Abstract
The subfamily Barnadesioideae (Compositae) is endemic to South America, comprising 10 genera and 80 species of mostly spiny herbs, subshrubs, shrubs, trees, or woody vines distributed from Venezuela to Argentina. Three genera, Dasyphyllum (27 species), Chuquiraga (22 spp.) and Barnadesia (19 spp.) contain 85% of the species, while the other seven genera (Archidasyphyllum, Arnaldoa, Doniophyton, Duseniella, Fulcaldea, Huarpea, and Schlechtendalia) are represented by up to three species each. Most species are found in xeric areas in the Andean and Patagonian regions—as in the Páramos, Puna and Patagonian steppe vegetation—with a secondary center of diversity in eastern South America. Previous phylogenetic hypotheses have clarified the relationships within the subfamily, showing that there are many non-monophyletic groups in different taxonomic ranks. As a result, taxonomic changes have been proposed over recent decades in order to reflect classifications comprising only monophyletic groups. In the present study, we provide a generic synopsis of the subfamily Barnadesioideae based on the most recent generic circumscriptions, including a key, expanded morphological descriptions, information on geographical distribution and habitat, photographs and taxonomic notes for all genera.
References
<p>Bohm, B.A. & Stuessy, T.F. (1995) Flavonoid chemistry of Barnadesioideae (Asteraceae). <em>Systematic Botany</em> 20: 22–27. https://doi.org/10.2307/2419629</p>
<p>Bohm, B.A. & Stuessy, T.F. (2001) <em>Flavonoids of the Sunflower Family (Asteraceae)</em>. Springer-Verlag Wien, Vienna, 831 pp. https://doi.org/10.1007/978-3-7091-6181-4</p>
<p>Bremer, K. (1994) <em>Asteraceae: cladistics and classification</em>. Timber Press, Portland, 752 pp.</p>
<p>Bremer, K. & Jansen, R.K. (1992) A new subfamily of the Asteraceae. <em>Annals of the Missouri Botanical Garden </em>79: 414–415. https://doi.org/10.2307/2399777</p>
<p>Cabrera, A.L. (1951) <em>Huarpea</em>, nuevo genero de Compuestas. <em>Boletín de la Sociedad Argentina de Botánica</em> 4: 129–132.</p>
<p>Cabrera, A.L. (1959) Revisión del género <em>Dasyphyllum </em>(Compositae). <em>Revista</em> <em>del Museo de</em> <em>La Plata,</em> <em>Sección botánica</em> 38: 21–100.</p>
<p>Cabrera, A.L. (1961) Compuestas Argentinas. Clave para la determinación de los géneros. <em>Revista del Museo Argentino de Ciencias Naturales. Botánicas</em> 2: 291–362.</p>
<p>Cabrera, A.L. (1962) Compuestas andinas nuevas. <em>Boletín de la Sociedad Argentina de Botánica</em> 10: 21–45.</p>
<p>Cabrera, A.L. (1977) Mutisieae—systematic review. <em>In</em>: Heywood, V.H., Harborne, J.B. & Turner, B.L. (Eds.) <em>The Biology and Chemistry of the Compositae</em>, vol. 2. Academic Press, London, pp. 1039–1066.</p>
<p>Candolle, A.P. de (1838) <em>Prodromus Systematis Naturalis Regni Vegetabilis</em>, vol. 7. Treuttel & Würtz, Paris, 801 pp.</p>
