Note: An version of this paper was published in

Arnaldoa 8(2): 25-44. 2001(2002)

TRIBAL CLASSIFICATION AND DIVERSITY

IN THE ASTERACEAE OF PERU

Michael O. Dillon
Department of Botany
The Field Museum
Chicago, IL 60605 USA
dillon@sacha.org

Abundio Sagástegui Alva
Museo de Historia Natural
Universidad Antenor Orrego
Casilla 1001
Trujillo, Perú
asagasteguia@upao.edu

 

Abstract

The Asteraceae is one of the largest families of angiosperms with +1500 genera and +20,000 species. Within the Peruvian flora it is one of the largest and most conspicuous elements and well-represented in all environments with the exception of the low-land tropics. This paper provides a synopsis of the current tribal classification of the Peruvian Asteraceae and diversity figures for South American representatives. The Peruvian Asteraceae has yielded additional records and new species since the last listing in 1993 when 222 genera and 1432 species were recorded. In the subsequent decade, the numbers have climbed to 239 genera and 1542 species.

 

We recognize 15 tribes within the Peruvian Asteraceae represented with native or naturalized representatives: Anthemideae (9 gen., ca. 15 spp.); Astereae (18 gen., ca. 170 spp.); Barnadesieae (5 gen., ca. 28 spp.); Cardueae (3 gen., 5 spp.); Eupatorieae (46 gen., ca. 325 spp.); Gnaphalineae (13 gen., ca. 50 spp.; Helenieae 5 gen., ca. 7 spp.; Heliantheae (59 gen., ca. 300 spp.; Lactuceae (8 gen., ca. 33 spp.; Liabeae (13 gen., ca. 80 spp.; Mutisieae (16 gen., ca. 90 spp.; Plucheeae (4 gen., ca. 7 spp.); Senecioneae (16 genera, ca. 340 spp.); Tageteae (5 gen., ca. 12 spp.); and Vernonieae (22 gen., ca. 70 spp.). The Calenduleae have only cultivated species (Calendula officinalis L.), and tribes Arctotoideae and Inuleae (sensu stricto) contain no native nor naturalized representatives in Peru.

 

The flora has had changes in the endemic genera present with 14 currently recognized: Ascidiogyne, Caxamarca, Ellenbergia, Holoschkuhria, Hughesia, Nothobaccharis, Uleophytum, Syncretocarpus, Bishopanthus, Chionopappus, Pseudonoseris, Chucoa, Schizotrichia, and Aynia. The genus Arnaldoa has been discovered in southern Ecuador. New, non-endemic genera for the Peruvian flora include Laestadia (Astereae), Microseris (Lactuceae), Dillandia (Liabeae), Stenopadus (Mutisieae), Talamancalia (Senecioneae), Trepadonia (Vernonieae), and Xenophyllum (Senecioneae) amongst others. A listing of recent or important literature references is provided for each tribe. A list of 113 additional species for the Peru is provided, including a website with frequent up-dates on new species:

http://www.sacha.org/Asteraceae_adiciones.htm>

 

Resumen

Las Asteráceas representan una de las familias más numerosas de angiospermas con más de1500 géneros y más de 20,000 especies. Dentro de las flora peruana, la familia Asteraceae es una de las más grandes y se distribuyen en casi todos los ambientes con excepción de la selva baja. El presente trabajo provee una sinopsis de la clasificación de las tribus actuales que componen la familia Asteraceae en el Perú así como cifras de su diversidad representada en Perú y Sudamerica. Las investigaciones recientes de las Asteráceas peruanas han producido registros adicionales y nuevas especies desde el último listado en 1993 en el cual se registran 222 géneros y 1432 especies. En la década siguiente, el número se ha elevado ha 239 géneros y 1542 especies.

 

Quince tribus están representadas por taxa nativos o naturalizados en el Perú: Anthemideae (9 gén., ca. 15 spp.); Astereae (18 gén., ca. 170 spp.); Barnadesieae (5 gén., ca. 28 spp.); Cardueae (3 gén., 5 spp); Eupatorieae (46 gén., ca. 325 spp.); Gnaphalineae (13 gén.; ca. 50 spp.); Helenieae 5 gén., ca. 7 spp.); Heliantheae (59 gén., ca. 300 spp.); Lactuceae (8 gén.; ca. 33 spp.); Liabeae (13 gén., ca. 80 spp.); Mutisieae (16 gén.; ca. 90 spp.); Plucheeae (4 gén., ca. 7 spp.); Senecioneae (16 gén. ca.. 340 spp.); Tageteae (5 gén., ca. 12 spp.); y Vernonieae (22 gén., ca. 70 spp.). La tribu Calenduleae esta representada solamente por una especie cultivada (Calendula officinalis L.) y tribus Arctotoideae e Inuleae (s.s.) no tienen ningún representante en Perú.

 

Catorce géneros son endémicos de Perú: Ascidiogyne, Caxamarca, Ellenbergia, Holoschkuhria, Hughesia, Nothobaccharis, Uleophytum, Syncretocarpus, Bishopanthus, Chionopappus, Pseudonoseris, Chucoa, Schizotrichia, y Aynia. El género Arnaldoa (Barnadesieae) ha sido descubierto en el sur de Ecuador, de la misma manera que el género Crossothamnus. Entre los nuevos géneros no endémicos para la flora peruana se incluye a Chiliotrichiopsis (Astereae), Microseris (Lactuceae), Dillandia (Liabeae), Laestadia (Astereae), Stenopadus (Mutisieae), Talamancalia (Senecioneae), Trepadonia (Vernonieae) y Xenophyllum (Senecioneae), entre otros.

