (holotype, E not seen). Additional specimen examined: Argentina. Prov. Neuquen: Cordillera del Viento, La Corona, F. Pastore 83 (LP).

26. Nassauvia planifolia Wedd., Chlor. andina 1: 48. 1855. TYPE: Argentina. E side of Andes, Bridges 1148 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Mendoza: San Rafael, Valle Hermoso, subida hacia la Vega de los Piuquenes, A. Ruiz Leal 27357 (LP).

27. Nassauviapuleherrima Cabrera, Notas Mus. La Plata, Bot. 22: 157. 1939. TYPE: Argentina. Prov. Rio Negro: Dept. Bariloche, Cerro Catedral, s.d., N. J. J. Neumeyer s.n. (holotype, LP!).

28. Nassauvia pygmaea (Cass.) Hook. f., Fl. antarct. 2: 319. 1847. Basionym: Triachne pygmaea Cass. TYPE: Argentina. Patagonia australis, Cerro Toro, 1826? 1896?, O. Nordenskjold s.n. (isotype, K not seen). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Isla de los Estados, Puerto Cook, M. A. Torres 1199 (LP).

29. Nassauvia pyramidalis Meyen, Reise Chile 1: 356. 1834. TYPE: Chile. Cordillera de Santiago, Valle del Yeso, Jan 1868, R. Philippi s.n. (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquen: Dept. Minas, Cordillera del Viento, cruzando de Tricao Malal al Cajon de Butalo, O. Boelcke 11598 (LP).

30. Nassauvia ramosissima DC., Prodr. 7: 49. 1838. TYPE: Chile. Prov. Colchagua: Talcaregue, 1831, C. Gay s.n. (holotype, G not seen, G photo at LP!).

31. Nassauvia revoluta D. Don, Philos. Mag. Ann. Chem. 11: 390. 1832. TYPE: Argentina. Prov. Mendoza: Andes of Mendoza, ascent to Planchon, J. Gillies 35 (holotype, E not seen, E photo at LP!). Additional specimen examined: Argentina. Prov. Neuquen: Dept. Norquin, Termas de Copahue, A. L. Cabrera 6282 (LP).

32. Nassauvia ruizii Cabrera, Notas Mus. La Plata, Bot. 22: 162. 1939. TYPE: Argentina. Prov. Mendoza: Dept. Tunuyan, Rincones del Cerro Morado, A. Ruiz Leal 3183 (holotype, LP!).

33. Nassauvia sceptrum Dusen, Rep. Princeton Univ. Exp. Patagonia, Botany, Suppl.: 286. 1914. TYPE: Argentina. Prov. Santa Cruz: Lago Viedma, P. Dusen 5887 (isotype, SI not seen, SI photo at LP!).

34. Nassauvia serpens d'Urv., Mem. Soc. Linn. Paris 4: 610. 1825. TYPE: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), 1825, d'Urville s.n. (holotype, P not seen; isotype, B not seen, B photo F n. 16039 at LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Islas Malvinas (Falkland Islands), East Falkland, Mt. Urbourne, D. M. Moore 583 (LP).

35. Nassauvia sprengelioides DC., Prodr. 7: 49. 1838. TYPE: Argentina. Prov. Mendoza: de la Cordillera del Planchon, s.d., M. Nee s.n. (holotype, G not seen). Additional specimen examined: Chile. Prov. Curico: Teno, La Montana, E. Barros 3801 (LP).

36. Nassauvia sublobata Cabrera, Darwiniana 24: 323. 1982. TYPE: Argentina. Prov. Neuquen: cordon del Chapelco, nacientes del arroyo Chapelco Grande, al N del pequeno lago La Kika, M. Gentili & P. Gentili 754 (isotype, LP!). Additional specimen examined: Argentina. Prov. Mendoza: Alto Valle de Calmuco, A. Burkart et al. 14401 (LP).

37. Nassauvia ulicina (Hook. f.) Macloskie, Rep. Princeton Univ. Exp. Patagonia, Botany 8: 885. 1906. Basionym: Trianthus ulicinus Hook. f. TYPE: Argentina. Prov. Santa Cruz: Cape Fairweather, s.d., Capt. King s.n. (holotype, K not seen). Additional specimen examined: Argentina. Prov. Chubut: Dept. Rawson, Rawson, Punta Norte, M. M. Job 3009 (LP).

38. Nassauvia uniflora (D. Don) Hauman, Anales Soc. Ci. Argent. 86: 327. 1918. Basionym: Panargyrum unoqorum D. Don. TYPE: Argentina. Prov. Mendoza: Andes of Mendoza, Paramillo de las Cuevas, s.d., J. Gillies s.n. (holotype, E not seen). Additional specimen examined: Argentina. Prov. Mendoza: Las Cuevas, Quebrada Benjamin Matienzo, R. Perez Moreau 12753 (LP).

Observation. Species of Nassauvia are dominant elements in vast areas of Patagonia.

Literature. Cabrera (1982), Kalin Arroyo and Marticorena (1988), Freire et al. (1993), Katinas at al. (2008a).

Oxyphyllum Phil.

Fl. atacam.: 28. 1860. TYPE: Oxyphyllum ulicinum Phil. 63 of Figs. 63-66.

Etymology. From the Greek oxys, sharp, pungent, and phyllon, leaf, referring to the spiny, sharp-pointed leaves.

Shrubs. Leaves imbricate; sessile, clasping; blades linear to obovate, entire, pinnatisect to reduced to three-parted spines, pinnately veined, margin entire, involute, pubescent. Capitulescences densely corymbose; capitula short-pedunculate, homogamous, discoid; receptacle paleate, paleae embracing the marginal florets; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate to slightly constricted at the apex, pubescent; pappus with scabrid to plumose, capillary bristles.

Pollen Description. Pollen spheroidal, with polar caps in equatorial view, circular in polar view, medium size, P x E = (28 - 33 x 26 - 30) [micro]m. Tricolporate, colpi long with scarcely microgranulate membrane. Exine Oxyphyllum type, scabrate, 3-4 [micro]m thick, thickened at the poles. Nexine 0.5-1 [micro]m thick (see also Parra & Marticorena, 1972; Crisci, 1974a; Telleria et al., 2003).

Monotypic genus endemic to the desert areas of northern Chile.

[FIGURES 63-66 OMITTED]

Species list

1. Oxyphyllum ulicinum Phil., F1. atacam.: 173. 1860. TYPE: Chile. Prov. Antofagasta: region del litoral, en Cachinal de la Costa, Paposo, etc., R. Philippi s.n. (type, probably SGO). Additional specimen examined: Hueso Parado, cerca de Taltal, O. Zo1lner 3892 (LP).

Literature. Reiche (1905).

Panphalea Lag.

Amen. nat. Espan.: 34. 1811. TYPE: Panphalea commersonii Cass. 64 of Figs. 63-66. Ceratolepis Cass., Bull. Sci. Soc. Philom. Paris: 111. 1819, nom. nudum. Pamphalea DC. Ann. Mus. Natl. Hist. Nat. 19: 68. 1812, an orthograhic variant of the name Panphalea (see Observation).

Etymology. From Greek pan, all, whole, and phalos, shiny, bright, probably in reference to the white florets or to the leaves.

Herbs annual or perennial, caulescent. Leaves glabrous to pubescent; basal leaves rosulate; petiolate, sessile to clasping; blades linear, lanceolate, ovate to orbicular, palmately to pinnately veined, margin entire, lyrate to pinnatisect; upper leaves alternate, reduced; clasping. Capitulescences racemose to corymbose; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre one- to three-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, pubescent; pappus absent.

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

Habitat and Distribution. Genus of nine South American species distributed in humid places of southern Brazil, eastern Paraguay, Uruguay, and northeastern Argentina.

Species list

1. Panphalea araucariophila Cabrera, Notas Mus. La Plata, Bot. 16: 232. 1953. TYPE: Brazil. State Rio Grande do Sul: Taimbesinho, p. Sao Francisco de Paula, B. Rambo 49335 (holotype, LP!).

2. Panphalea bupleurifolia Less., Linnaea 5: 8. 1830. TYPE: Brazil. Brasilia meridionalis, s.d., Sello s.n. (holotype, B not seen, B photo F n. 16094 at LP!). Additional specimen examined: Argentina. Prov. Buenos Aires: arroyo Juan Blanco-Ruta 11, J. V. Crisci 542 (LP).

3. Panphalea cardaminfolia Less., Linnaea 5: 9. 1830. TYPE: Brazil. Brasilia meridionalis. Uruguay. Montevideo, Sello 1553 (holotype B, not seen, B photo F n. 16095 at LP!). Additional specimen examined: Brazil. State Santa Catarina: Mun. Lajes, 6 km W of Capao Alto, S. B. Smith & R. M. Klein 11341 (LP).

4. Panphalea commersonii Cass., Bull. Sci. Soc. Philom. Paris: 111. 1819. TYPE: Uruguay. Pres de Montevideo, s.d., s. leg., herb. Jussieu s.n. (holotype, P not seen). Additional specimen examined: Uruguay. Dept. Montevideo: Montevideo, alrededores del cerro de Montevideo, J. Chebataroff 232 (LP).

5. Panphalea heterophylla Less., Linnaea 5: 8. 1830. TYPE: Brazil. Brasilia meridionalis. Uruguay. Montevideo, Sello 1328 (holotype, B not seen, B photo F n. 16096 at LP!). Additional specimen examined: Argentina. Prov. Entre Rios: Dept. Concordia, Salto Grande, A. L. Cabrera & A. Sagastegui 19295 (LP).

6. Panphalea maxima Less., Linnaea 5: 9. 1830. TYPE: Brazil. Brasilia meridionalis. Uruguay. Montevideo, Sello 1365 (holotype, B not seen, B photo F n. 16097 at LP!). Additional specimen examined: Uruguay. Dept. Salto: Paysandfi, Cueva del Tigre, J. Chebataroff 7001 (LP).

7. Panphalea missionum Cabrera, Notas Mus. La Plata, Bot. 16: 235. 1953. TYPE: Argentina. Prov. Misiones: Dept. Candelaria, Loreto, J. E. Montes 2515 (holotype, LP!). Additional specimen examined: Argentina. Prov. Misiones: Campo de las Cuyas, Jan 1907, C. L. Spegazzini s.n. (LP).

8. Panphalea ramboi Cabrera, Notas Mus. La Plata, Bot. 16: 229. 1953. TYPE: Brazil. State Rio Grande do Suh Taimbesinho, p. Sao Francisco de Paula, B. Rambo SJ 49420 (holotype, LP!).

9. Panphalea smithii Cabrera, Bol. Soc. Argent. Bot. 7: 200. 1959. TYPE: Brazil. Santa Catarina: Mun. Bom Retiro, bog, Fazenda Campo dos Padres, Campo dos Padres, S. B. Smith & P. Reitz 10417 (holotype, LP!).