<p>Ccana-Ccapatinta, G.V., Monge, M., Ferreira, P.L. & Da Costa, F.B. (2018) Chemistry and medicinal uses of the subfamily Barnadesioideae (Asteraceae). <em>Phytochemistry Reviews</em> 17: 471–489. https://doi.org/10.1007/s11101-017-9544-y</p>
<p>Chung, I.C. (1965) <em>Revision of</em> <em>Barnadesia</em>. Published by the author, Chicago, Illinois.</p>
<p>Cialdella, A.M. & López de Kiesling, A.G. (1981) Cariología de <em>Schlechtendalia luzulaefolia</em> (Compositae). <em>Darwiniana</em> 23: 357–360.</p>
<p>Cristóbal, C.L. (1986) El número cromosómico de dos Compositae–Mutisieae. <em>Boletín de la Sociedad Argentina de Botánica</em> 24: 363–380.</p>
<p>DeVore, M.L., Zhao, Z., Jansen, R.K. & Skvarla, J.J. (2000) Utility of trends in pollen morphology for phylogenetic analyses: an example using subfamilies Barnadesioideae and Cichorioideae (Asteraceae). <em>In</em>: Harley, M.M., Morton, C.M. & Blackmore, S. (Eds.) <em>Pollen and Spores: morphology and biology</em>. Kew Royal Botanic Gardens, London, pp. 399–412.</p>
<p>Diers, L. (1961) Der Anteil an Polyploiden in den Vegetationsgürteln der Westkordillera Perus. <em>Zeitschrift für Botanik</em> 49: 437–488.</p>
<p>Don, D. (1830) Descriptions of the new Genera and Species of the Class Compositae belonging to the Floras of Peru, Mexico, and Chile. <em>Transactions of the Linnean Society of London </em>16: 169–304. https://doi.org/10.1111/j.1095-8339.1829.tb00136.x</p>
<p>Erbar, C. & Leins, P. (2000) Some interesting features in the capitulum and flower of <em>Arnaldoa macbrideana</em> Ferreyra (Asteraceae, Barnadesioideae). <em>Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie</em> 122: 517–537.</p>
<p>Ezcurra, C. (1985) Revisión del género <em>Chuquiraga</em> (Compositae–Mutisieae). <em>Darwiniana</em> 26: 219–284.</p>
<p>Ferreira, P.L., Saavedra, M.M. & Groppo, M. (2019) Phylogeny and circumscription of <em>Dasyphyllum</em> (Asteraceae: Barnadesioideae) based on molecular data with the recognition of a new genus, <em>Archidasyphyllum</em>. <em>PeerJ</em> 7: e6475. https://doi.org/10.7717/peerj.6475</p>
<p>Ferreyra, R. (1995) Family Asteraceae: Part IV, Tribe Mutiseae. <em>In</em>: Macbride, J.F. <em>et al.</em> (Eds.) <em>Flora of Peru</em>. Fieldiana Botany. N.S. 35: 1–101.</p>
<p>Funk, V.A. & Roque, N. (2011) The monotypic Andean genus <em>Fulcaldea</em> (Compositae, Barnadesioideae) gains a new species from Northeastern Brazil. <em>Taxon</em> 60: 1095–1103. https://doi.org/10.1002/tax.604012</p>
<p>Grace, O.M., Pérez-Escobar, O.A., Lucas, E.J., Vorontsova, M.S., Lewis, G.P., Walker, B.E., Lohmann, L.G., Knapp, S., Wilkie, P., Särkinen, T., Darbyshire, I., Lughadha, E.N., Monro, A., Woudstra, Y., Demissew, S., Muasya, A.M., Díaz, S., Baker, W.J. & Antonelli, A. (2021) Botanical Monography in the Anthropocene. <em>Trends in Plant Science</em> 26: 433–441. https://doi.org/10.1016/j.tplants.2020.12.018</p>
<p>Granda, P.A. (1997) Uma nueva espécie de <em>Chuquiraga</em> (Asteraceae-Mutiseae) del Perú. <em>Kurtziana </em>25: 151–156.</p>