Las referencias de literatura reciente y más importantes son suministradas para cada tribo. Una lista de 113 especies descubiertas desde 1993 es proveída. Esta lista cerá actualizada frecuentemente y esta disponible a través de la WWW:

<http://www.sacha.org/Asteraceae_adiciones.htm>

Introduction

The exact number of species for the Asteraceae has been variously estimated. Mabberley (1987) estimated the size of the family at 1314 genera and 21,000 species (Mabberley, 1987), Turner and Nesom (1989) calculated 1500 genera and +25,000 species (Turner & Nesom, 1989), and Bremer (1994) estimated 1535 genera and ca. 23, 000 species. South America contains high levels of generic diversity with over 450 genera or roughly 30% of the generic diversity in the entire family (Bremer 1994). Equally high levels of species diversity are common throughout South America, where the family typically represents 10% or more of the total flowering plant diversity (Turner & Nesom, 1989). The continent of South America has been suggested as a potential geographic origin of the Asteraceae (Raven & Axelrod 1974; Turner 1977; Bremer 1994) and it contains most of the diversity of the Barnadesieae (subfamily Barnadesioideae), a tribe considered basal in the Asteraceae (Bremer 1994; Jansen et al. 1991). In South America, the largest concentration of genera is to be found in the tribes Eupatorieae (ca. 116 genera), Heliantheae (ca. 90 genera), Vernonieae (ca. 57 genera), Mutisieae s.s. (ca. 54 genera), and Astereae (ca. 34 genera). Among the smallest tribes are the Barnadesieae with 9 genera and 92 species, the Liabeae with 16 genera and over 150 species, the Gnaphalineae with 20 genera and ca. 100 species, and the Plucheeae with 6 genera and 26 species.

The Asteraceae of Peru is one of the largest and most diverse families in the flora. Dillon and Hensold (1993) compiled the first detailed listing of the Asteraceae for the Peruvian flora and recorded 222 genera and ca. 1432 species. That figure lacked precision due to overlooked species. Now, after nearly 10 years, the diversity figures for the Asteraceae have changed considerably. In a recent publication, Beltrán and Baldeon (2001) reported 245 genera and 1530 species for the Peruvian Asteraceae. In the current study, we have attempted to account for newly described and validly published names, and in most cases have examined a voucher. We report here 239 genera and ca. 1542 species. Our totals would reach 245 genera or more, if we included Carthamnus, Calendula, Gerbera, Helichrysum, Madia, and Monticalia.

We recognize 15 tribes within the Peruvian Asteraceae represented with native or naturalized representatives: Anthemideae with 9 genera, ca. 15 species; Astereae with 18 genera, ca. 170 species; Barnadesieae with 5 genera, ca. 28 species; Cardueae with 3 genera, 5 species; Eupatorieae with 46 genera, ca. 325 species; Gnaphalineae with 13 genera, ca. 50 species; Helenieae with 5 genera, ca. 7 species; Heliantheae with 59 genera, ca. 300 species; Lactuceae with 8 genera, ca. 33 species; Liabeae with 13 genera, ca. 80 species; Mutisieae with 16 genera, ca. 90 species; Plucheeae with 4 genera, ca. 7 species; Senecioneae with 16 genera, ca. 340 species; Tageteae with 5 genera, ca. 12 species; and Vernonieae with 22 genera, ca. 70 species Only tribes Arctotoideae, Calenduleae (Calendula officinalis L.), and Inuleae (sensu stricto) contain no native nor naturalized representatives in Peru. While the tribe Cardueae contains only introduced species, it is included here since some species are widely distributed and naturalized.

Endemism in the Peruvian Asteraceae is high with over 750 species (ca. 50% of the total) and 14 genera considered endemic to Peru. The tribe Eupatorieae has 5 endemic genera (Ascidiogyne, Ellenbergia, Hughesia, Nothobaccharis, Uleophytum), Helenieae (Holoschkuhria), Heliantheae only one (Syncretocarpus), Liabeae has 3 endemic genera (Bishopanthus, Chionopappus, Pseudonoseris), Mutisieae only one (Chucoa), Senecioneae only one (Caxamarca), Tageteae only one (Schizotrichia), and Vernonieae only one (Aynia). The largest genus within the flora is Senecio (Senecioneae) with ca. 180 species, followed by Mikania (Eupatorieae) with ca. 84 species, Verbesina (Heliantheae) with ca. 56 species, Gynoxys (Senecioneae) with ca. 48 species, Ageratina (Eupatorieae) with 43 species, Pentacalia (Senecioneae) with ca. 40 species, and Diplostephium (Astereae) with ca. 40 species. Endemic genera are in bold-face in the lists for each tribe.

In Peru, the Asteraceae is an important member in nearly every vegetational community from ocean strand to high-elevation habitats in excess of 4500 m, and every habitat in between. Their diversity is not equally distributed and the family overall finds its greatest species and generic diversity in strongly seasonal habitats with pronounced dry/wet cycles of intermontane valleys or strongly diurnal regimes such as jalca and ceja de la montaña. Their representation in arid or semi-arid habitats is also notable and the coastal lomas formations contain + 70 Asteraceae species, many endemic (Dillon, unpubl. data). The only environments containing lower numbers of Asteraceae are the lowland rain forests where few species are found on the forest floor but lianas occupy canopy sites and others are restricted to disturbed habitats of roadsides, tree gaps or riparian sites.

With the advent of molecular techniques (Jansen & Palmer, 1987) and cladistic analysis (Bremer, 1994), the internal structure and classification of the Asteraceae has changed over the last 20 years. The family is now considered to contain three subfamilies: (1) the Barnadesioideae (tribe Barnadesieae), (2) the Cichorioideae (tribes Mutisieae, Cardueae, Lactuceae, Vernonieae, Liabeae, Arctoteae), and (3) the Asteroideae (tribes Inuleae, Plucheeae, Gnaphalieae, Calenduleae, Astereae, Anthemideae, Senecioneae, Helenieae, Heliantheae, Tageteae, Eupatorieae). The following discussion gives a summary of tribes recognized within the Peruvian flora and their generic and species diversity. We are trying to make the relevant publications on the family Asteraceae available and update the listing of tribes and genera. New species are continually being described and many more were omitted from the Catalogue of the Flowering Plants and Gymnosperms of Peru (Dillon & Hensold, 1993). For this reason, we provide a listing of some of the new records and species reported for the Peruvian flora.

We are also maintaining a webpage via the internet, where new Peruvian records and synonymy are being maintained as an ongoing resource to researchers:

<http://www.sacha.org/Asteraceae_adiciones.htm>.

References

Beltrán, H. & S. Baldeon. 2001. Adiciones a las Asteraceas del Peru. Dilloniana 1(1): 9-14.