Observation. It can be found in the literature the use of the name Pamphalea instead of the original name Panphalea. The basic problem is that the final letter of "pan" is a dental. In Greek, when a dental (the final "n" of "pan") comes up against a labial, it changes to a labial ("m"), hence "pam." The International Code of Botanical Nomenclature (Greuter et al., 2000) in article 60.1 suggests maintaining the original spelling, not imposing linguistically correct spellings. Therefore the original spelling, Panphalea, must be maintained. Those who want to use "pam" must propose conservation, documenting overwhelming usage of "pam" (Dan Nicolson, Smithsonian Institution, personal communication).

Literature. Cabrera (1953b, 1959a), Katinas and Crisci (2008).

Perezia Lag.

Amen. nat. Espan.: 31. 1811. TYPE: Perdicium magellanicum L. f. [= Perezia magellanica (L. f.) Lag.]. 65 of Figs. 63-66.

Calorezia Panero, Phytologia 89: 199. 2007. TYPE: Calorezia nutans (Less.) Panero [= Perezia nutans Less.].

Clarionea Lag. ex DC., non Cass., Ann. Mus. Natl. Hist. Nat. 19: 65. 1812. TYPE: Clarionea magellanica Lag. ex DC. [= Perezia magellanica (L. f) Lag.].

Clarionia D. Don, Trans. Linn. Soc. London 16: 204. 1830, an orthographic variant of the name Clarionea.

Clarionema Phil., Linnaea 28: 717. 1858. TYPE: Clarionema humilis Phil. [= Perezia pedicularidifolia Less.].

Drozia Cass., Dict. Sci. Nat. 34: 217. 1825. TYPE: Drozia dicephala Cass. [= Perezia pungens (Humb. & Bonpl.) Less.].

Dumerilia Less., Linnaea 5: 13. 1830, non Lag. ex DC., 1812, nora. illeg.

Heteranthus Cass., Dict. sci. nat. 21: 110. 1821, an admitted orthographic variant of the name Homoianthus (Simpson Vuilleumier, 1969).

Homanthis Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. 4 (folio ed.): 10. 1818, probably an orthographic variant of the name Homoianthus.

Homoeanthus Spreng., Syst. veg. 3: 503. 1826, an orthographic variant of the name Homoianthus.

Homoianthus DC., Ann. Mus. Natl. Hist. Nat. 19: 65. 1812. TYPE: Homoianthus pungens (Humb. & Bonpl.) DC., based on Chaetanthera pungens Humb. & Bonpl. [= Perezia pungens (Humb. & Bonpl.) Less.].

Isanthus DC., Prodr. 7: 63. 1838, nora. nudum.

Pogonura DC. ex Lindl., Intr. nat. syst. bot.: 263. 1836, nom. nudum.

Scolymanthus DC., Prodr. 7: 63. 1838, nom. nudum.

Stenophyllum Less., Syn. gen. Compos.: 412. 1832, a name frequently listed as a synonym of Perezia, but it was actually used only as a sectional name by Lessing (Simpson Vuilleumier, 1969).

Etymology. In honor of Lazaro (or Laurentio) Perez, a former pharmacist from Toledo, Spain.

Herbs perennial, caulescent or acaulescent. Leaves pinnately veined or parallel veined when sheathing, glabrous to densely pubescent; basal leaves rosulate; pseudopetiolate to clasping; blades linear, oblong to ovate, margin entire to lyrate or pinnately partite; upper leaves alternate, numerous to absent, reduced to bracts to broadly oblong or ovate, margin entire, ciliate to lacerate; clasping. Capitulescences monocephalous, corymbose to paniculate; capitula pedunculate to subsessile, homogamous, discoid or radiate; receptacle epaleate; involucre multiseriate. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, glabrous to villose; pappus of scabrid, capillary bristles.

Pollen Description. Perezia (114 of Figs. 112-116) (Parra & Marticorena, 1972). Pollen subprolate to spheroidal-oblate, medium size, tricolporate, exine Oxyphyllum type (see also Simpson Vuilleumier, 1969; Crisci, 1974a; Crisci & Marticorena, 1978; Markgraf & D'Antoni, 1978; Wingenroth & Heusser, 1983).

Habitat and Distribution. Genus of 32 South American species mainly from the Andes and Patagonia, from Colombia to the Islas Malvinas (Falkland Islands), with a few species in eastern Argentina, Brazil, Paraguay, and Uruguay.

Species list

1. Perezia bellidifolia (Phil.) Reiche, Anales Univ. Chile 116: 437. 1905. Basionym: Homoeanthus bellidifolius Phil. TYPE: Chile. Prov. Valdivia: Cordillera de Valdivia, Huahim, O. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Argentina. Prov. Neuquen: Dept. Lacar, Cerro Chapelco, A. L. Cabrera et al. 22987 (LP).

2. Perezia calophylla (Phil.) Reiche, Anales Univ. Chile 116: 435. 1905. Basionym: Homoeanthus calophyllus Phil. TYPE: Chile. Prov. Valdivia: Pucaullu, O. Philippi 2254 (isotype, LP!). Additional specimen examined: Argentina. Prov. Neuquen: San Martin de los Andes, Bridarolli 2229 (LP).

3. Perezia carduncelloides Griseb., Pl. lorentz.: 150. 1874. TYPE: Argentina. Prov. Tucuman: Cienaga, P. G. Lorentz 320 (holotype, B not seen, B photo F n. 16065 at LP!; isotype, CORD not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, Volcan, subida al Cerro Horqueta, A. L. Cabrera et al. 16994 (LP).

4. Perezia carthamoides (D. Don) Hook. & Am., Comp. Bot. Mag. 1: 34. 1835. Basionym: Clarionea carthamoides D. Don. TYPE: Argentina. Prov. Mendoza: Alto de La Laguna, 1821, J. Gillies s.n. (holotype, K not seen). Additional specimen examined: Argentina. Prov. Mendoza: Cordillera del Tigre, King 330 (LP).

5. Perezia ciliaris D. Don ex Hook. & Am., Comp. Bot. Mag. 1: 34. 1835. TYPE: Chile: s.d., J. Dombey s.n. (lectotypes designated by Simpson Vuilleumier, 1969: 95, P, G, not seen, P photo at LP!, G photo F n. 8278 at LP!). Additional specimen examined: Argentina. Prov. Tucumam: Quebradas del Baron, H. A. Fabris 1382 (LP).

6. Perezia ciliosa (Phil.) Reiche, Anales Univ. Chile 116: 426. 1905. Basionym: Clarionea ciliosa Phil. TYPE: Chile. Tarapaca, Cerro de Copacoya, F. Philippi 2250 (isotype, LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Susques, Alto Chorrillos, A. L. Cabrera & H. Schwabe 124 (LP).

7. Perezia coerulescens Wedd., Chlor. andina 1: 39. 1855. TYPE: Peru. Cuzco, Oct 1839-Feb 1840, C. Gay s.n. (holotype, P not seen). Additional specimen examined: Peru. Dept. Puno: Prov. Melgao, Hacienda Huaycho, E. Antazana 51 (LP).

8. Perezia delicata Vuilleum., Contr. Gray Herb. 199: 128. 1969. TYPE: Argentina. Prov. Rio Negro: region del Lago Nahuel Huapi, Paso de las Nubes, A. L. Cabrera 5917 (holotype, LP!). Additional specimen examined: Argentina. Prov. Rio Negro: Cerro L6pez, H. A. Fabris 2161 (LP).

9. Perezia eryngioides (Cabrera) Crisci & Martic., J. Arnold Arbor. 59: 352. 1978. Basionym: Trixis eryngioides Cabrera. TYPE: Brazil. State Santa Catarina: Mun. Irani, campo de Irani, X B. Smith & R. M. Klein 7777 (holotype, LP!*).

10. Perezia fonkii (Phil.) Reiche, Anales Univ. Chile 116: 434. 1905. Basionym: Clarionea fonkii Phil. TYPE: Chile. Cordillera del Doce, Fonk 59 (isotype, LP!). Additional specimen examined: Argentina. Prov. Rio Negro: Dept. Bafiloche, Cerro Lopez, O. Boelcke 1963 (LP).

11. Perezia kingii Baker, Fl. bras. 6: 380. 1884. TYPE: Uruguay. Concepcion, Puerto de Salamanca, Oct 1875, P G. Lorentz s.n. (holotype, B not seen, B photo F n. 16074 at LP!; isotype, GH not seen). Additional specimen examined: Uruguay. Dept. Florida: San Pedro del Timote, arroyo Timote, A. Gallinal et al. PE-5-334 (LP).

12. Perezia lactucoides (Vahl) Less., Linnaea 5: 22. 1830. Basionym: Perdicium laetucoides Vahl. TYPE: Chile. Prov. Magallanes: s.d., D. Commerson s.n. (isotype, C not seen, C photo F n. 22606 at LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Ushuaia, N. Correa & R. Perez Moreau 1853 (LP).

13. Perezia linearis Less., Syn. gen. Compos.: 412. 1832. TYPE: Chile. Prov. Bio Bio: near Antuco, E. F. Poeppig 767 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Rio Negro: Dept. Bariloche, Cerro Lopez, H. A. Fabris 2179 (LP).

14. Perezia lyrata (J. Remy) Wedd., Chlor. andina 1: 42. 1855. Basionym: Homoianthus lyratus J. R6my. TYPE: Chile. Prov. Colchagua: Yalcaregue, C. Gay 283 (holotype, P not seen, P photo at LP!). Additional specimen examined: Chile. Prov. Talca: Laguna del Maule, O. Boelcke et al. 10430 (LP).

15. Perezia magellanica (L. f.) Lag., Amen. nat. Espan.: 31. 1811. Basionym: Perdicium magellanicum L. f. TYPE: Argentina. Prov. Tierra del Fuego, s.d., Forster s.n. (holotype, LINN 1003.5 not seen). Additional specimen examined: Argentina. Prov. Santa Cruz, Dept. Lago Argentino, Cerro Fitz Roy, P. Agostini B-30 (LP).

16. Perezia mandonii Rusby, Mem. Torrey Bot. Club. 3: 66. 1893. TYPE: Bolivia. Oruro, Capi, M. Bang 777 (holotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, Volcan, subida al cerro Horqueta, A. L. Cabrera et al. 16996 (LP).

17. Perezia megalantha Speg., Revista Fac. Agron. Univ. Nac. La Plata 3: 540. 1897. TYPE: Argentina. Prov. Santa Cruz: Lago Argentino, 1890, Tonini del Furia s.n. (holotype, LP!).

18. Perezia multiflora (Humb. & Bonpl.) Less., Linnaea 5: 19. 1830. Basionym: Chaetanthera multiflora Humb. & Bonpl. TYPE: Ecuador. Prov. Pichincha: Volcan, Antisana, s.d., s. leg. (holotype, P not seen). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Tumbaya, Aguada Cerro Abra Morada, A. L. Cabrera et al. 16997 (LP).