<p>Gruenstaeudl, M., Urtubey, E., Jansen, R.K., Samuel, R., Barfuss, M.H.J. & Stuessy, T.F. (2009) Phylogeny of Barnadesioideae (Asteraceae) inferred from DNA sequence data and morphology. <em>Molecular Phylogenetics and Evolution</em> 51: 572–587. https://doi.org/10.1016/j.ympev.2009.01.023</p>
<p>Gustafsson, M.H.G., Pepper, A.S.R., Albert, V.A. & Källersjö, M. (2001) Molecular phylogeny of the Barnadesioideae (Asteraceae). <em>Nordic Journal of Botany</em> 21: 149–160. https://doi.org/10.1111/j.1756-1051.2001.tb01352.x</p>
<p>Hansen, H.V. (1991) Phylogenetic studies in Compositae tribe Mutisieae. <em>Opera Botanica </em>109: 1–50.</p>
<p>Harling, G. (1991) Compositae-Mutisieae. <em>In</em>: Harling, G. & Andersson, L. (Eds.) <em>Flora of Ecuador</em>. University of Gothenburg, Gothenburg, pp. 1–105.</p>
<p>Heiser, C.B. (1963) Numeración cromosómica de plantas ecuatorianas. <em>Ciencia y Naturaleza </em>6: 2–6.</p>
<p>Hind, D.J.N. (2001) A New Species of <em>Barnadesia</em> (Compositae: Barnadesieae) from Bolivia. <em>Kew Bulletin</em> 56: 705–710. https://doi.org/10.2307/4117698</p>
<p>Hoffmann, O. (1893) Tubuliflorae–Mutisieae. <em>In</em>: Engler, A. & Prantl, K. (Eds.) <em>Die natürlichen Pflanzenfamilien</em>, vol. 4 (5). Engelmann, Leipzig, pp. 333–350.</p>
<p>Jansen, R.K. & Palmer, J.D. (1987) Chloroplast DNA from lettuce and <em>Barnadesia</em> (Asteraceae): structure, gene localization, and characterization of a large inversion. <em>Current Genetics</em> 11: 553–564. https://doi.org/10.1007/BF00384619</p>
<p>Jansen, R.K. & Palmer, J.D. (1988) Phylogenetic implications of Chloroplast DNA restriction site variation in the Mustiseae (Asteraceae). <em>American Journal of Botany</em> 75: 753–766. https://doi.org/10.1002/j.1537-2197.1988.tb13496.x</p>
<p>Jansen, R.K., Michaels, H.J., Wallace, R.S., Kim, K-J., Keeley, S.C., Watson, L.E. & Palmer, J.D. (1992) Chloroplast DNA Variation in the Asteraceae: Phylogenetic and Evolutionary Implications. <em>In</em>: Soltis, P.S., Soltis, D.E. & Doyle, J.J. (Eds.) <em>Molecular Systematics of Plants</em>. Springer, Boston, pp. 252–279. https://doi.org/10.1007/978-1-4615-3276-7_11</p>
<p>Jussieu, A.L. (1789) <em>Genera Plantarum Secundum Ordines Naturales Disposita</em>. Herrisant & Barrois, Paris, 498 pp.</p>
<p>Katinas, L. & Stuessy, T.F. (1997) Revision of <em>Doniophyton</em> (Compositae, Barnadesioideae). <em>Plant Systematics and Evolution</em> 206: 33–45. https://doi.org/10.1007/BF00987939</p>
<p>Kunth, C.S. (1820) Compositae. <em>In</em>: Humboldt, F.W.H.A., Bonpland, A.J.A. & Kunth, K.S. (Eds.) <em>Nova genera et species plantarum</em>, vol. 4. Librariae Graeco-Latino-Germanico, Paris, 312 pp. + 113 tab.</p>
<p>Lessing, C.P. (1832) <em>Synopsis generum compositarum</em>. Duncker & Humblot, Berlin.</p>