Bremer, K. 1994. Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon. 752 pps.

Bremer, K., R.K. Jansen, P.O. Karis, M. Kallersjo, S.C. Keeley, K.-J. Kim, H.J. Michaels, J.D. Palmer, & R.S. Wallace. 1992. A review of the phylogeny and classification of the Asteraceae. Nordic Journal of Botany 12: 141-148.

Dillon, M.O. & N. Hensold. 1993. Family Asteraceae. In L. Brako & J. L. Zarucchi. Catalogue of the Flowering Plants and Gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45: 103-189.

Jansen, R. K., H.J. Michaels, & J. D. Palmer. 1991. Phylogeny and character evolution in the Asteraceae based upon chloroplast DNA restriction site mapping. Syst. Bot. 16: 98-115.

Jansen, R. K. & J. D. Palmer. 1987. A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). Proc. Natl. Acad. Sci. USA 84: 5818-5822.

Mabberley, D.J. 1987. The Plant Book, A Portable Dictionary of the Higher Plants. Cambridge University Press, Cambridge.

Raven, P.H. & D. I. Axelrod. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Garden 61: 539-673.

Turner, B.L. & G.L. Nesom. 1989. Asteraceae, the largest family of vascular plants: an extrapolation from census of the species found in Mexico and Central America. Abst. Amer. J. Bot. 76(6): 277.

 

I. ANTHEMIDEAE

Worldwide, the Anthemideae is a large tribe consisting of 109 genera and about 1,740 species (Bremer, 1994), occurring primarily in the temperate Northern Hemisphere. It is not an important tribe in the Neotropics. In Peru, the tribe is represented by 9 genera and 15 species, most of which are introduced weeds or escapees from cultivation; however, Cotula and Soliva contain native species. There are no endemic genera in Peru.

Genera: Achillea, Artemisia, Cotula, Dendranthema, Leucanthemum, Matricaria, Santolina, Soliva, Tanacetum.

References

Bremer, K. 1994. Tribe Anthemideae. Pp. 435-478. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Bremer, K., & C.J. Humphries. 1993. Generic monograph of the Asteraceae-Anthemideae. Bull. Nat. Hist. Mus. Lond. (Bot.) 23:71-177.

Cabrera, A. L. 1949. Sinopsis del género Soliva (Compositae). Notas Mus. La Plata, Bot. 14: 123-139. Caro, J. A. 1961. Las especies de Cotula (Compositae) del centro de la Repulica Argentina. Kurtiziana 1: 289-298.

Dillon, M. O. 1981. Family Compositae: Part II. Tribe Anthemideae. In J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 7, 1-21.

Heywood, V. H. & C. J. Humphries. 1977. Anthemideae - Systematic Review. In V. H. Heywood et al. (eds.), The Biology and Chemistry of the Compositae. Pp. 852-898. Academic Press, London.

 

II. ASTEREAE

Worldwide, the Astereae is one of the largest tribes in the family with an estimated 170 genera and nearly 3000 species. Centers of diversity are to be found in North America, South Africa, New Zealand and Australia. In South America, the tribe contains approximately 34 genera primarily confined to the Andean Cordillera. In Peru, the tribe is represented by 18 genera and ca. 170 species, with highest concentration of species in Baccharis with 75 species, followed by Diplostephium with 40 species. It should be noted that Plagiocheilus has been moved from Anthemideae to the Astereae, and Novenia has been moved from the Gnaphalineae to the Astereae (Nesom 1994). The description of Chiliotrichiopsis from southwestern Peru (Dept. Ayacucho) is a new interesting generic disjunction from Argentina. There are no endemic genera within the Astereae in Peru.

Genera: Baccharis, Chiliotrichiopsis, Conyza, Diplostephium, Egletes, Erigeron, Grindelia, Haplopappus, Laennecia, Laestadia, Lepidophyllum, Llerasia, Noticastrum, Novenia, Orithrophium, Parastrephia, Plagiocheilus, Psilactis.

References

Bremer, K. 1994. Tribe Astereae. Pp. 377-434. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Cuatrecasas, J. 1986. Un género nuevo de Astereae, Compositae de Colombia. Anales Jard. Bot. Madrid 42(2): 415-426.

Nesom, G. 1994. Subtribal classification of the Astereae (Asteraceae). Phytologia 76: 193-274.

Nesom, G.L., H. Robinson & A. Granda. 2001. A new species of Chiliotrichiopsis (Asteraceae: Astereae) from Peru. Brittonia 53: 430-434.

 

III. BARNADESIEAE

The Barnadesieae is a newly created tribe, but one that is quite well marked both morphologically and chemically (Bremer, 1994; Gustafsson et al., 2001; Jansen & Palmer, 1987). The distinctions are considered so great that the tribe has been placed in its own subfamily (Barnadesioideae). The members of this tribe were previously considered part of the Mutisieae and treated there in the Flora of Peru treatment (Ferreyra, 1995). The tribe contains 9 genera and 92 species and is confined to South America. In Peru, the tribe is represented by five genera and 28 species. Until recently, Arnaldoa was considered to be a Peruvian endemic; however, a new species was discovered in southern Ecuador (Ulloa et al., 2002). While Harling (1991) stated that Fulcaldea was endemic to Ecuador, it undoubtedly has Peruvian populations (Ferreyra, 1995).

Genera: Arnaldoa, Barnadesia, Chuquiraga, Dasyphyllum, Fulcaldea.

References

Bremer, K. 1994. Tribe Barnadesieae. Pp. 49-60. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Díaz-Piedrahita, S. & C. Vélez-Nauer. 1993. Revisión de las tribus Barnadesieae y Mutisieae (Asteraceae) para la Flora de Colombia. Monogr. Jard. Bot. José Celestino Mutis 1: i-xi, 1-162.

Ferreyra, R. 1995. Family Asteraceae: Part VI, Mutisieae. In J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 35: 1-101

Gustafsson, M.H.G., A. S.-R. Pepper, V.A. Albert & M. Källersjö. 2001. Molecular phylogeny of the Barnadesioideae (Asteraceae). Nord. J. Bot. 21: 149-160.