19. Perezia nutans Less., Syn. gen. Compos.: 409. 1832. TYPE: Chile. Antuco, meseta de Antuco, E. F. Poeppig 896 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Neuquen: Dept. Minas, cajon del arroyo Covunco, O. Boelcke et al. 11460 (LP).

20. Perezia pedicularidifolia Less., Syn. gen. Compos.: 410. 1832. TYPE: Chile. Prov. Bio Bio: Andes de Antuco, Pico de Pilque, s.d., E. F. Poeppig s.n. (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Rio Negro: Dept. Bariloche, laguna Frias, Cerro Riggi, A. L. Cabrera 6052 (LP).

21. Perezia pilifera (D. Don) Hook. & Am., Comp. Bot. Mag. 1: 34. 1835. Basionym: Clarionea pilifera D. Don. TYPE: Argentina. Prov. Mendoza: Andes de Mendoza, Cerro de Pulcura, s.d., J. Gillies s.n. (holotype, K not seen). Additional specimen examined: Chile. Prov. Nuble: Termas de Chillan, Pirigallo, A. L. Cabrera 3642 (LP).

22. Perezia pinnatifida (Humb. & Bonpl.) Wedd., Chlor. andina 1: 40. 1855. Basionym: Chaetanthera pinnatifida Humb. & Bonpl. TYPE: Lectotype, designated by Simpson Vuilleumier, 1969: 140, the illustration in Pl. Aequin. 2: 170, pl. 136. 1809. Additional specimen examined: Bolivia. La Paz: Larecaja, vicinity of Sorata, between Pongo and Amlaya, Mandon 20 (US, not seen).

23. Perezia poeppigii Less., Syn. gen. Compos.: 411. 1832. TYPE: Chile. Boreal Andes, s.d., E. F. Poeppig s.n. (holotype, P not seen). Additional specimen examined: Chile. Prov. Santiago: Alhue, Monte Cantillana, E. Barros 2025 (LP).

24. Perezia prenanthoides Less., Syn. gen. Compos.: 409. 1832. TYPE: Chile. Prov. Bio Bio: Andes de Antuco, Valle de Quilai, E. F. Poeppig 923 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Rio Negro: Cerro Lopez, A. Corte 226 (LP).

25. Perezia pungens (Humb. & Bonpl.) Less., Linnaea 5: 20. 1830. Basionym: Chaetanthera pungens Humb. & Bonpl. TYPE: Ecuador. Prov. Pichincha: Mt. Pichincha near Quito, s.d., F. Humboldt & A. Bonpland s.n. (holotype, P not seen; isotype, B not seen, B photo F n. 16084 at LP!). Additional specimen examined: Peru. Prov. Cajamarca: Dept. Cajamarca, Challuayaco, entre Kumulca y Celendin, I. Sanchez V. & W. Ruiz V. 468 (LP).

26. Perezia purpurata Wedd., Chlor. andina 1: 43. 1855. TYPE: Bolivia. Potosi, d'Orbigny 1420 (holotype, P not seen, P photo at LP!). Additional specimen examined: Argentina. Prov. Salta: Sierra del Cajon, D. Rodriguez 1376 (LP).

27. Perezia pygmaea Wedd., Chlor. andina 1: 40. 1855. TYPE: Bolivia. Prov. La Paz: ravine of Chuquiaguillo, near La Paz, 1851, H. A. Weddell s.n. (holotype, P not seen, P photo F n. 38128 at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Humahuaca, Mina Aguilar, A. L. Cabrera et al. 19789 (LP).

28. Perezia recurvata (Vahl) Less., Linnaea 5: 21. 1830. Basionym: Perdicium recurvatum Vahl. TYPE: Chile. Prov. Magallanes: strait of Magallan, s.d., D. Commerson s.n. (holotype, C not seen, C photo F n. 22607 at LP!). Additional specimen examined: Argentina. Prov. Tierra del Fuego: Ushuaia, Rio Grande, N. Alboff 374 (LP).

29. Perezia spathulata (D. Don) Hook. & Am., Comp. Bot. Mag. 1: 33. 1835. Basionym: Clarionea spathulata D. Don (see Observations). TYPE: Chile: s. d., H. Ruiz & J. A. Pavon s.n. (type, probably MA). Additional specimen examined: Argentina. Prov. Neuquen: Dept. Lacar, Hua-Hum, camino al Lago Queni, J. Diem 3252 (LP).

30. Perezia squarrosa (Vahl) Less., Linnaea 5: 15. 1830. Basionym: Perdicium squarrosum Vahl. TYPE: Uruguay. Dept. Canelones: sommet du Morne de Montevideo, 1767, D. Commerson s.n. (holotype C, not seen). Additional specimen examined: Paraguay. Paraguari: Ybytimi, Montes 12951 (LP).

31. Perezia sublyrata Domke, Notizbl. Bot. Gart. Berlin-Dahlem 13: 248. 1936. TYPE: Peru. Puno, Santa Rosa, Stafford 345 (holotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Yilcara, subida al Abra de Remate, H. Sleumer 4075 (LIL).

32. Perezia volcanensis Cabrera, Fl. Prov. Jujuy 10: 660. 1978. TYPE: Argentina. Prov. Jujuy: Dept. Tumbaya, Estacion Volcan, L. Castillon 8155 (holotype, LIL, photo at LP!: see Observations).

Observations. Perezia spathulata is cited as a synonym of P. viscosa Less. in the revision of Perezia (Simpson Vuilleumier, 1969). However, as mentioned by Cabrera (1971b), P. spathulata is the correct name by priority.

The citation of the type specimen of Perezia volcanensis as L. Castillon 7020 in Flora of Jujuy (Cabrera, 1978) is a typographic error (Katinas, 1995; Freire & Iharlegui, 2000).

The genus Calorezia, containing two species of Perezia, was recently proposed by Panero (2007) based on a chloroplast DNA coding regions study in preparation by Panero et al. Since there are not clear morphological characters that support this new genus, it is included in the synonymy of Perezia.

Literature. Simpson (1973), Simpson Vuilleumier (1969), Crisci and Marticorena (1978), Katinas (1995).

Pleocarphus D. Don

Trans. Linn. Soc. London 16: 228. 1830. TYPE: Pleocarphus revolutus D. Don. 66 of Figs. 63-66.

Etymology. From the Greek pleos, plenty, full, filled, and karphos, any small dry body, in reference to the receptacle completely paleate.

Shrubs, stems with brachyblasts and macroblasts. Leaves alternate; sessile; blades linear to linear-lanceolate, pinnately veined, margin sinuate to dentate, revolute, glabrescent. Capitulescences densely paniculate; capitula sessile to shortly pedunculate, homogamous, discoid; receptacle paleate, paleae embracing all the florets; involucre uniseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex, pubescent; pappus of scabrid, capillary bristles.

Pollen Description (Parra & Marticorena, 1972). Pollen prolate-subprolate, medium size, tricolporate, exine Trixis type (see also Crisci, 1974a).

Habitat and Distribution. Monotypic genus endemic to rocky soils of central and northern Chile.

Species list

Pleocarphus revolutus D. Don, Trans. Linn. Soc. London 16: 229. 1830. TYPE: Chile. Prov. Coquimbo: s.d., Caldcleugh s.n. (holotype, G not seen, G photo at LP!, US!). Additional specimen examined: Chile. Prov. Coquimbo: alrededores de La Serena, Quebrada Romeral, A. L. Cabrera 11407 (LP).

Literature. Gay (1847).

Polyachyrus Lag.

Amen. nat. Espan.: 37. 1811. TYPE: Cephaloseris poeppigii Kuntze ex Less. [= Polyachyrus poeppigii (Kunze ex Less.) Less.] 67 of Figs. 67-70.

[FIGURES 67-70 OMITTED]

Bridgesia Hook., Bot. misc. 2: 228. 1831. TYPE: Bridgesia echinopsoides Hook. [= Polyachyrus sphaerocephalus D. Don].

Cephaloseris Poepp. ex Rchb., Consp. regn. veg. 1: 100. 1828. TYPE: Cephaloseris poeppigii Kunze ex Less., Linnaea 5: 5. 1830 [= Polyachyrus poeppigii Kunze ex Less. vat. poeppigii].

Diaphoranthus Meyen, Reise Chile 1: 404. 1834. TYPE: Diaphoranthus fuscus Meyen [= Polyachyrus fuscus (Meyen) Walp.].

Polyachrus Hook. & Am., Comp. Bot. Mag. 1: 36. 1835, an orthographic variant of the name Polyachyrus.

Polyachurus DC., Ann. Mus. Natl. Hist. Nat. 19: 69. 1812, an orthographic variant of the name Polyachyrus.

Polyachyris Walp., Repert. Bot. Syst. (Walpers) 2: 681. 1843, an orthographic variant of the name Polyachyrus.

Etymology. From Greek polys, many, and achyron, chaff, husks, referring to the number of pappus bristles.

Subshrubs erect or decumbent. Leaves pinnately veined, pubescent; basal leaves rosulate to sub-rosulate; decurrent to amplexicaul; blades linear to oblong, pinnately lobed to pinnately partite, auriculate at the base, margin spiny to unarmed; upper leaves alternate, similar to the basal leaves but reduced. Capitulescences a corymbose pseudocephalium; capitula shortly-pedunculate, homogamous, discoid, with two florets, rarely three, surrounded by an involucre of five phyllaries: four enclosing one floret and one enclosing the other floret or all florets, the phyllary between florets interpreted as a palea; receptacle paleate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs Cypselae truncate to constricted at the apex, glabrescent to pubescent; pappus dimorphic, scabrid to plumose bristles, shorter in the inner floret, capillary to somewhat paleaceous.

Pollen Description. Polyachyrus (Parra & Marticorena, 1972). Pollen subprolatespheroidal, medium size, tricolporate, exine Oxyphyllum type (see also Telleria et al., 2003).

Habitat and Distribution. Genus of seven species endemic to the arid regions of northern and central Chile and Peru.

Species list

1. Polyachyrus annuus I. M. Johnst., Contr. Gray Herb. 8: 134. 1929. TYPE: Chile. Prov. Antofagasta: Dept. Tocopilla, Tocopilla, ca. 6 km N of port and about opposite Caleta Duendes, I. M. Johnston 3538 (isotype, CONC!). Additional specimen examined: Peru. Dept. Arequipa: Prov. Camana, R. Ferreyra 2518 (LP).

2. Polyachyrus carduoides Phil., Fl. atacam.: 202. 1860. TYPE: Chile. Prov. Antofagasta: Dept. Taltal, Valle Sandon, s.d., R. A. Philippi s.n. (holotype, SGO not seen). Additional specimen examined: Chile. Prov. Atacama: Encantada, s.d., G. E. Gulig s.n. (LP).

