Baccharis Systematics (Astereae) Gustavo Heiden & José Rubens Pirani Universidade de São Paulo, Brasil Baccharis macrophylla Dusén Compositae: 12 subfamilies, more than 40 tribes (Funk et al. 2009, Funk & Bonifacino 2010) Subfamily Asteroideae Tribe Astereae Monophyletic (Brouillet et al., 2009; Funk et al., 2005; Goertzen et al., 2003; Jansen et al., 1991; Kim et al., 1992; Kim & Jansen, 1995; Lane et al., 1996; Noyes & Rieseberg, 1999; Panero & Funk, 2002, 2008; Zhang & Bremer, 1993) Astereae: 18 subtribes, 222 genera, ~3.100 spp. (Brouillet et al., 2009; Nesom & Robinson, 2006) 31 genera and ~740 spp. in South America Ecaudate and ecalcarate anther bases Two distinct marginal stigmatic lines and deltate to triangular or lanceolate style appendages, glabrous adaxially and with sweeping hairs abaxially Astereae in Brazil: 8 subtribes, 16 genera, ~220 spp. (120 endemic) Baccharidinae Chrysopsidinae Conyzinae Hysterionica 3 spp. Noticastrum 8 spp. Baccharis 160 spp. Neja 3 spp. A.A.Schneider Apopyros 2 spp. Grangeinae Machaerantherinae Conyza 12 spp. Leptostelma 4 spp. Solidaginae www.cnip.org.br www.cnip.org.br Egletes 1 sp. Podocominae Grindelia 7 spp. Symphyotrichinae Solidago 1 sp. Podocoma 6 spp. Inulopsis 4 spp. Sommerfeltia 1 sp. Asteropsis 1 sp. Microgyne 1 sp. Symphyotrichum 5 spp. Astereae: Baccharidinae Variability in the sexual systems Homogamous or heterogamous capitula, not in the same plant Archibaccharis Heering Baccharis L. 32 spp. ~400 spp. www.sbs.utexas.edu Archibaccharis schiedeana (Benth.) J. Jacks Baccharis leucocephala Dusén Monoecious Dioecious Flowers hermaphrodite and unisexual Flowers unisexual Cypselae 2-ribbed, flattened Cypselae >5-ribbed, cylindrical Baccharis L. Southeastern Canada (Nova Scotia) and Western USA (Oregon) to Eastern Brazil and Southern Tierra del Fuego in Argentina and Chile Baccharis salicifolia (Ruiz & Pav.) Pers. Baccharis ciliata Gardner Several widespread and many narrow endemic species Baccharis >90% of the species occurs in South America Two mains centers of diversity: Andes and Southeastern South America Mainly in grasslands, savannas and montane vegetation Baccharis From the sea level to the summits of mountains above 5000 m... Baccharis arenaria Baker Baccharis uncinella DC. Meireles E. ...the 400 species of Baccharis occur in a great diversity of environments, playing an important role in several plant communities Baccharis Herbaceous, subshrubby, shrubby, arboreal and lianescent species lianescent and arboreal shrubby, subshrubby, Herbaceous, B. gracilis DC. reflects in a diverse array of morphological morphological diverse array a of in reflects The wide range of environments inhabited inhabited environments of range wide The features within genus the within features B. riograndensis Teodoro & Vidal B. granndimucronata Teodoro B. longiattenuata A.S.Oliveira Baccharis junciformis DC. B. anomala DC. Baccharis Useful... Baccharis dracunculifolia DC. (Lam.) Pers. (Lam.) B. articulataB. ...or weeds “caused the death of 18 big bulls” DC. Baccharis coridifolia DC. Baccharis coridifolia Baccharis Baccharis Since Candolle (1836), Baccharis has never been revised taxonomically as a whole Ir. Teodoro Luis / Malagarriga Heras (1977) Nomenclator Baccharidinarum omnium Müller (2010) World checklist of Baccharis L. (Compositae-Astereae) Baccharis Most of the modern taxonomic studies on the genus are geographically limited Venezuela Steyermark 1953 Colombia Aristeguieta 1964 Cuatrecasas 1967, 1969 Bolivia Brazil Müller 2006 Barroso 1976 Oliveira et. al. 2006 Heiden & Schneider 2010, 2011 Uruguay Chile Arechavaleta 1906-1908 Heering 1905 Hellwig 1990 Argentina Giuliano & Ariza Espinar 1999 Giuliano 2000, 2005 Baccharis s.s.: Hellwig (1993, 1996) Segregation into new, reinstated or recircumscribed smaller genera Baccharis Neomolina F.H.Hellw. L. (Lam.)Pers. (Ruiz & Pav.) Pers. & (Ruiz Pav.) Hook. & Arn.) F.H.Hellw. Hook.& Arn.) B. magellanicaB. B. linearisB. Pingraea ( racemosa Neomolina Cass. Stephananthus Lehm. Lehm. (DC.) F.H.Hellw. (DC.) (Lam.) F.H.Hellw. (Lam.) Pingraea anomala Pingraea Pingraea articulataPingraea Stephananthus junceusStephananthus Baccharis: traditional circumscription Baccharis (~400 spp.) Heterothalamus (3 spp.) ? Baccharidastrum (2 spp.) Baccharidiopsis (1 sp.) Baccharis fide Nesom (1988) Baccharis (~400 spp.) Heterothalamus (3 spp.) ? Baccharis sect. Baccharidastrum (2 spp.) Baccharidiopsis (1 sp.) Baccharis s.l.: Müller (2006) Baccharis (~400 spp.) Baccharis subgen. Molina sect. Heterothalamus (3 spp.) ? Baccharis subgen. Molina sect. Baccharidastrum (2 spp.) Baccharis subgen. Pteronioides [incl. Baccharidiopsis = B. hirta] Baccharis s.l.: what support the adoption of a “broad” circumscription? Baccharis macrophylla Dusén