Jansen, R. K. & J. D. Palmer. 1987. A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). Proc. Natl. Acad. Sci. USA 84: 5818-5822.

Harling, G. 1991. Compositae-Mutisieae. In: G. Harling & L. Andersson (eds.), Fl. Ecuador 42: 1-105.

Stuessy, T. F. & A. Sagástegui A. 1993. Revisión de Arnaldoa (Compositae, Barnadesioideae), género endémico del norte del Perú. Arnaldoa 1(4): 9--21.

Stuessy, T. F., T. Sang & M. L. DeVore. 1996. Phylogeny and biogeography of the subfamily Barnadesioideae with implications for early evolution of the Compositae. Pp. 463-490 In D. J. N. Hind (ed.), Compositae: Systematics, Proceedings of the international Compositae conference, Kew, 1994. Vol. 1. Royal Botanic Gardens, Kew.

Ulloa-Ulloa, C., P. M. Jørgensen & M. O. Dillon. 2002. Arnaldoa argentea (Barnadesioideae: Asteraceae) a new species and a new generic record for Ecuador. Novon. 12(3): 415-149.

 

IV. CARDUEAE

Worldwide, the Cardueae contains approximately 83 genera and ca. 2500 species, and is primarily distributed throughout the Old World in North Africa and Eurasian area. In South America, the tribe is represented primarily by Old World weeds and the 3 genera and 5 species represented in Peru are all introduced and often naturalize weeds in the New World. Vouchers of Carthamnus lanatus L. (~Kentrophylum), a native of the Mediterranean region, have not been examined. Cynara scolymus L. (artichoke, alcachofa) is not usually encountered outside of cultivation.

Genera: Centaurea, Cirsium, Silybum.

References

Bremer, K. 1994. Tribe Cardueae. Pp. 112-156. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Dittrich, M. 1977. Cynareae-systematic review. In V. H. Heywood, et al. (eds.), The Biology and Chemistry of the Compositae. Pp. 999-1015. Academic Press, London.

Dillon, M. O. 1982. Family Compositae: Part IV. Tribe Cardueae. In J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 10, 1-8.

Garcia-Jacas, T Garnatje, A. Susanna, & R. Vilatersana. 2002. Tribal and subtribal delimitation and phylogeny of the Cardueae (Asteraceae): A combined nuclear and chloroplast DNA analysis. Molecular Phylogenetics & Evolution, 22: 51-64.

 

V. EUPATORIEAE

The tribe Eupatorieae is a large tribe with approximately 170-180 genera and 2400 species worldwide, with centers of diversity in Mexico, Central and South America. The South American Andes are one of the major centers of generic diversity with 116 genera recorded. In Peru, the tribe is one of the largest with 46 genera and 325 species. Five genera are considered endemic to Peru. The largest Peruvian genus is Mikania with 84 species, followed by Ageratina with 43 species, and Stevia with ca. 30 species.

Genera: Adenostemma, Ageratina, Ageratum, Amboroa, Aristeguietia. Ascidiogyne, Asplundianthus, Austroeupatorium, Ayapana, Ayapanopsis, Badilloa, Bartlettina, Brickellia, Chromolaena, Condylidium, Critonia, Critoniella, Cronquistianthus, Crossothamnus, Dasycondylus, Ellenbergia, Ferreyrella, Fleischmannia, Grosvenoria, Guevaria, Gymnocoronis, Hebeclinium, Helogyne, Heterocondylus, Hughesia, Idiothamnus, Isocarpha, Kaunia, Koanophyllon, Mikania, Neocuatrecasia, Nothobaccharis, Ophyryosporus, Phalacraea, Polyanthina, Praxelis, Raulinoreitzia, Sciadocephala, Stevia, Uleophytum, Urolepis.

References

Bremer, K., A. A. Anderberg, P. O. Karis, & J. Lundberg. 1994. Tribe Eupatorieae. Pp. 625-680. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Holmes, W. C. & S. McDaniel. 1982. Family Asteraceae: Part III. Genus Mikania, Tribe Eupatorieae. In, J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 9, 1-56.

King, R. M. & H. Robinson. 1987. The Genera of the Eupatorieae (Asteraceae). Monographs in Systematic Botany, Missouri Botanical Garden 22: 1-581.

Sánchez-Vega, I. & M. O. Dillon. 2000 (2001). Una nueva especie de Mikania (Eupatorieae: Asteraceae) de Piura, Perú. Arnaldoa 7(1-2): 7-12.

 

VI. GNAPHALINEAE

The subtribe Gnaphaliinae (sensu Anderberg) contains ca. 180 genera and ca. 2000 species, with centers of diversity in Africa, Asia, and the Neotropics. Bremer (1987) concluded that the traditional tribe Inuleae was paraphyletic and suggested that three tribes were justified. While the classification is still controversial, recent chloroplast DNA restriction site mapping studies by Jansen et al. (1990, 1991) and others (Keeley & Jansen 1991) support the hypothesis that the Gnaphalieae and Plucheeae are distinct from the Inuleae. In a series of papers, Anderberg (1989, 1991) dismantled the Inuleae (sensu lato) and treated the constituent genera in the three newly constituted tribes that roughly corresponded to the traditional subtribes: Inuleae (sensu stricto), Plucheeae, and Gnaphalieae.

The Peruvian genera formerly attributed to the tribe Inuleae (cf. Dillon & Sagástegui, 1991) are now assigned to two new tribes: Gnaphalineae with 13 genera and ca. 50 species, and Plucheeae with 4 genera and 7 species. The popular ornamental, Helichrysum bracteatum (Vent.) Andrews, is rarely encountered outside of cultivation. The distribution of Mniodes is essentially endemic to Peru (*), but there are reports of the genus from extreme northern Chile.

Genera: Achyrocline, Antennaria, Chevreulia, Cuatrecasasiella, Facelis, Gamochaeta, Jalcophila, Loricaria, Lucilia, Luciliocline, Mniodes*, Pseudognaphalium, Stuckertiella.

References

Anderberg, A. A. 1989. Phylogeny and reclassification of the tribe Inuleae (Asteraceae). Canadian J. Bot. 67: 2277-2296.