3. Polyachyrus cinereus Ricardi & Weldt, Gayana, Bot. 26: 26. 1974. TYPE: Chile. Prov. Antofagasta: region about Aguada Cachina, waterhole in Quebrada Cachina, ca. 6 km in land from Caleta Esmeralda, I. M. Jhonston 5726 (holotype, GH not seen). Additional specimen examined: Chile. Prov. Copiapo: Pan de Azucar, S. Roig 19 (LP).

4. Polyachyrus fuscus (Meyen) Walp., Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 1: 288. 1843. Basionym: Diaphoranthus fuscus Meyen. TYPE: Chile. Cordillera de Copiapo, 1833, Meyen s.n. (holotype, B not seen, B photo F n. 15992 at LP!). Additional specimen examined: Chile. Prov. Antofagasta: la Chimba, E. Barros 2083 (LP).

5. Polyachyrus gayi J. Rdmy, Fl. chil. 3: 372. 1847. TYPE: Chile. Costas de las provincias septentrionales, s.d., C. Gay s.n. (holotype, P not seen, P photo F n. 15993 at LP!; isotype, SGO not seen). Additional specimen examined: Chile. Prov. Antofagasta: la Chimba, E. Barros 2079 (LP).

6. Polyachyrus poeppigii (Kunze ex Less.) Less., Linnaea 5: 5. 1830. Basionym: Cephaloseris poeppigii Kunze ex Less. TYPE: Chile. Prov. Valparaiso: Concon, E. F. Poeppig 221 (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Chile. Prov. Valparaiso: Quebrada en ruta 5 y Puente Juan Soldado, M. Bonifacino 1366 (LP).

7. Polyachyrus sphaerocephalus D. Don, Trans. Linn. Soc. London 16: 229. 1830. TYPE: Chile. Prov. Tarapaca: Dept. Iquique, Usmagama, Feb 1885, F. Philippi s.n. (holotype, B not seen, B photo F n. 15996 at LP!). Additional specimen examined: Peru. Dept. Lima: Infiernillo, A. L. Cabrera 10891 (LP).

Literature. Ricardi and Weldt (1974), Katinas and Crisci (2000), Katinas et al. (2008a).

Proustia Lag.

Amen. nat. Espan.: 33. 1811. TYPE: Proustia pyrifolia DC. 68 of Figs. 67-70.

Etymology. In honor of Ludovico Proust, a chemist from Madrid, Spain, of the early nineteenth century.

Shrubs or little trees, erect or scandent, some branches modified in thorns to unarmed, stems with brachyblasts and macroblasts. Leaves alternate; petiolate to subsessile; blades linear, oblong to ovate, pinnately veined, margin entire to spinosedentate, glabrous to pubescent. Capitulescences densely racemose to paniculate; capitula subsessile, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth to slightly papillose; style bifid, branches rounded at the apex, dorsally papillose. Cypselae truncate to slightly constricted at the apex, glabrous to pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Proustia (Telleria et al., 2003). Pollen subprolate, medium size, tricolporate, exine Trixis type and both Proustia and Trixis type in a single specimen, with polar caps only in P pyrifolia (see also Parra & Marticorena, 1972; Crisci, 1974a).

Habitat and Distribution. Genus of three South American species distributed in the Andean regions of southern Peru, Bolivia, central Chile, and Argentina.

Species list

1. Proustia cuneifolia D. Don, Trans. Linn. Soc. London 16: 202. 1830. TYPE: Chile. Coquimbo, s.d., Caldcleugh s.n. (type, probably G). Additional specimen examined: Argentina. Prov. Salta: Dept. Rosario de Lerma, Santa Rosa de Yastil, A. L. Cabrera 9008 (LP).

2. Proustia ilicifolia Hook. & Am., Bot. Beechey Voy.: 28. 1840. TYPE: Chile. Coquimbo, s.d., s. leg. (type, probably K). Additional specimen examined: Chile. Prov. Atacama: Dept. Copiapo, La Puerta, M. Ricardi & C. Marticorena 3667 (CONC).

3. Proustia pyrifolia DC., Ann. Mus. Natl. Hist. Nat. 19: 70. 1812. TYPE: Hab. in Chili prope Talcahuano, s.d., Lagasca s.n. (type, not localized). Additional specimen examined: Chile. Prov. Valparaiso: Cerro de la Campana, April 1933, Grandjot s.n. (CONC).

Observation. Proustia has sometimes been placed in the subtribe Mutisiinae (Cabrera, 1961; Fabris, 1968) because of its rounded, dorsally papillose style branches. The type of exine stratification in the pollen of Proustia (Crisci, 1974a; Telleria et al., 2003) clearly indicates the position of this genus within the subtribe Nassauviinae (= tribe Nassauvieae).

Literature. Fabris (1968), Crisci (1974a).

Triptilion Ruiz & Pav.

Fl. peruv, prodr.: 102. 1794. TYPE: Triptilion spinosum Ruiz &Pav. 69 of Figs. 67-70. Triptilium DC., Ann. Mus. Natl. Hist. Nat. 19: 59. 1812, an orthographic variant of the name Triptilion.

Etymology. From the Greek treis, tria, three, and ptilon, feather, referring to the three scales of the pappus.

Herbs annual or perennial, caulescent to acaulescent. Leaves pinnately veined, pubescent; basal leaves rosulate to spread; sessile; blades linear-lanceolate to orbicular, entire to pinnatisect, margin spiny to unarmed; upper leaves alternate, similar to the basal leaves but reduced. Capitulescences corymbose, paniculate to glomerulose or a pseudocephalium; capitula sessile to pedunculate, homogamous, discoid, five-flowered; receptacle epaleate; involucre two- to three-seriate. Florets isomorphic, bisexual, corolla bilabiate; anther apical appendages acute, tails smooth; style bifid, branches apically truncate, crowned by collector hairs. Cypselae truncate at the apex, glabrous to pubescent; pappus with three, rarely two or four, narrowly paleaceous bristles, longitudinally plicate, apically plumose to laciniate, rarely crown-like.

Pollen Description. Triptilion (Parra & Marticorena, 1972). Pollen spheroidal to suboblate, small to medium size, tricolporate, exine Oxyphyllum type (see also Crisci, 1974a).

Habitat and Distribution. Genus of seven species endemic to Chile and to the subantarctic forests of southwestern Argentina.

Species list

1. Triptilion achilleae DC., Prodr. 7: 51. 1838. TYPE: Chile. Tagua-Tagua, C. Bertero 708 (holotype, P!). Additional specimen examined: Argentina. Prov. Neuquon: Dept. Huiliches, Lago Huechulaufquen, A. L. Cabrera 11286 (LP).

2. Triptilion benaventii J. Remy, Fl. chil. 3: 357. 1848. TYPE: Chile. Concepcion, C. Gay 252 (holotype, P!). Additional specimen examined: Chile. Malleco, camino de Icalma a Liucura, Marimenuco, en las inmediaciones de la confluencia del arroyo Chanchoco con el Bio Bio, Pfister 7382 (CONC).

3. Triptilion berteroi Phil., Anales Univ. Chile 87: 91. 1894. TYPE: Chile. Rancagua, 1818, C. Bertero s.n. (holotype, SGO!). Additional specimen examined: Chile. Santiago, Colina, cerro en la vecindad de Los Banos, s.d., Behn s.n. (CONC).

4. Triptilion capillatum (D. Don) Hook. & Am., Comp. Bot. Mag. 2: 43. 1836. Basionym: Nassauvia capillata D. Don. TYPE: Chile. Baths of Collina, s.d., Macrae s.n. (phragment of holotype, K!). Additional specimen examined: Chile. Quillota, Cerro La Campana, O. Zollner 7280 (LP).

5. Triptilion cordifolium Lag. ex Lindl., Bot. Reg. 10: 853. 1824. TYPE: Lectotype, designated by Katinas et al., 1992: 112, the illustration in Bot. Reg. 10: 852. 1824. Additional specimen examined: Chile. Petorca: Zapallar, Kausel 4530 (LP).

6. Triptilion gibbosum J. Remy, Fl. chil. 3: 356. 1848. TYPE: Chile. Colchagua, s.d., C. Gay s.n. (type, P!). Additional specimen examined: Chile. Elqui, La Serena, E. Barros 2429 (LP).

7. Triptilion spinosum Ruiz &Pav., Syst. veg. fl. peruv, chil. 1: 185. 1798. TYPE: Chile. Concepcion, 1782, H. Ruiz & J. A. Pavon s.n. (holotype, MA not seen, MA photo at LP!). Additional specimen examined: Chile. Prov. Antofagasta: Cerro Moreno, O. Zollner 3367 (LP).

Literature. Katinas et al. (1992, 2008a).

Trixis P. Browne

Civ. nat. hist. Jamaica: 312. 1756. TYPE: Trixis inula Crantz. 70 of Figs. 67-70.

Bowmania Gardner, London J. Bot. 2: 9. 1843. TYPE: Bowmania verbascifolia Gardner [= Trixis nobilis (Vell.) Katinas].

Castra Vell., Fl. flumin.: 343. 1829 (1825). TYPE: Castra praestans Vell. [= Trixis praestans (Vell.) Cabrera].

Prionanthes Schrank, Pl. rar. hort. monac. 2: 51. 1820. TYPE: Prionanthes antimenorrhoea Schrank [= Trixis divaricata (Kunth) Spreng. var. divaricata].

Tenorea Colla, Hortus ripul.: 137. 1824, non Raf., 1814, Gasp., 1845, nec K. Koch, 1869, nom. illeg. TYPE: Tenorea berteri Colla [= Trixis inula Crantz].

Etymology. From the Greek trixos, trissos, three times, in reference to the tridentate outer lip of the corolla.

Shrubs, subshrubs, erect to climbing, or herbs perennial, caulescent. Leaves alternate, rarely basal leaves rosulate; petiolate, sessile, decurrent to auriculate; blades linear, elliptic to ovate, pinnately veined, margin mucronulate to serrate, sometimes revolute, pubescent. Capitulescences corymbose, paniculate to spicate; capitula sessile to pedunculate, homogamous, discoid or radiate; receptacle epaleate; involucre one- to five-seriate. Florets isomorphic or dimorphic, bisexual; marginal florets with corolla bilabiate, outer lip expanded or not; central florets with corolla bilabiate; anther apical appendages acute, tails smooth to papillose; style bifid, branches apically truncate, crowned by collector hairs. Cypselae constricted at the apex to longly rostrate, pubescent; pappus of scabrid, capillary bristles.

Pollen Description. Trixis (115 of Figs. 112-116) (Parra & Marticorena, 1972). Pollen prolate, medium to large size, tricolporate, exine Trixis type (see also Crisci, 1974a).

Habitat and Distribution. Neotropical genus of 38 species distributed from southern United States to central Chile and Argentina.

Species list

1. Trixis aggregata Rusby, Mere. Torrey Bot. Club 6: 71. 1896. TYPE: Bolivia. Bolivian palteau, M. Bang 1150 (isotype, BM!). Additional specimen examined: Bolivia. Dept. Cochabamba, Prov. Cercado, Cerro San Pedro, J. Steinbach 9705 (MO).