<p>Mandel, J.R., Dikow, R.B., Siniscalchi, C.M., Thapa, R., Watson, L.E. & Funk, V.A. (2019) A fully resolved backbone phylogeny reveals numerous dispersals and explosive diversifications throughout the history of Asteraceae. <em>Proceedings of the National Academy of Sciences</em> 116: 14083 LP-14088. https://doi.org/10.1073/pnas.1903871116</p>
<p>Mendiondo, M.E., Juárez, B.E. & Seeligmann, P. (1997) Flavonoid patterns of some Barnadesioideae (Asteraceae). Eventual chemosystematic significance. <em>Biochemical Systematics and Ecology</em> 25: 673–674. https://doi.org/10.1016/S0305-1978(97)00060-4</p>
<p>Mendiondo, M.E., Juárez, B.E. & Seeligmann, P. (2000) Flavonoid profiles of some Argentine species of <em>Chuquiraga</em> (Asteraceae). <em>Biochemical Systematics and Ecology </em>28: 283–285. https://doi.org/10.1016/S0305-1978(99)00061-7</p>
<p>Mutis, J.C.B. (1781) Polyadelphia Pentandria. <em>In</em>: Linné, C. (Ed.) <em>Supplementum Plantarum</em>. Brunsvigae, pp. 55.</p>
<p>Nabhan, A.R. & Sarkar, I.N. (2012) The impact of taxon sampling on phylogenetic inference: a review of two decades of controversy. <em>Briefings in bioinformatics</em> 13: 122–134. https://doi.org/10.1093/bib/bbr014</p>
<p>Nakajima, J.N., Dematteis, M., Loeuille, B., Teles, A.M., Heiden, G., Schneider, A., Ritter, M., Oliveira, C.T., Hattori, E.K.O., Roque, N., Ferreira, S.C., Magenta, M., Bringel, J.B.A., Esteves, R., Almeida, G.S.S., Saavedra, M.M., Monge, M., Soares, N.P., Sancho, G., Mondin, C.A., Fernandes, A.C., Pereira, A.C.M., Kutschenko, D.C., Filho, L.A.F.S., Prieto, P.V., Borges, R.A.X., Penedo, T.S.A., Messina, T., Moraes, M.M.V., Moraes, M.A. & Coelho, M.A.C. (2013) Asteraceae. <em>In</em>: Martinelli, G. & Moraes, M.A. (Eds.) <em>Livro vermelho da Flora do Brasil</em>. Centro Nacional de Conservação da Flora, Rio de Janeiro, pp. 203–286.</p>
<p>Olsen, J. (1980) Chromosome number reports LXVII. <em>Taxon</em> 29: 366–367. https://doi.org/10.1002/j.1996-8175.1980.tb00607.x</p>
<p>Padin, A.L., Calviño, C.I. & Ezcurra, C. (2015a) Morfología y anatomía foliar comparada de <em>Chuquiraga</em> y géneros afines (Asteraceae). <em>Brittonia</em> 67: 150–165. https://doi.org/10.1007/s12228-015-9364-6</p>
<p>Padin, A.L., Calviño, C.I. & Ezcurra, C. (2015b) Molecular Phylogeny of <em>Chuquiraga</em> (Asteraceae-Barnadesioideae): Infrageneric Classification and Generic Affinities. <em>Systematic Botany</em> 40: 316–326. https://doi.org/10.1600/036364415X686602</p>
<p>Poeppig, E.F. & Endlicher, S.L. (1835–1836) <em>Nova genera ac species plantarum</em>, vol. 1. F. Hofmeister, Leipzig, 62 pp.</p>
<p>Poiret, J.L.M. (1817) <em>Fulcaldea</em> [under <em>Turpinia</em>]. <em>In</em>: Lamarck, M. & Poiret, J.L.M. (Eds.) <em>Encyclopedie Methodique,</em> Botanique, suppl., vol. 5. Paris, pp. 575.</p>
<p>QGIS Development Team (2019) QGIS Geographic Information System, version 3.4. Open Source Geospatial Foundation Project. Available from: http://qgis.osgeo.org (accessed 20 February 2019)</p>
<p>Radford, A.E., Dickson, W.C., Massey, W.C. & Bell, C.R. (1974) <em>Vascular Plant Systematics</em>. Harper & Row, New York, 891 pp.</p>