Anderberg, A. A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Bot. 104: 1-195.

Anderberg, A.A.. 1994. Tribe Gnaphalieae. Pp. 304-364. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Anderberg, A. & S. Freire. 1991. A cladistic and biogeographic analysis of the Lucilia group (Asteraceae, Gnaphalieae). J. Linn. Soc. Bot. 106: 173-198.

Dillon, M. O. & A. Sagástegui A. 1986. Jalcophila, a new genus of Andean Inuleae (Asteraceae). Brittonia 38: 162-167.

Dillon, M. O. & A. Sagástegui A. 1991. Family Asteraceae: Part V. Tribe Inuleae. In J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 26, 1-70.

Dillon, M. O. & A. Sagástegui A. 1991 (1992). Sinopsis de los Géneros de Gnaphaliinae (Asteraceae-Inuleae) de Sudamerica. Arnaldoa 1(2): 5-91.

 

VII. HELENIEAE

The tribe Helenieae has traditionally been separated from the Heliantheae by the lack of paleae on the receptacle and a pappus of scales or absent. A recent study utilizing nuclear rDNA evidence (Baldwin et al., 2002) has changed the constitution of the Helenieae. At this moment, we have chosen to place the members of the traditional tribe Helenieae Benth. & Hook. (Helenium, Hymenoxys) and the tribe Bahieae B.G. Baldwin (Schkuhria) together. Madia (Madieae Jeps.) has been reported for Peru but no voucher has been examined. Villanova Lagasca (1816) was conserved over Villanova Ortega (1797), which inturn moved Vasquezia R.A. Philippi (1860) into synonymy. In the study by Baldwin et al. (2002), the placement of Villanova is still controversial. In Peru, five genera contain seven species.

Genera: Helenium, Holoschkuhria, Hymenoxys, Schkuhria, Villanova.

References

Baldwin, B.G., B.L. Wessa, & J.L. Panero. 2002. Nuclear rDNA Evidence for Major Lineages of Helenioid Heliantheae (Compositae) Syst. Bot. 27: 161-198.

Karis, P. O. & O. Ryding. 1994. Tribe Helenieae. Pp. 521-558. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Robinson, H. Holoschkuhria, a new genus of the Hymenopappinae (Helenieae) from Peru. Compositae Newsletter 38: 47-51.

 

VIII. HELIANTHEAE

Worldwide, the Heliantheae contain 189 genera and nearly 2500 species. The highest diversity in the tribe is in the Neotropics and the Andean Cordillera. In Peru, the Heliantheae has ca. 300 species and 59 genera and marks the highest generic diversity for any tribe represented. Xanthium strumarium L. has been reported from Colombia to Paraguay and one Peruvian voucher exists at MO. The largest genera are Verbesina with 50 species, Coreopsis with 35 species and Pappobolus with 31 species.

Genera: Acanthospermum, Acanthoxanthium, Acmella, Alloispermum, Ambrosia, Aphanactis, Aspilia, Baltimora, Bidens, Blainvillia, Borrichia, Calea, Chrysanthellum, Clibadium, Complaya, Coreopsis, Cosmos, Delilia, Eclipta, Eleutheranthera, Encelia, Enhydra, Ericentrodea, Flourensia, Galinsoga, Garcilassa, Helianthus, Heliopsis, Heterosperma, Hidalgoa, Ichthyothere, Jaegeria, Lagascea, Melanthera, Milleria, Monactis, Montanoa, Neurolaena, Oblivia, Oyedaea, Pappobolus, Parthenium, Perymenium, Polymnia, Salmea, Schistocarpha, Schizoptera, Siegesbeckia, Simsia, Smallanthus, Spilanthes, Syncretocarpus, Tridax, Verbesina, Viguiera, Wedelia, Wulffia, Xanthium, Zinnia.

References

Baldwin, B.G., B.L. Wessa, & J.L. Panero. 2002. Nuclear rDNA Evidence for Major Lineages of Helenioid Heliantheae (Compositae) Syst. Bot. 27: 161-198.

Robinson, H. 1981. A revision of the tribal and subtribal limits of the Heliantheae (Asteraceae). Smithsonian Contrib. 51: 1-102.

Karis, P. O. & O. Ryding. 1994. Tribe Heliantheae. Pp. 559-624. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Stuessy, T. F. 1977. Heliantheae-systematic review. Pp. 621-671. In V.H. Heywood, J.B. Harborne & B.L. Turner (eds.), The Biology and Chemistry of the Compositae. Academic Press, London.

 

IX. LACTUCEAE

Worldwide, the Lactuceae contains 98 genera and more than 1550 species. It is primarily a Northern Hemisphere tribe with centers of diversity in the Mediterranean area, Central Asia, and southwestern North America. In Peru, the tribe is represented by 8 genera and 33 species. There are no endemic genera in Peru.

Genera: Crepis, Hieracium, Hypochoeris, Lactuca, Microseris, Picrosia, Sonchus, Taraxacum

References

Bremer, K. 1994. Tribe Lactuceae. Pp. 157-201. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

 

X. LIABEAE

The Liabeae is a well defined Neotropical tribe that contains approximately 16 genera and +180 species distributed in a wide variety of habitats throughout Mexico, Central America, West Indies (Cuba, Jamaica, Hispaniola), and Andean South America (Robinson, 1983; Funk et al. 1996). Peru contains the greatest diversity in the Liabeae, where no fewer than 14 genera and perhaps 80 species are represented. Its acceptance as a distinct tribe was not immediate and its component taxa have, until recently, been variously placed in the Vernonieae or Senecioneae. Robinson and Brettell (1973, 1974) and (Nordenstam 1977) accepted the tribe and positioned it near the Vernonieae. Phylogenetic studies of Bremer (1987, 1994) and Jansen et al. (1991) have supported the monophyly of the tribe and its placement near the Vernonieae. Funk and Robinson (2001) have recently described the new genus, Dillandia, largely based upon its nuclear ribosomal DNA sequence data and a suite of morphological characters.

Genera: Bishopanthus, Cacosmia, Chionopappus, Chrysactinium, Dillandia, Erato, Ferreyranthus, Liabum, Munnozia, Oligactis, Paranephelius, Philoglossa, Pseudonoseris.