2. Trixis alata D. Don, Trans. Linn. Soc. London 16: 192. 1830. TYPE: Mexico. State Morelos: Ayacapixtl (Yecapixtla), s.d., M. Sesse 62 Mocino s.n. (holotype, G-DEL not seen).

3. Trixis angustifolia DC., Prodr. 7: 69. 1838. TYPE: Mexico. State San Luis Potosi: pied des monts de Catorce, J. L. Berlandier 1353 (isotype, NY not seen, NY photo at LP!).

4. Trixis cacalioides (Kunth) D. Don, Trans. Linn. Soc. London 16: 187. 1830. Basionym: Perdicium cacalioides Kunth. TYPE: Peru. Andes, s.d., A. Bonpland s.n. (holotype, P!). Additional specimen examined: Peru. Dept. Amazonas, Prov. Chachapoyas, Rio Maranon at Balsas, Hutchinson & Wright 5407 (MO).

5. Trixis calcicola B. L. Rob., Proc. Amer. Acad. Arts 49: 516. 1913. TYPE: Mexico. State Guerrero: Iguala Canyon, near Iguala, C. G. Pringle 13921 (isotype, US not seen, US photo at LP!).

6. Trixis californica Kellogg, Proc. Calif. Acad. Sci. 2: 182. 1862. TYPE: Mexico. State Baja California Norte: Cedros Island, Veatch, s.d., s. leg. (holotype, CAS not seen). Additional specimen examined: Mexico. Baja California: Sierra de la Giganta, Pilon de las Parras, W of Loreto, A. Carter 62 H. Sharsmith 4237 (LP).

7. Trixis calycina D. Don, Trans. Linn. Soc. London 16: 187. 1830. TYPE: Brazil. State Bahia, s.d., C. Martius s.n. (holotype, M not seen, M photo at LP!). Additional specimen examined: Brazil. State Bahia: Mun. Morro da Chapeu, Serra do Tombador, G. Hatschbach & Guimaraes 42354 (LP).

8. Trixis chiapensis C. E. Anderson, Brittonia 23: 347. 1971. TYPE: Mexico. State Chiapas: 24 mi SE of Comitan, W. Anderson & C. Anderson 5494 (holotype, MICH not seen).

9. Trixis divaricata (Kunth) Spreng., Syst. veg. 3: 501. 1826. Basionym: Perdicium divarieatum Kunth. TYPE: Peru. Carico, F. Humboldt & A. Bonpland 3650 (holotype, P not seen, P photo F n. 37850 at LP! MO!). Additional specimen examined: Argentina. Prov. Tucuman: Dept. Trancas, El Cadillal, S. Venturi 377 (LP).

10. Trixis glaziovii Baker, Fl. bras. 6: 391. 1884. TYPE: Brazil. State Rio de Janeiro: Itatiaia, Campo Feio, A. F. M. Glaziou 6582 (holotype, P!). Additional specimen examined: Brazil. State Minas Gerais: Caldas, 10 Jan 1919, Hoehne s.n. (SP).

11. Trixis glutinosa D. Don, Trans. Linn. Soc. London 16: 189. 1830. TYPE: Brazil. State Minas Gerais: inter Minas Novas et fluvium Sao Francisci, s.d., C. Martius s.n. (holotype, M not seen, M photo at LP!); State Goias, rod. br. 153, I0 km W de Anapolis, G. Hatschbach 36682 (LP!).

12. Trixis grandibracteata C. E. Anderson, Brittonia 23: 348. 1971. TYPE: Mexico. State Guerrero: 21 km E of Petatlan, on road to Tlapa, at km 121, W. Anderson & C. Anderson 5729 (isotype, NY not seen, NY photo at LP!).

13. Trixis grisebachii Kuntze, Revis. gen. pl. 3:183. 1898. TYPE: Argentina. Prov. Jujuy: Jujuy, April 1871, P. G. Lorentz & G. H. E. W. Hieronymus s.n. (isotype, NY not seen, NY photo at LP!). Additional specimen examined: Argentina. Prov. Santiago del Estero: Dept. Rio Hondo, camino de San Miguel de Tucuman a Rio Hondo, L. Katinas et al. 97 (LP!).

14. Trixis haenkei Sch. Bip., Bot. Voy. Herald: 314. 1856. TYPE: Mexico. State Sinaloa: probably from the mountains along the Sinaloa-Durango border, Seemann 2029 (holotype, K not seen).

15. Trixis hassleri Chodat, Bull. Herb. Boissier 3: 782. 1903. TYPE: Paraguay. Dept. Amambay: Rio Apa, Hassler 8318 (isotype, GH!). Additional specimen examined: Paraguay. Dept. Amambay: 30 km SE de Bella Vista, A. Krapovickas & A. Schinini 32645 (SI).

16. Trixis hyposericea S. Watson, Proc. Amer. Acad. Arts 25: 157. 1890. TYPE: Mexico. State Jalisco: in the barranca near Guadalajara, C. G. Pringle 1741 (isotype, US not seen, US photo at LP!).

17. Trixis inula Crantz, Inst. rei herb. 1: 329. 1766. TYPE: Jamaica, s.d., s. leg. (holotype, LINN 1000.3 not seen, LINN photo at LP!). Additional specimen examined: Colombia. Dept. Atlantico: Puerto Colombia, Elias 1158 (LP).

18. Trixis lessingii DC., Prodr. 7: 70. 1838. TYPE: Brazil. Brasilia aequinoctialis, F. Sellow 2068 (lectotype, designated by Katinas, 1996b: 72, K!). Additional specimen examined: Brazil. State Santa Catarina: Matos Costa, R. M. Klein 3569 (LP).

19. Trixis megalophylla Greenm., Proc. Amer. Acad. Arts 41: 270. 1905. TYPE: Mexico. State Guerrero: between Sochi and Tlakinsala, E. W. Nelson 2038 (isotype, US not seen, US photo at LP!).

20. Trixis mexicana Lex., Nov. veg. descr. 1: 27. 1824. TYPE: not localized. Additional specimen examined: Mexico. State Michoacan: about 17 mi W of Jiquilpan, A. Cronquist 9757 (LP).

21. Trixis michuacana Lex., Nov. veg. descr. 1. 28. 1824. TYPE: Mexico. State Michoacan: prope Vallisoletum, s.d., LaLlave s.n. (holotype, G-DEL not seen). Additional specimen examined: Mexico. State Jalisco: Barranca du Rio Santiago, Dec 1912, Diguet s.n. (LP).

22. Trixis nelsonii Greenm., Proc. Amer. Acad. Arts 41: 270. 1905. TYPE: Mexico. State Chiapas: between San Crist6bal and Teopsica, E. W. Nelson 3459 (isotype, US not seen, US photo at LP!).

23. Trixis nobilis (Vell.) Katinas, Darwiniana 34: 74. 1996. Basionym: Castra nobilis Vell. TYPE: Lectotype, designated by Katinas, 1996b: 74, the illustration in Fl. flumin, icon. 8: tab. 82. 1831 (1827). Additional specimen examined: Brazil. State Goias: Serra dos Pirineus, 20 km of Corumba de Goias, on road to Niquelandia, Irwin et al. 18746 (SP).

24. Trixis ophiorhiza Gardner, London J. Bot. 6: 461. 1847. TYPE: Brazil. State Minas Gerais: bewteen the Rio de Sao Francisco and Formigas, G. Gardner 4959 (holotype, BM!). Additional specimen examined: Brazil. Federal district: Fundacao Zoobotanica, Pires et al. 9583 (LP).

25. Trixis pallida Less., Linnaea 5: 30. 1830. TYPE: Brazil. s.d., Sello 3267 (isotype, K not seen, K photo at LP!). Additional specimen examined: Argentina. Prov. Chaco: Dept. Sargento Cabral, Colonia La Elisa, Schulz 1050 (LP!).

26. Trixis parviflora C. E. Anderson, Brittonia 23: 351. 1971. TYPE: Mexico. State Oaxaca: ca. 12 km N of Amusgos, on road from Pinotepa Nacional to Tlaxiaco, near km 205, W. Anderson & C. Anderson 5636 (isotype, NY not seen, NY photo at LP!).

27. Trixis peruviana Katinas, Darwiniana 34: 83. 1996. TYPE: Peru. Dept. Lima: Cajatambo, Banos de Churin, Landeman 5425 (holotype, K!).

28. Trixis praestans (Vell.) Cabrera, Revista Mus. La Plata, Secc. Bot. 1: 61. 1936. Basionym: Castra praestans Veil. TYPE: Lectotype, designated by Katinas, 1996b: 61, the illustration in Fl. flumin, icon. 8: tab. 80. 1831 (1827). Additional specimen examined: Brazil. State Minas Gerais: Grao Mogol, G. Hatschbach 41513 (LP).

29. Trixis pringlei B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 40: 10. 1904. TYPE: Mexico. State Oaxaca: Tomellin Canyon, C. G. Pringle 5894 (holotype, GH not seen).

30. Trixis proustioides Hieron., Bot. Jahrb. Syst. 28: 653. 1901. TYPE: Colombia. Triana 1508 (holotype, B not seen, B photo F n. 16108 at LP!). Additional specimen examined: Argentina. Prov. Jujuy: Dept. Santa Barbara, camino de Santa Clara a El Fuerte, Abra de los Morteros, A. L. Cabrera et al. 27968 (LP).

31. Trixis pruskii D. J. N. Hind, Kew Bull. 55: 382. 2000. TYPE: Brazil. State Bahia: Mun. Rio Contas, Carrapato, beira Rio da Agua Suja, Ganev 2483 (isotype, US not seen, US photo at LP!).

32. Trixis pterocaulis B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 40: 8. 1904. TYPE: Mexico. State Colima: Manzanillo, E. Palmer 897 (isotype, US not seen, US photo at LP!).

33. Trixis sagasteguii Cabrera, Bol. Soc. Argent. Bot. 10: 42. 1962. TYPE: Peru. Dept. La Libertad: Prov. Bolivar, Catal, ruta Rio Maranon-Longotea, A. Lopez & A. Sagastegui 3163 (holotype, LP!).

34. Trixis silvatica B. L. Rob. & Greenm., Proc. Amer. Acad. Arts 40: 12. 1904. TYPE: Mexico. State Oaxaca: unterhalb Totolapam, Seler 1633 (lectotype, designated by Anderson, 1972: 55, GH not seen; isotypes, NY not seen, NY photo at LP!, US not seen, US photo at LP!). Additional specimen examined: Mexico. Oaxaca: 35 mi W of Tehuantepec, along the Pan-American highway, A. Cronquist 9681 (LP).