<p>Saavedra, M.M. (2011) <em>Sistemática de Dasyphyllum (Asteraceae)</em>. Phd Thesis. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro, pp. 1–247.</p>
<p>Saavedra, M.M. (2020) <em>Dasyphyllum</em> in Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB5291 (accessed 20 March 2021)</p>
<p>Saavedra, M.M., Guimarães, E.F., Loeuille, B. & Forzza, R.C. (2018) Taxonomic Revision of <em>Dasyphyllum</em> sect. <em>Macrocephala</em> (Asteraceae: Barnadesioideae). <em>Systematic Botany</em> 43: 297–315. https://doi.org/10.1600/036364418X696888</p>
<p>Saavedra, M.M., Monge, M., Guimarães, E.F. (2014) <em>Dasyphyllum diamantinense</em> (Asteraceae, Barnadesioideae): A new species from the Chapada Diamantina, Bahia State, Brazil. <em>Phytotaxa</em> 174 (4): 231–236. https://doi.org/10.11646/phytotaxa.174.4.4</p>
<p>Sagástegui, A.A. (1980) Compuestas andinas peruanas nuevas para la ciencia. <em>Boletín de la Sociedad Argentina de Botánica</em> 19 (1–2): 61–68.</p>
<p>Sagástegui, A.A. & Dillon, M. (1985) Four new species of Asteraceae from Peru. <em>Brittonia </em>37 (1): 6–13. https://doi.org/10.1007/BF02809659</p>
<p>Sagástegui, A.A. & Sánchez, V.I. (1991) Una nueva espécie de <em>Chuquiraga</em> (Asteraceae-Mutisieae) del Norte del Peru. <em>Arnaldoa</em> 2: 1–4.</p>
<p>Schumann, K.M. (1902) Neue Arten des Siphonogamen. <em>Just’s botanischer Jahresbericht</em> 28 (Abt. 1): 410–498.</p>
<p>Skvarla, J.J., Turner, B.L., Patel, V.C. & Tomb, A.S. (1977) Pollen morphology in the Compositae and in morphologically related families. <em>In</em>: Heywood, V.H., Harborne, J.B. & Turner, B.L. (Eds.) <em>The Biology and Chemistry of the Compositae</em>. Academic Press, London, pp. 141–248.</p>
<p>Strother, J.L. & Panero, J.L. (1994) Chromosome studies: Latin American Compositae. <em>American Journal of Botany</em> 81: 335–577. https://doi.org/10.2307/2445657</p>
<p>Stuessy, T.F. & Sagástegui, A.A. (1993) Revision de <em>Arnaldoa</em> (Compositae, Barnadesioideae) género endémico del norte del Peru. <em>Arnaldoa </em>1: 9–21.</p>
<p>Stuessy, T.F. & Urtubey, E. (2006) Phylogenetic implications of corolla morphology in subfamily Barnadesioideae (Asteraceae). <em>Flora: Morphology, Distribution, Functional Ecology of Plants</em> 201: 340–352. https://doi.org/10.1016/j.flora.2005.07.009</p>
<p>Stuessy, T.F. & Urtubey, E. (2007) Barnadesieae. <em>In</em>: Kadereit, J.W. & Jeffrey, C. (Eds.) Kubtizki, K., <em>The Families and genera of vascular plants</em>, vol. 8. Springer-Verlag, Berlin & Heidelberg, pp. 87–90.</p>
<p>Stuessy, T.F., Sang, T. & DeVore, M.L. (1996) Phylogeny and biogeography of the subfamily Barnadesioideae with implications for early evolution of the Compositae. <em>In</em>: Hind, D.J.H. & Beentje, H.J. (Eds.)<em> Compositae: Systematics</em>. <em>Proceedings of the International Compositae Conference, Kew, 1994</em>, vol. 1. Royal Botanical Garden, Kew, pp. 463–490.</p>