References

Bremer, K. 1994. Tribe Liabeae. Pp. 234-245. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Dillon, M. O., & A. Sagástegui A. 1995. Estudios en la tribu Liabeae (Asteraceae) en Peru: 1. Revision de Ferreyranthus. Arnaldoa 2(2): 7-23.

Dillon, M. O., & A. Sagástegui A. 1995. Estudios en la tribu Liabeae (Asteraceae) en Peru: 2. Una nueva especie de Oligactis procedente del norte del Peru y sur del Ecuador. Arnaldoa 2(2): 25-30.

Funk, V. A. & H. Robinson. 2001. A bully new genus from the Andes (Compositae: Liabeae). Sys. Bot. 26: 216-225.

Funk, V. A., & M. F. Zermoglio. 1999. A revision of Chrysactinium (Compositae: Liabeae). Syst. Bot. 24: 323-338.

Funk, V. A., H. Robinson, & M. O. Dillon. 1996. Liabeae: Taxonomy, Phylogeny and Biogeography. In D.J.N. Hind & H.J. Beentje (eds.). Compositae: Systematics. Proceedings of the International Compositae Conference, Kew. 1994. Vol. 1. Pp. 545-567. Royal Botanic Gardens, Kew.

Nordenstam, B. 1977. Senecioneae and Liabeae -- Systematic review. Pp. 799-830. In The Biology and Chemistry of the Compositae, eds. V. H. Heywood, J. B. Harborne, and B. L. Turner. London: Academic Press.

Robinson, H. 1978. 190 (2), Compositae -- Liabeae. Flora of Ecuador, 8: 1-62.

Robinson, H. 1983. A generic review of the tribe Liabeae (Asteraceae). Smithsonian Contr. Bot. 54: 1-69.

Robinson, H. & R. D. Brettell. 1973. Tribal revisions in the Asteraceae. III. A new tribe, Liabeae. Phytologia 25: 104-107.

Robinson, H. & R. D. Brettell. 1974. Studies in the Liabeae (Asteraceae), II: Preliminary survey of the genera. Phytologia 28: 43-63.

Sagastegui-A., A., & M.O. Dillon. 1995. Estudios en la tribu Liabeae (Asteraceae) en Peru: 3. Una nueva especie de Chrysactinium del norte del Peru. Arnaldoa 2(2): 31-35.

 

XI. MUTISIEAE

The Mutisieae is primarily a Neotropical tribe with 76 genera and 970 species and greatest diversity in austral South America. In Peru, the tribe contains 16 genera and ca. 90 species. The removal of the genera now treated in the Barnadesieae is one of the more recent changes in this tribe. Gerbera jamesonii Bojus ex Hook.f. (Transvaal daisy) is a popular ornamental. The rare genus Chucoa, known from only two collections, was recently re-collected near the type locality in Prov. Santiago de Chuco, Dept. La Libertad (Sagástegui et al. 16626, HAO). Stenopadus andicola Pruski has been reported from Peru (Beltrán & Pruski, 2002); a photograph has been seen, but no voucher.

Genera: Chaetanthera, Chaptalia, Chucoa, Gochnatia, Jungia, Leucheria, Lophopappus, Lycoseris, Mutisia, Onoseris, Perezia, Plazia, Polyachyrus, Proustia, Stenopadus, Trichocline, Trixis.

References

Beltrán, H., & J. F. Pruski. 2002. Stenopadus andicola (Asteraceae: Mutisieae): un nuevo registro genérico para la flora del Perú. Abst. VIII CONABOT, p. 112.

Bremer, K. 1994. Tribe Mutisieae. Pp. 71-111. In K. Bremer (ed.), Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Cabrera, A. L. 1977. Mutisieae - Systematic review. Pp. 141-248. In V. H. Heywood, et al. (eds.), The Biology and Chemistry of the Compositae, Academic Press, London.

Ferreyra, R. 1995. Family Asteraceae: Part VI. Tribe Mutisieae. In J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 35, 1-101.

Harling, G. 1991. 190(10). Compositae - Mutisieae. Pp. 1-106. In G. Harling & L. Andersson (eds.), Flora of Ecuador, no. 42, Swedish Natural Science Research Council, Stockholm.

 

XII. PLUCHEEAE

Worldwide, the tribe Plucheeae contains 28 genera and ca. 220 species. It is a relatively small group in South America and represented by six genera containing perhaps 36 species. In Peru, the tribe is represented by 4 genera and 7 species. This tribe finds its greatest generic and species diversity in essentially low or mid-elevation habitats of austral South America. This is in sharp contrast to the Gnaphalieae which finds its greatest species diversity in predominately high-elevation genera of the Andean Cordillera.

Genera: Pluchea, Pseudoconyza, Pterocaulon, Tessaria.

References

Anderberg, A. A. 1991. Taxonomy and phylogeny of the tribe Plucheeae (Asteraceae). Pl. Sys. Evol. 176: 145-177.

Anderberg, A. A. 1994. Tribe Plucheeae. Pp. 292-303. In K. Bremer (ed.) Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Nesom, G. L. 1989. New species, new sections, and a taxonomic overview of American Pluchea (Compositae: Inuleae). Phytologia, 67: 158-167.

 

XIII. SENECIONEAE

Worldwide, the Senecioneae contains 120 genera and more than 3000 species. It is an important tribe in South America and represented in Peru by 16 genera and ca. 340 species. Senecio with ca. 180 species is the largest genus in the Peruvian flora (Vision & Dillon, 1996). Other large genera include Gynoxys with 48 species and Pentacalia with 40 species. No endemic genera are known from Peru.

Genera: Aequatorium, Caxamarca, Chersodoma, Dendrophorbium, Dorobaea, Erechtites, Gynoxys, Lasiocephalus, Misbrookia, Paracalia, Pentacalia, Pseudogynoxys, Senecio, Talamancalia, Werneria, Xenophyllum.

References

Beltran, H. & J.F. Pruski. 2000. Talamancalia y Rolandra (Asteraceae): dos nuevos registros para el Peru. Arnaldoa 7(1-2): 13-18.