35. Trixis spicata Gardner, London J. Bot. 6: 462. 1847. TYPE: Brazil. State Minas Gerais: campos near the Rio Claro, G. Gardner 4957 (holotype, BM!). Additional specimen examined: Paraguay. Dept. Amambay: Sierra de Amambay, T. Rojas 10251 (LP).

36. Trixis thyrsoidea Dusen ex Malme, Kongl. Svenska Vetensk. Acad. Handl. 12: 116. 1933. TYPE: Brazil. State Parana: Parana, Rio Branco, P. Dusen 13836 (holotype, S not seen, S photo at LP!). Additional specimen examined: Brazil. State Santa Catarina: Vacas Gordas, Urubici, P. Reitz 6685 (LP).

37. Trixis vauthieri DC., Prodr. 7: 69. 1838. TYPE: Brazil. State Minas Gerais: Tejuco, M. Vauthier 278 (isotype, GH!). Additional specimen examined: Brazil. State Minas Gerais: Serra do Curral, Nova Lima, 5 miles SE of Belo Horizonte, Williams & Assis 8018 (MO).

38. Trixis verbascifolia (Gardner) S. F. Blake, Contr. U. S. Natl. Herb. 22: 655. 1924. Basionym: Bowmania verbascifolia Gardner. TYPE: Brazil. State Rio de Janeiro: Organ Mtns., G. Gardner 5797 (holotype, BM!). Additional specimen examined: Brazil. State Rio de Janeiro: Serra dos Orgaos, A. L. Cabrera 12255 (LP*).

Literature. Anderson (1972), Katinas (1996b), Hind (2000).

Tribe Stifflieae D. Don

D. Don, Trans. Linn. Soc. London 16: 291. 1830. TYPE: Stifftia J. C. Mikan.

Stifftioideae (D. Don) Panero, pro parte, Phytologia 89: 356. 2007. TYPE: Stifftia J. C. Mikan.

Wunderlichioideae Panero & V. A. Funk, pro parte, Phytologia 89: 357. 2007. Wunderlichieae Panero & V. A. Funk, Phytologia 89: 357. 2007. TYPE: Wunderlichia Riedel ex Benth.

Simple to branched subshrubs, shrubs, trees, or vines; stems woolly to glabrous. Leaves simple, alternate, often clustered at stem apex, unarmed; usually petiolate, the petioles, when present, often clasping; blades oblanceolate, less commonly linear, orbicular to elliptic, often thickly coriaceous, glabrous and eglandular, or tomentose to woolly beneath. Capitulescences monocephalous, terminal, less commonly cymose, with few to many capitula. Capitula usually large, homogamous, discoid, with one to ca. 100 flowers; involucre multiseriate, the series graduate; receptacle paleate, partially paleate to epaleate. Florets bisexual, corollas actinomorphic, tubular-funnelform to sub-campanulate, deeply five-lobed, exceptionally bilabiate; anthers mostly exserted, apical appendages acuminate, tails long, smooth to papillose, free to those of adjacent anthers often connate; style cleft into two short, bilobed branches, the branches erect, apically rounded, dorsally rugulose to papillose with the branch apex often smooth and the papillae extending beyond the bifurcation point onto the shaft, occasionally styles bifid, branches recurved and dorsally smooth. Cypselae usually prismatic and glabrous; pappus of scabrid capillary bristles, basally connate or not, inner series apically broadened to plumose, unequal to subequal in length. Pollen with exine structure heterogeneous, most genera with compact ectosexine constituted by very densely arranged columellae, and endosexine with stout, ramified columellae; exine surface perforate, microechinate to echinate.

Characters that define the tribe are in italics.

Stifftieae include six genera and 48 species mostly centered in the Guayana Highlands in Colombia, Guyana, and Venezuela. Some species are distributed in the Andes of Ecuador and Peru, and central-eastern Brazil.

Genera: Chimantaea Maguire, Steyerm. & Wurdack, Quelchia N. E. Br., Stenopadus S. F. Blake, Stifftia J. C. Mikan, Stomatochaeta (S. F. Blake) Maguire & Wurdack, Wunderlichia Riedel ex Benth.
Key to Genera

1a. Leaves caducous; plants flowering in the leafless condition;
pappus connate basally, deciduous as a ring, bristles barbellate to
plumose Wunderlichia

1b. Leaves commonly long-persistent; plants thus flowering when leafy;
pappus basally free, persistent, bristles scabrid.

2a. Corolla lobes stiffly erect to very slightly recurved at extreme
apex; corolla cream-colored, not reddish.

3a. Branched or unbranched subshrubs or shrubs; leaves tomentose to
woolly beneath; corolla lobes frequently pilose at tip; anther tails
free; cypselae often pilose Chimantaea

3b. Much-branched subshrubs or trees; leaves glabrous or, if
pubescent, then generally rapidly glabrate; corolla lobes apically
glabrous; tails of adjacent anthers appressed or truly connate;
cypselae usually glabrous Stomatochaeta

2b. Corolla lobes strongly coiled to flexuous, never stiffly erect;
corolla cream-colored, yellowish, or reddish.

4a. Capitula one-flowered, in axillary to terminal glomerate cymes;
anther tails papillose Quelchia

4b. Capitula many-flowered in monocephalous or in lax capitulescences;
anther tails smooth to occasionally papillose.

5a. Shrubby trees or vines; receptacles epaleate; corolla tube
five-nerved; anther tails free Stifftia

5b. Shrubs to big trees; receptacles sometimes paleate; corolla tube
ten- to 15-nerved; anther tails often connate Stenopadus


Chimantaea Maguire, Steyerm. & Wurdack

Mem. New York Bot. Gard. 9: 428. 1957. TYPE: Chimantaea mirabilis Maguire, Steyerm. & Wurdack. 71 of Figs. 71-74.

Etymology. Named for the Chimanta Massif (state Bolivar, Venezuela), the world's largest Massif of closely clustered tepuis, and where all the species of the genus occur.

[FIGURES 71-74 OMITTED]

Shrubs or subshrubs unbranched to branched. Leaves alternate, densely inserted, commonly clustered at stem apex; sessile to petiolate; blades linear, rhomboidal to obovate, thickly coriaceous, pinnately veined, margin entire, frequently revolute, glabrous above except sometimes along midrib, tomentose to woolly beneath. Capitulescences monocephalous, terminal; capitula sessile, homogamous, discoid; receptacle epaleate or with a few deciduous marginal paleae; involucre multiseriate. Florets isomorphic, bisexual, actinomorphic, tubular-funnelform, deeply five-lobed, lobes stiffly erect to slightly recurved, frequently pilose at the tip; anthers yellow to reddish, generally exserted, apical appendages acute to acuminate, tails smooth, free; style yellow to reddish, bilobed, branches dorsally rugulose to papillose much below the bifurcation point. Cypselae often densely pilose; pappus of scabrid bristles, the innermost somewhat longer, apically broadened.

Pollen Description. Chimantaea (116 of Figs. 112-116). Pollen subprolate, elliptic in equatorial view, circular in polar view, large size. Tricolporate, long colpi with scabrate microgranulate membrane. Exine Mutisia type, microechinate. Nexine ca. 2 [micro]m thick (see also Carlquist, 1957b). Chimantaea cinerea (Gleason & S. F. Blake) Maguire, Steyerm. & Wurdack. Pollen size P x E = (64-68 x 50 54) [micro]m. Exine 6 [micro]m thick. Ratio ectosexine/endosexine: ca. 1:1. Chimantaea huberi. Pollen size P x E = (48-57 x 37-42) [micro]m. Exine 5.5-6 [micro]m thick, ectosexine slightly thickened at the poles. Ratio ectosexine/endosexine: 1:1; 1.5:1. SEM: tectum very perforate, apertural membrane scabrate (116 of Figs. 112-116). Note: pollen similar to that of Neblinaea promontorium, Stifftia uniflora, Stenopadus campestris, S. colveii and S. huachamacari (see also Telleria, 2008).

Habitat and Distribution. Chimantaea is a genus of nine species endemic to the eastern Venezuelan Guayana Highland. All nine species of Chimantaea occur on the Macizo del Chimanta, but subspecies of two species are disjunct, occurring outside of the massif on Auyan-tepui and Murisipan-tepui.

Species list

1. Chimantaea acopanensis Steyenn., Ann. Missouri Bot. Gard. 71: 335. 1984. TYPE: Venezuela. State Bolivar: Piar, Massif of Chimanta, plateau of Acopantepui, J. A. Steyermark et al. 129932 (isotype, US!).

2. Chimantaea cinerea (Gleason & S. F. Blake) Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 429. 1957. Basionym: Stenopadus cinereus Gleason & S. F. Blake. TYPE: Venezuela. State Bolivar, Auyantepui, G. H. H. Tate 1111 (holotype, NY!). Additional specimen examined: Venezuela. Bolivar, B. K. Host 3745 (MO).

3. Chimantaea eriocephala Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 431. 1957. TYPE: Venezuela. State Bolivar: Chimanta massif Churitepui (Muru-tepui), J. J. Wurdack 34225 (holotype, NY!).

4. Chimantaea espeletoidea Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 433. 1957. TYPE: Venezuela. State Bolivar: Chimanta massif, below upper falls of Rio Tirica, J. A. Steyermark & J. J. Wurdack 574 (isotype, NY!).

5. Chimantaea huberi Steyerm., Ann. Missouri Bot. Gard. 71: 335. 1984. TYPE: Venezuela. State Bolivar: Piar, J. A. Steyermark et al. 128815 (isotype, US!). Additional specimen examined: Venezuela. State Bolivar: Mun. Gran Sabana, macizo de Chimanta, SE del Apacara-tepui, O. Huber 12034 (LP*).

6. Chimantaea humilis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 432. 1957. TYPE: Venezuela. State Bolivar: Chimanta massif, central section, between W and E branches of headwaters of Rio Tirica, J. A. Steyermark & J. J. Wurdack 747 (holotype, NY!).

7. Chimantaea lanocaulis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 430. 1957. TYPE: Venezuela. State Bolivar: Apacara-tepui, J. A. Steyermark 75912 (isotype, US!).

8. Chimantaea mirabilis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 432. 1957. TYPE: Venezuela. State Bolivar: Rio Tirica, J. A. Steyermark & J. J. Wurdack 821 (isotype, US!).

9. Chimantaea rupicola Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 431. 1957. TYPE: Venezuela. State Bolivar: Rio Tirica, J. A. Steyermark & J. J. Wurdack 748 (holotype, US!).

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

Quelchia N. E. Br.

Trans. Linn. Soc. London, Bot. 6: 41. 1901. TYPE: Quelchia conferta N. E. Br. 72 of Figs. 71-74.

Etymology. Named in honor of John Quelch who, together with Frederick McConnell, climbed Mount Roraima in 1894 and 1898 and collected the type material.