<p>Stuessy, T.F., Urtubey, E. & Gruenstaeudl, M. (2009) Barnadesieae (Barnadesioideae). <em>In</em>: Funk, V.A., Susanna, A., Stuessy, T. & Bayer, R.J. (Eds.) <em>Systematics, evolution, and biogeography of Compositae</em>. IAPT, Vienna, pp. 215–228.</p>
<p>Svoma, E., Mayer, V., Stuessy, T.F. & Urtubey, E. (2019) Staminal features in Barnadesioideae (Asteraceae): description, evolution and function. <em>Botanical Journal of the Linnean Society</em> 192: 474–497. https://doi.org/10.1093/botlinnean/boz091</p>
<p>Tellería, MC., Palazzesi, L. & Barreda, V. (2015) Evolutionary significance of exine ultrastructure in the subfamily Barnadesioideae (Asteraceae) in the light of molecular phylogenetics. <em>Review of Palaeobotany and Palynology</em> 221: 32–46. https://doi.org/10.1016/j.revpalbo.2015.05.008</p>
<p>Thiers, B. (2019) Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 20 February 2019)</p>
<p>Ulloa Ulloa, C., Jørgensen, P.M. & Dillon, M.O. (2002) <em>Arnaldoa argentea </em>(Barnadesioideae: Asteraceae), a new species and a new generic record for Ecuador. <em>Novon</em> 12: 415–419. https://doi.org/10.2307/3393091</p>
<p>Urtubey, E. & Stuessy, T.F. (2001) New hypotheses of phylogenetic relationships in Barnadesioideae (Asteraceae) based on morphology. <em>Taxon</em> 50: 1043–1066. https://doi.org/10.2307/1224720</p>
<p>Urtubey, E. (1997) Morfología del pólen de <em>Barnadesia</em> (Asteraceae, Barnadesioideae). <em>Boletín de la Sociedad Argentina de Botánica</em> 33: 69–75.</p>
<p>Urtubey, E. (1999) Revisión del género <em>Barnadesia </em>(Barnadesioideae, Asteraceae). <em>Annals of the Missouri Botanical Garden </em>86: 57–111. https://doi.org/10.2307/2666218</p>
<p>Urtubey, E. & Tellería, M.C. (1998) Pollen morphology of the subfamily Barnadesioideae (Asteraceae) and its phylogenetic and taxonomic significance. <em>Review of Palaeobotany and Palynology</em> 104: 19–37. https://doi.org/10.1016/S0034-6667(98)00049-9</p>
<p>Watanabe, K., Yahara, T., Hashimoto, G., Nagatani, Y., Soejima, A., Kawahara, T. & Nakazawa, M. (2007) Chromosome numbers and karyotypes in Asteraceae. <em>Annals of the Missouri Botanical Garden</em> 94: 643–654. https://doi.org/10.3417/0026-6493(2007)94[643:CNAKIA]2.0.CO;2</p>
<p>Weddell, H.A. (1855) <em>Chloris andina: essai d’une flore de la región alpine des Cordillères de l’Amérique de Sud.</em> Bertrand, Paris. https://doi.org/10.5962/bhl.title.217</p>
<p>Wulff, A.F. (1984) Estudios cromosómicos en Compuestas de las floras patagónica y bonaerense. <em>Darwiniana</em> 25: 17–26.</p>
<p>Wulff, A.F. (1990) Estudios cromosómicos en Barnadesiinae (Mutisieae, Asteraceae). I. <em>Chuquiraga</em> y <em>Doniophyton</em>. <em>Darwiniana</em> 30: 185–193.</p>
<p>Zhao, Z., Skvarla, J.J., Jansen, R.K. & DeVore, M.L. (2000) Phylogenetic implications of pollen morphology and ultrastructure in the Barnadesioideae (Asteraceae). <em>Lundellia</em> 3: 26–40. https://doi.org/10.25224/1097-993X-3.1.26</p>