Bremer, K. 1994. Tribe Senecioneae. Pp. 479-520. In Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Dillon, M. O. & A. Sagástegui A. 1999. Caxamarca, A New Monotypic Genus of Senecionieae (Asteraceae) from Northern Peru. Novon 9(2): 156-161.

Nordenstam, B. 1977. Senecioneae - Systematic Review. In V. H. Heywood, et al (eds.), The Biology and Chemistry of the Compositae, Academic Press, London.

Nordenstam, B.1978. Taxonomic studies in the tribe Senecioneae (Compositae). Opera Bot. 44: 1-83.

Nordenstam, B. & J. F. Pruski. 1995. Additions to Dorobaea and Talamancalia (Compositae: Senecioneae). Compositae Newsl. 27: 31-42.

Vision, T. J. & M. O. Dillon. Sinopsis de Senecio L. (Senecioneae, Asteraceae) para el Perú. Arnaldoa 4 (1): 23-46. 1996.

 

XIV. TAGETEAE

The Tageteae contain 34 genera and +250 species native to the New World. The tribe is characterized by highly dissected leaves and phyllaries with pellucid secretory cavities producing aromatic oils (Strother, 1977). In the treatment by Karis & Ryding (1994), its members were placed in the tribe Helenieae. The results of Baldwin et al. (2002) re-establish the validity of the tribe. In Peru, it is represented by 5 genera and 12 species with one endemic genus.

Genera: Flaveria, Pectis, Porophyllum, Schizotrichia, Tagetes.

References

Baldwin, B.G., B.L. Wessa, & J.L. Panero. 2002. Nuclear rDNA Evidence for Major Lineages of Helenioid Heliantheae (Compositae) Syst. Bot. 27: 161-198.

Karis, P. O., & O. Ryding 1994. Tribe Helenieae. Pp. 521-558. In K. Bremer (ed.) Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Robinson, H. 1981. A Revision of the Tribal and Subtribal Limits of the Heliantheae (Asteraceae). Smithsonian Contrib. Bot. 51:1-102.

Strother, J. L. 1977. Tageteae -systematic review. In V. H. Heywood, et al. (eds.), The Biology and Chemistry of the Compositae Pp. 769-783. Academic Press, London.

Turner, B.L. y A. M. Powell. 1977. Helenieae-systematic review. Pp. 699-737. In V.H. Heywood, J.B. Harborne y B.L. Turner (eds.), The Biology and Chemistry of the Compositae. Academic Press, London.

 

XV. VERNONIEAE

Worldwide, the tribe Vernonieae contains ca. 70 genera and 1500 species, most of which are in the core genus Vernonia. The tribe is distributed throughout the New and Old World tropics and extends into temperate North America. In Peru, the tribe is represented by 22 genera and 70 species. The genus Vernonia, before its fragmentation, contained ca. 38 species. H. Robinson (1989, 1996, 1999) has created and re-instated segregate genera for elements formerly treated as Vernonia (Dillon & Sagástegui, 1998).

Genera: Aynia, Centrantheum, Chrysolaena, Critoniopsis, Cuatrecasanthus, Cyanthillium, Cyrtocymura, Eirmocephala, Elephantopus, Lepidaploa, Lessingianthus, Quechualia, Pacourina, Piptocarpha, Pollalesta, Pseudelephantopus, Rolandra, Struchium, Trichospira, Trepadonia, Vernonanthura, Xiphochaeta (~Stilpnopappus).

References

Bremer, K. 1994. Tribe Vernonieae. Pp. 202-233. In Asteraceae, Cladistics and Classification. Timber Press, Portland, Oregon.

Dillon, M. O. 1982. Family Compositae: Additions to the Tribe Vernonieae. In J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 11, 1-7.

Dillon, M. O. & A. Sagástegui A. 1998. Una breve revisión del género Vernonia (sensu lato) del Perú. Arnaldoa 5(1): 25-33.

Jones, S. B. 1977. Vernonieae--Systematic review. Pp. 503-521. In V. H. Heywood, etal. (eds), The Biology and Chemistry of the Compositae, Academic Press, London.

Jones. S. B. 1980. Family Compositae: Tribe Vernonieae. In J. Francis Macbride & Collaborators, Flora of Peru, Fieldiana: Botany, N.S. 5, 22-73.

Robinson, H. 1989. Two new genera of Vernonieae (Asteraceae) from the northern Andes with dissected corolla limbs, Cuatrecasanthus and Joseanthus. Rev. Acad. Colomb. Cienc. 17(65): 207-213.

Robinson, H. 1996. The status of generic and subtribal revisions in the Vernonieae. In D.J.N. Hind & H.J. Beentje (eds.). Compositae: Systematics. Proceedings of the International Compositae Conference, Key, 1994. Vol. 1. Pp. 511-529. Royal Botanic Gardens, Kew.

Robinson, H. 1999. Generic and subtribal classification of American Vernonieae. Smithsonian Contrib. Bot. 89: 1-116.

Robinson. H. & H. Beltrán. 2000. A new species of Trepadonia (Asteraceae: Vernonieae) from Peru. Sida, Contrib. Bot. 19: 111-113

Sagástegui-Alva, A. & M. O. Dillon 1998. Una nueva especie de Critoniopsis (Vernonieae: Asteraceae) de Cajamarca, Perú. Arnaldoa 5(1): 19-24.

Sagástegui-Alva, A. & M. O. Dillon. 2001. Una nueva especie de Critoniopsis (Vernonieae: Asteraceae) del Norte de Perú. Arnaldoa 8(1): 25-35.

 

Appendix 1. Adiciones del Asteraceae Peruana

 

The Catalogue of the Flowering Plants and Gymnosperms of Peru (Brako & Zarucchi 1993) was a benchmark in the recounting of Peruvian floristic diversity. Since its publication in 1993, additions and omissions to the flora have been encountered (Beltrán & Baldeon, 2001). The taxa listed here are not found in Brako & Zarucchi (1993) and are provided with the proper author citation as a resource to researchers. No judgment as to their taxonomic validity is implied or intended. New genera are in boldface.

Aphanactis hutchisonii H. Rob.

Ayapanonsis wurdackiana H. Rob.