Shrubs or small trees monopodial to few-branched. Leaves alternate, often clustered at the stem apex; petiolate; blades oblanceolate, elliptic to obovate, coriaceous, pinnately veined, margin entire, glabrous to densely woolly beneath. Capitulescences cymose, glomerate, axillary or terminal; capitula long-pedunculate to subsessile, homogamous, one-flowered; receptacle epaleate; involucre multiseriate. Floret bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed to rarely becoming nearly bilabiate, lobes coiled at apex; anthers sometimes red-streaked, partly to mostly exserted, apical appendages acuminate, tails papillose, free; style bifid, sometimes reddish, branches dorsally smooth. Cypselae glabrous to slightly pilose; pappus of scabrid, capillary bristles, the innermost slightly apically broadened and longer.

Pollen Description. Quelchia bracteata Maguire, Steyerm. & Wurdack (117 of Figs. 117-118). Pollen subprolate, elliptic-subrectangular in equatorial view, circular in polar view, large size, P x E = (55-62 x 45-48) [micro]m. Tricolporate, colpi long with margin and microgranulate membrane. Exine Gongylolepis type, 7-10 [micro]m thick in the whole grain, spines ca. 2.5 [micro]m length. Ratio ectosexine/endosexine: 1:2; 1:2.5. Nexine ca. 2 [micro]m thick. SEM: tectum very perforate. Pollen similar to that of Duidaea marahuacensis and Glossarium rhodanthum (see also Carlquist, 1957b; Telleria, 2008).

Habitat and Distribution. Four species plus one named hybrid, all endemic to the Guayana Highland in southern Venezuela.

Species list

1. Quelchia bracteata Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 435. 1957. TYPE: Venezuela. State Bolivar: Chimanta, J. A. Steyermark & J. J. Wurdack 683 (holotype, NY!). Additional specimens examined: Venezuela. State Bolivar: Dist. Piar, macizo del Chimanta, O. Huber 11591 (US); id., J. Pipoly et al. 7217 (US*).

2. Quelchia cardonae Steyerm., Fieldiana, Bot. 28: 670. 1953. TYPE: Venezuela. State Bolivar: cumbre del cerro Acopan, Rio Caroni, Guayana, F. Cardona 2288 (holotype, US!).

3. Quelchia conferta N. E. Br., Trans. Linn. Soc. London, Bot. 6: 41. 1901. TYPE: Border of Roraima, Brazil, Guyana, and Bolivar, Venezuela: Mt. Roraima, F. McConnell & J. Quelch 652 (holotype, K!).

4. Quelchia eriocaulis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 435. 1957. TYPE: Venezuela. State Bolivar: Toroni-tepui, J. A. Steyermark & J. J. Wurdack 675 (isotype, US!).

5. Quelchia xgrandifolia Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 436. 1957. TYPE: Venezuela. State Bolivar: Chimanta Massif, Torono-tepui, J. A. Steyermark & J. J. Wurdack 756 (isotype, NY!).

Observation. Quelchia combines characteristics of Stifftieae and Mutisieae. Species of the genus have smooth style branches, and capitula with corollas that range from tubular, slightly zygomorphic, to bilabiate.

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

Stenopadus S. E Blake

Bull. Torrey Bot. Club 58: 489. 1931, emend. Maguire & Wurdack, and collaborators, Mem. New York Bot. Gard. 9: 379. 1957. TYPE: Stenopadus talaumifolius S. F. Blake. 73 of Figs. 71-74.

Etymology. From the Greek steno, narrow, and padus, foot, in reference to the narrow paleae at the "foot" of the cypselae as often seen in the type species.

Shrubs or big trees. Leaves alternate; petiolate; blades oblanceolate to orbicular, coriaceous, pinnately veined, margin entire, often revolute, usually glabrous. Capitulescences monocephalous or cymes two- to four-headed, terminal; capitula sessile to short-pedunculate, homogamous, discoid; receptacle moderately paleate to epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla actinomorphic, tubular-funnelform, deeply five-lobed, tube ten- to 15-nerved, lobes strongly coiled to less commonly flexuous; anthers sometimes green to purplish, exserted, apical appendages acuminate, tails smooth to papillose, often connate with the tails of adjacent stamens; style sometimes reddish, bilobed, branches dorsally papillose much below the bifurcation point. Cypselae glabrous; pappus of scabrid, capillary bristles to flattened and widened at the apex, the innermost longer.

Pollen Description. Stenopadus (118 of Figs. 117-118). Pollen prolate, elliptic or subrectangular in equatorial view, circular-subcircular in polar view, large size. Tricolporate, colpi long or short, with psilate or microgranulate membrane, with or without diffuse mesoaperture. Exine Stenopadus type and Mutisia type, microechinate, equally thickened in the whole grain or with polar caps, scabrate. Ratio ectosexine/endosexine: 1:1; 1:1.5 (see also Carlquist, 1957b; Barroso & Maguire, 1973; Telleria, 2008). Stenopadus chimantensis Maguire, Steyerm. & Wurdack (118A, B of Figs. 117-118). Pollen size P x E = (62-72 x 36-52) [micro]m. Colpi 45-50 [micro]m length with membrane densely microgranulate around the endoaperture. Exine Stenopadus type, with polar caps, 3-6 [micro]m thick at the equator and 7-9 [micro]m at the poles. Ratio ectosexine/endosexine: 1:1. Nexine 2-2.5 [micro]m thick. SEM: tectum psilate at the poles and very perforate with striate aspect at the equator. Pollen similar to that of Stenopadus connellii, S. jauaensis, S. sericeus, S. talaumifolius, and Stomatochaeta condensata (see also Telleria, 2008). Stenopadus connellii (N. E. Br.) S. F. Blake (118E of Figs. 117-118). Pollen size P x E = (67-75 x 42-50) [micro]m. Colpi 40-50 [micro]m length with granulate membrane. Exine Stenopadus type, psilate, 5-5.5 [micro]m thick in the whole grain. Nexine 2 [micro]m thick. SEM: tectum very perforate, with striate aspect and very small microspines. Note: some grains have more than three apertures, and pyramidal or quadrangular shape. Pollen similar to that of Stenopadus chimantensis, S. jauensis, S. sericeus, S. talaumifolius, and Stomatochaeta condensata (see also Telleria, 2008). Stenopadus huachamacari Maguire (118D of Figs. 117-118). Pollen size P x E = (52-54 x 36-43) [micro]m. Colpi long with psilate margin and scabrate membrane, mesoaperture diffuse. Exine Mutisia type, 4-6 [micro]m thick, slightly thickened at the poles. Nexine 2 [micro]m thick. SEM: tectum very perforate. Pollen similar to that of Chimantaea huberi, C. cinerea, Neblinaea promontorium, Stifftia uniflora, Stenopadus campestris, and S. colveii (see also Telleria, 2008). Stenopadus talaumifolius (118C of Figs. 117-118). Pollen size P x E = (81.5-90 x 50-61) [micro]m. Colpi 53-64 [micro]m length with membrane psilate or scarcely microgranulate. Exine Stenopadus type, scabrate, 6-8 [micro]m thick in the whole grain. Nexine ca. 2.5 [micro]m thick. Note: pollen similar to that of Stenopadus chimantensis, S. connelli, S. jauensis, S. sericeus, and Stomatochaeta condensata (see also Telleria, 2008).

Habitat and Distribution. Stenopadus is a genus of 15 species, 14 of these found in the sandstone areas of the Guayana Highland in Colombia, Venezuela, Guyana, and Brazil, and one species found in the Andes of Ecuador and Peru.

Species list

1. Stenopadus andicola Pruski, Novon 8: 67. 1998.. TYPE: Ecuador. Zamora-Chinchipe: Canton Nangaritza, valle del Rio Nangaritza, Miazi, W. Palacios 6712 (holotype, US!).

2. Stenopadus aracaensis Pruski, Ann. Missouri Bot. Gard. 76: 999. 1989. TYPE: Brazil. State Amazonas: plateau of northern massif of Serra Araca, G. T. Prance et al. 29184 (holotype, IMPA!).

3. Stenopadus campestris Maguire & Wurdack, Mem. New York Bot. Gard. 9: 383. 1957. TYPE: Venezuela. State Amazonas: Puerto Pimichin, B. Maguire & J. J. Wurdack 35578 (holotype, NY!).

4. Stenopadus chimantensis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 9: 427. 1957. TYPE: Venezuela. State Bolivar: Chimanta, Torono-tepui, J. A. Steyermark & J. J. Wurdack 652 (holotype, NY!). Additional specimen examined: Venezuela. State Bolivar: Distr. Piar, macizo del Chimanta, seccion centro del Churi-tepui, J. Pipoly et al. 7142 (MO*).

5. Stenopadus colombianus Cuatrec. & Steyerm., Bot. Mus. Leafl. 17: 99. 1955. TYPE: Colombia. Vaupes: Rio Kananari, cerro Isibukuri, R. E. Schultes & I. Cabrera 15031 (holotype, F!).

6. Stenopadus colveei (Steyerm.) Pruski, Ann. Missouri Bot. Gard. 76: 998. 1989. Basionym: Stomatochaeta colveei Steyerm. TYPE: Venezuela. State Bolivar: cumbre Cerro Guaiquinima, J. A. Steyermark et al. 117407 (holotype, VEN!).

7. Stenopadus connellii (N. E. Br.) S. F. Blake, Bull. Torrey Bot. Club 58: 490. 1931. Basionym: Stifftia connellii N. E. Br. TYPE: Border of Roraima, Brazil; Guyana; and Venezuela. Mount Roraima, F. McConnell & J. Quelch 661 (holotype, K!). Additional specimen examined: Venezuela. State Bolivar: Kukenan-tepui, R. Liesner 23109 (MO*).

8. Stenopadus cucullatus Maguire & Wurdack, Mem. New York Bot. Gard. 9: 387. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Yutaje, B. Maguire & C. K. Maguire 35053 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Dept. Atures, Carlo Yutaje, at S base of Serrania de Yutaje, Holst & R. Liesner 3043 (MO).

9. Stenopadus huachamacari Maguire, Mem. New York Bot. Gard. 8: 151. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Huachamacari, B. Maguire et al. 30026 (holotype, NY!). Additional specimen examined: Venezuela. Terr. Fed. Amazonas: Cerro Huachamacari, Rio Cunucunuma, B. Maguire et al. 30116 (MO*).

10. Stenopadus jauensis Aristeg., Acta Bot. Venez. 3: 40. 1968. TYPE: Venezuela. Meseta de Jaua: cerro Jaua, cumbre de Ika, J. A. Steyermark 97867 (isotype, NY!).

11. Stenopadus kunhardtii Maguire, Mem. New York Bot. Gard. 8: 151. 1953. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Sinapo, B. Maguire & L. Politi 28304 (holotype, US!).

12. Stenopadus megacephalus Pruski, Ann. Missouri Bot. Gard. 76: 1001. 1989. TYPE: Guayana. Upper Mazaruni River basin, Mt. Ayanganna, S. S. Tillett et al. 45100 (holotype, NY!).