Tufted indumentum with adjoining basal cells Volkens (1890) Volkens Florets functionally unisexual Baccharis s.l.: infrageneric relationships? ? Given the great diversity and wide geographic distribution of Baccharis, the recognition of smaller and morphologically well-circumscribed infrageneric groups is critical to achieve strong advances in taxonomy Baccharis s.l.: infrageneric taxonomy Candolle (1836): 8 sections Followed by Weddel (1855), Baker (1882) and Hoffmann (1890) Gray (1886): 4 informal groups Influenced Heering (1903, 1904a, 1904b): 5 subgenera, several sections and subsections Cuatrecasas (1967, 1982): circumscription of 24 sections based mainly on Colombian species Influenced Ariza (1973), Nesom (1988, 1990), Giuliano (2001, 2005) and Giuliano & Nesom (2003) Barroso (1976): 15 informal groups, similar to the sections of Cuatrecasas Müller (2006): 4 subgenera with several informal groups and 3 groups not ascertain to a subgenus Giuliano (2011): followed Müller subgenera, described 15 new sections + 3 new combinations to section level ***several sections and groups are monotypic*** Search for all infrageneric names published Provisional placement into the described subgenera, sections, subsections, series and informal groups for further testing of monophyly of the proposed groups Baccharis subgen. Baccharis • 14 to 20 sections • ~200 species • USA (Oregon) and Canada (Nova Scotia) to Patagonia DC. B. brevifolia brevifolia B. Wawra Müller 2007 Müller B. itatiaiae itatiaiae B. Male floret with style apex bearing sweeping hairs of unequal size DC. Female corolla 5-dentate Female pappus multiseriate and deciduous B. uncinella uncinella B. Cypselae glabrous Baccharis subgen. Molina Heering • 16 to 18 sections • ~100 species • Southern USA to Patagonia (Lam.)Pers. Baker B. serrulata B. B. glaziovii glaziovii B. Müller 2007 Müller Male floret with style apex bearing sweeping hairs of equal size Female corolla truncate or up to 5-dentate (F.H.Hellw.) Joch. Mull. Joch. (F.H.Hellw.) Female pappus uniseriate and not acrescent Cypselae glabrous or with twin hairs B. wagenitzii wagenitzii B. Baccharis subgen. Pteronioides Heering • 8 to 10 sections • ~60 species • Southern USA to Patagonia Baker Baccharis maxima maxima Baccharis Müller 2006 Müller Male floret with style apex bearing sweeping hairs of equal size DC. Female pappus multiseriate, acrescent and persistent Cypselae large, brownish to dark, with uniseriate hairs Baccharis gracilis gracilis Baccharis Baccharis subgen. Stephananthus (Lehm.) Heering • 2 sections • 4 species • Southern Peru to Southern Brazil and Patagonia (Lehm.)Desf. Müller 2006 Müller Leaves fleshy Baccharis juncea Baccharis Male floret with style apex bearing Baker sweeping hairs of equal size Female pappus multiseriate, acrescent and persistent Cypselae with twin hairs Baccharis arenaria Baccharis Baccharis subgen. Tarchonanthoides Heering • 4 to 6 sections • ~20 species • Bolivia and Southeastern Brazil to Central Argentina Heering MalmeHeering ex Hook. & Arn. Hook.& Baccharis curytibensis Baccharis Müller 2006 Müller Baccharis artemisioides Baccharis Cup-shaped involucre of male capitula Baker Female floret with apically broadened and 5-dentate corolla Female pappus multiseriate, acrescent and persistent Baccharis gibertii Baccharis Cypselae with twin hairs Baccharidinae and Baccharis phylogenetic relationships Zanowiak (1991): cpDNA RFLP markers Baccharidinae and Baccharis phylogenetic relationships Lane et al. (1996): cpDNA RFLP markers Baccharidinae and Baccharis phylogenetic relationships Karaman-Castro & Urbatsch (2009): ITS Baccharidinae and Baccharis phylogenetic relationships Brouillet et al. (2009): ITS Two of five subgenera and 1.5% of the Baccharis species sampled Current knowledge based on a limited dataset does not provide enough data American lineages South on the intergeneric relationships of genus Baccharis, its monophyly and the relationships of its infrageneric groups Hence, a comprehensive phylogenetic study within the genus is in need to refuse or corroborate its alleged monophyly Prospects Include more outgroups Sample at least one species per infrageneric group, preferably the type one Teodoro Reduce the geographic bias of sampling Search for more informative sequences Consider including morphological data pseudovaccinioides Baccharis The future recognition of smaller monophyletic and morphologically well circumscribed groups will make possible the advance of taxonomic knowledge on the genus, free of geographic limitations Assembling the big picture of the Baccharis phylogeny will provide answers on the diversification, the wide morphological diversity and the evolution of sexual expression within one of the largest genera of Compositae .