Baccharis articulata Pers.

Baccharis jhonwurdackiana H. Rob.

Chaptalia exscapa (Pers.) Baker

Caxamarca M.O.Dillon & Sagást.

Caxamarca sanchezii M.O.Dillon & Sagást.

Chersodoma deltoidea Sagást. & M.O.Dillon

Chiliotrichiopsis peruviana Nesom, H. Rob. & Granda

Chrysactinium breviscapum M.O.Dillon & Sagást.

Chrysactinium wurdackii Zermoglio & V.A.Funk

Chuquiraga raimondiana Granda

Chuquiraga oblongifolia Sagást. & Sánchez

Coreopsis canescentifolia Sagást.

Coreopsis dentifolia Sánchez, Sagást. & Crawford

Coreopsis dilloniana Sánchez, Sagást. & Crawford

Coreopsis ferreyrae Sagást & Sánchez

Coreopsis helleborifolia Sánchez, Sagást. & Crawford

Critoniopsis ayabacensis Sagást. & M.O.Dillon

Critoniopsis boliviana (Britton) H.Rob.

Critoniopsis gynoxiifolia H.Rob.

Critoniopsis huairacajana (Hieron.) H.Rob.

Critoniopsis oblongifolia Sagást. & M.O.Dillon

Critoniopsis quilloensis H.Rob. Delilia biflora Kuntze

Dendrophorbium multinerve (Schultz-Bip.) C.Jeffrey (~Senecio)

Dillandia V.A.Funk & H.Rob.

Dillandia chachapoyensis (H.Rob.) V.A.Funk & H.Rob. (~Munnozia)

Dillandia subumbellata V.A.Funk & H.Rob.

Diplostephium rupestre (Kunth) Wedd.

Dorobaea laciniata (Kunth) B. Nord & J. Pruski

Emilia fosbergii Nicolson

Ferreyranthus gentryii H. Rob.

Gamochaeta lulioana S.E.Freire & Iharlegui

Gochnatia lanceolata H. Beltrán & Ferreyre

Gynoxys foliosa (Rusby) S.F.Blake

Holoschkuhria H.Rob.

Holoschkuhria tetramera H.Rob.

Ichthyothere macdanielii H.Rob.

Lactuca sativa L.

Laestadia lechleri (Schultz-Bip.) Wedd.

Laestadia muscicola Wedd.

Lepidaploa sanmartinensis H.Rob.

Lessingianthus coriaceus (Less.) H. Rob.

Microseris pygmaea D. Don.

Mikania hensoldiana Sánchez & M.O.Dillon

Munnozia luyensis H.Rob.

Mutisia mandoniana Wedd. ex Cabr.

Neurolaena lobata H. Rob.

Oligactis cuatrecasasii M.O.Dillon & Sagást.

Parthenium hysterophorus L.

Pentacalia brittonia (Hieron.) Cuatrec.

Pentacalia cutervonis Cuatrec. & H.Rob.

Pentacalia loretensis Cuatrec. (~Senecio)

Pentacalia maynasensis Cuatrec. & H.Rob.

Pentacalia mucronatifolia Cuatrec. & H.Rob.

Pentacalia nunezii Cuatrec. & H.Rob.

Pentacalia sagasteguii Cuatrec. & H.Rob.

Pentacalia tilletii Cuatre. & H.Rob.

Pentacalia todziae Cuatrec. & H.Rob.

Pentacalia vargasiana Cabr. (~Gynoxys)

Perymenium celendianum B.L.Turner

Perymenium huascaranum B.L.Turner

Pluchea symphytifolia (Miller) Gillis

Parthenium hysteriophorus L.

Pluchea symphytifolia (Miller) Gillis

Rolandra fruticola (L.) Kuntze

Sciadocephala schultze-rhonhofiae Matff.

Senecio algens Wedd.

Senecio ancashinus Cabr.

Senecio burkartii Cabr.

Senecio cantensis Cabr.

Senecio castanaefolius DC.

Senecio danai A. Gray

Senecio dombeyana DC.

Senecio expansus Wedd.

Senecio hohenackeri Schultz-Bip.

Senecio icaensis H.Beltrán & Galan de Mera

Senecio larahuinensis H.Beltrán & Galan de Mera

Senecio pascoensis Cabr.

Senecio pflanzii (Perkens) Cuatrec.

Senecio pininchense Cuatrec. (~Ecuador)

Senecio praeruptorium Schultz-Bip.

Senecio pseudodiscoides Schultz-Bip.(~Chile)

Senecio recurvatus Kunth

Senecio sericens (DC.) S.F. Blake

Senecio steinbachianus Cuatrec.

Solidago chilensis Meyen

Stenopadus andicola Pruski

Talamancalia putcalensis (Hieron.) B. Nord. & J. Pruski.

Trepadonia oppositifolia H. Rob. & H. Beltrán

Trichocline caulescens Phil.

Tridax cajamarcensis H.Rob.

Trixis montesecoensis Sagást. & M.O.Dillon

Verbesina albissima Sagást.

Verbesina ampliatifolia Sagást. & Quipuscoa

Verbesina ancashensis Sagást. & Quipuscoa

Verbesina aypatensis Sagást. & Quipuscoa

Verbesina brevilingua Sagást.

Verbesina brunnea Sagást. & Quipuscoa

Verbesina capituliparva Sagást.

Verbesina chachapoyensis Sagást. & Quipuscoa

Verbesina crassicephala Sagást. & Quipuscoa

Verbesina citrina Sagást. & Zapata

Verbesina huancabambae Sagást. & Quipuscoa

Verbesina leivae Sagást. & Quipuscoa

Verbesina monactinoides Sagást., Leiva & Lezama

Verbesina otuzcensis Sagást. & Quipuscoa

Verbesina pauciramea Sagást., Leiva & Lezama

Verbesina perlanata Sagást. & Quipuscoa

Xenophyllum poposum (Phil.) V.A.Funk

Xenophyllum staffordiae (Sandw.) V.A.Funk

Xenophyllum lycopodioides (S.F.Blake) V.A.Funk

Xenophyllum weddellii (Phil.) V.A.Funk

Werneria spathulata Wedd.