13. Stenopadus obconicus Maguire & Wurdack, Mem. New York Bot. Gard. 9: 381. 1957. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro de la Neblina, B. Maguire et al. 37041 (holotype, US!).

14. Stenopadus sericeus Maguire & Aristeg., Mem. New York Bot. Gard. 9: 385. 1957. TYPE: Venezuela. Ilo-tepui: Gran Sabana, B. Maguire 33490 (holotype, NY!).

15. Stenopadus talaumifolius S. F. Blake, Bull. Torrey Bot. Club 58: 491. 1931. TYPE: Venezuela. Terr. Fed. Amazonas: Cerro Duida, G. H. H. Tate 759 (holotype US!). Additional specimen examined: Venezuela. Dept. Atapabo, Huachamacari, R. Liesner 18346 (MO*).

Literature. Maguire et al. (1957a, b), Aristeguieta (1964), Pruski (1989b, 1991, 1997, 1998), Diaz-Piedrahita and Velez-Nauer (1993), Pruski and Beltran (2003).

Stifftia J. C. Mikan

Del. fl. faun. bras. 1: t. 1. 1820. TYPE: Stifftia chrysantha J. C. Mikan. 74 of Figs. 71-74.

Aristomenia Vell., Fl. flumin.: 8. 1829 (1825). TYPE: Aristomenia fruticosa Vell. [= Stifftia fruticosa (Vell.) D. J. N. Hind & Semir].

Augusta Leandro, Denkschr. Konigl. Akad. Wiss. Munchen 7: 235. 1820, non Pohl, 1828, nom. rejic. TYPE: Augusta grandiflora Leandro [= Stifftia chrysantha J. C. Mikan].

Mocinia DC. Prodr. 7: 26. 1838. TYPE: Mocinia mutisioides DC. [= Stifftia chrysantha J. C. Mikan].

Sanhilaria Leandro ex DC., Prodr. 7: 26. 1838, nom. nudum.

Etymology. Named for C. E. Stifft, geologist and author of several works from ca. 1807-1832.

Trees shrubby or vines. Leaves alternate; petiolate; blades elliptical to oblong, chartaceous or less commonly sub-coriaceous, pinnately veined, margin entire, completely glabrous to weakly arachnoid beneath. Capitulescences corymbose to paniculate, less commonly monocephalous; capitula pedunculate, homogamous, discoid; receptacle epaleate; involucre multiseriate. Florets isomorphic, bisexual, corolla actinomorphic, tubular-funnelform, deeply fivelobed, tube five-nerved, the lobes completely coiled or only near the apex; anther apical appendages acuminate, tails smooth to sometimes papillose, free; style bilobed, branches dorsally smooth to slightly rugulose at the apex. Cypselae glabrous to weakly puberulent; pappus of scabrid bristles, capillary to somewhat paleaceous, the innermost slightly apically broadened and longer.

Pollen Description. Stifftia (Fig. 119). Pollen spheroidal, elliptic in equatorial view, circular in polar view, large size. Tricolporate, colpi long with margin, and scabrate or psilate membrane. Exine Mutisia type, microechinate, with or without polar caps. Ratio ectosexine/endosexine: 1:1 (see also Carlquist, 1957b; Telleria, 2008). Stifftia chrysantha (Fig. 119). Pollen elliptic, subtriangular in polar view, planaperturate, P x E = (66.5-85 x 58-65) [micro]m. Exine with polar caps, 8-14 [micro]m thick at the equator and 11-16 [micro]m thick at the poles, ectosexine and internal tectum very thickened at the poles. Ratio ectosexine/endosexine: 2:1; 1:1. Nexine ca. 2 [micro]m thick. SEM: tectum with very small perforations. Stifftia uniflora Ducke. Pollen spheroidal, circular in polar view. P x E = (48-60 x 42-54) [micro]m. Exine 6-6.5 [micro]m thick in the whole grain. Nexine ca. 1.5 [micro]m thick. Note: some grains lack the endoaperture. Pollen similar to that of Chimantaea huberi, C. cinerea, Neblinaea promontoriorum, Stenopadus campestris, S. huachamacari and S. colveii (see also Telleria, 2008).

Habitat and Distribution. Stifftia is a genus of seven species occurring in Brazil and French Guiana. Stifftia is the only genus of this tribe occurring both north and south of the Amazon River.

Species list

1. Stifftia cayennensis H. Rob. & B. Kahn, Phytologia 58: 248. 1985. TYPE: French Guiana. Cayenne route de Saint Laurent a Paul Isnard, entre le PK 70 et Citron, au PK 118, G. Cremers 8153 (holotype, US!).

2. Stifftia chrysantha J. C. Mikan, Del. fl. faun. bras. 1:1. 1820. TYPE: Brazil. Rio de Janeiro, s.d., Mikan s.n. (isotype, NY!). Additional specimens examined: Brazil, Hutchinson 8832 (US); Rio de Janeiro, Jardin Botanico, Cabrera 12242 (LP*).

3. Stifftia fruticosa (Vell.) D. J. N. Hind & Semir, Kew Bull. 53: 618. 1998. Basionym: Aristomenia fruticosa Vell. Icon. fl. flumin. 8: tab. 84. 1831 (1827). TYPE: Lectotype, designated by Hind & Semir, 1998a: 618, the illustration Icon. fl. flumin. 8: tab. 84. 1831 (1827).

4. Stifftia hatschbachii H. Rob., Syst. Bot. 16: 687. 1991. TYPE: Brazil. Espirito Santo: Mun. Linhares, Reserva Forestal Sooretama, G. Hatschbach et al. 53506 (isotype, US!).

5. Stifftia parviflora (Leandro) D. Don, Trans. Linn. Soc. London 16: 291. 1833. Basionym: Augusta parviflora Leandro. TYPE: Brazil. State Rio de Janeiro: near Rio de Janeiro, s.d., Leandro s.n. (type, probably P). Additional specimens examined: Brazil. State Minas Gerais: Faz. do Brejao, Jose do Alfredo, Cordesburgo, Heringer 7680 (LP*); State Sao Paulo: Piassaguera, Lemos 17110 (LP*).

6. Stifftia racemosa H. Rob., Syst. Bot. 16: 691. 1991. TYPE: Brazil. State Minas Gerais: capoeira Ibirite, municipio de Betim, G. Mendes Magalhaes 974 (holotype, US!).

7. Stifftia uniflora Ducke, Arq. Inst. Biol. Veg. 2: 72. 1935. TYPE: Brazil. State Amazonas: Rio Negro, Santa Isabel, A. Ducke 24148 (isotype, US!). Additional specimen examined: Brazil. State Amazonas: Rio Negro, Santa Isabel, A. Ducke s.n. (LP*).

Literature. Maguire et al. (1957a), Robinson and Kahn (1985), Pruski (1991), Robinson (1991), Roque and Pirani (1997), Hind and Semir (1998a).

Stomatochaeta (S. F. Blake) Maguire & Wurdack

Mem. New York Bot. Gard. 9: 388. 1957, based on Stenopadus subgen. Stomatochaeta S. F. Blake, Bull. Torrey Bot. Club 58: 490. 1931. TYPE: Stenopadus crassifolius S. F. Blake [= Stomatochaeta crassifolia (S. F. Blake) Maguire & Wurdack]. 75 of Figs. 75-76.

Dermatophyllum Sch. Bip. ex M. R. Schomb., Reis. Br.-Guiana 3: 1079. 1848 (1849), non Scheele 1848, nom. nudum.

Etymology. From the diminutive of Greek stoma, mouth, and chaite, hair, in reference to the "hairy mouth" (technically the pubescent upper portion of the corolla throat) of the corollas.

Subshrubs or trees much branched. Leaves alternate, often clustered at the stem apex; petiolate; blades narrowly spathulate, oblanceolate to obovate, coriaceous, pinnately veined, margins entire, thickened, revolute, glabrous, if pubescent, then generally rapidly glabrate. Capitulescences monocephalous, terminal; capitula sessile to subsessile, homogamous, discoid; receptacle partially paleate to epaleate; involucre multiseriate. Florets isomorphic, bisexual, actinomorphic, tubular-funnelform, very deeply five-lobed, lobes very long, stiffly erect, glabrous; anthers generally exserted, apical appendages acuminate, tails smooth to very slightly papillose, free but appressed to the tails of adjacent stamens to truly connate; style bilobed, branches dorsally papillose much below the bifurcation point. Cypselae usually glabrous; pappus of scabrid, capillary bristles, the innermost apically broadened and longer.

Pollen Description. Stomatochaeta condensata (120 of Figs. 120-121). Pollen subprolate, spheroidal-elliptic in equatorial view, circular in polar view, large size, P x E = (52-55 x 40-45) [micro]m. Tricolporate, colpi long with microgranulate membrane. Exine Stenopadus type 4-5 [micro]m thick, slightly thickened at the poles. Nexine 2 [micro]m thick. Note: pollen similar to that of Stenopadus chimantensis, S. connelli, S. jauensis, S. sericeus and S. talaumifolius (see also Carlquist, 1957b; Telleria, 2008).

Habitat and Distribution. A genus of six species, all of which are endemic to the Guayana Highland of southern Venezuela, Guyana, and northern Brazil.

Species list

1. Stomatochaeta acuminata Pruski, Brittonia 41: 35. 1989. TYPE: Venezuela. State Bolivar: Piar, summit of Angasima, O. Huber 11374 (holotype, VEN!).

2. Stomatochaeta condensata (Baker) Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. Basionym: Stifftia condensata Baker. TYPE: Guyana. Roraima, Schomburgk 543 (holotype, K not seen). Additional specimens examined: Venezuela. State Bolivar: R. Liesner 19415 (MO*); cumbre de Cerro Guaiquinima, rio Carapo, J. Steyermark & E. Dunsterville 113254 (US).

3. Stomatochaeta crassifolia (S. F. Blake) Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. Basionym: Stenopadus crassifolius S. F. Blake. TYPE: Venezuela. Terr. Fed. Amazonas: Mt. Duida, central camp., G. H. H. Tate 1014 (isotype, US!).

4. Stomatochaeta cylindrica Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. TYPE: Venezuela. State Bolivar: Cerro Guaiquinima, Rio Paragua, 1 km E of Cumbre Camp, B. Maguire 32782 (holotype, NY!).

5. Stomatochaeta cymbifolia (S. F. Blake) Maguire & Wurdack, Mem. New York Bot. Gard. 9: 389. 1957. Basionym: Stenopadus cymbifolius S. F. Blake. TYPE: Venezuela. State Bolivar: Auyan-tepui, G. H. H. Tate 1322 (holotype, NY!).

6. Stomatochaeta steyermarkii Aristeg., Acta Bot. Venez. 3: 39. 1968. TYPE: Venezuela. Bolivar: Jaua meseta, cerro Jaua, J. A. Steyermark 97917 (isotype, US!).

[FIGURES 75-76 OMITTED]

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

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 4
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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