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
DC.
The wide range of environments inhabited reflects in a diverse array of morphological
features within the genus junciformis Baccharis
DC.
DC.
Teodoro
A.S.Oliveira
B. gracilis B.
B. anomala B.
Teodoro & Vidal & Teodoro
B. granndimucronata B.
B. longiattenuata B. B. riograndensis B.
Herbaceous, subshrubby, shrubby, arboreal and lianescent species Baccharis
Useful...
Baccharis dracunculifolia DC.
(Lam.) Pers. (Lam.)
B. articulata B.
...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 L. Neomolina F.H.Hellw.
(Lam.)Pers.
(Ruiz & Pav.) Pers. & (Ruiz Pav.)
Hook. & Arn.) F.H.Hellw. Hook.Arn.) &
B. magellanica B. B. linearis B.
Pingraea Cass. ( racemosa Neomolina
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.(Lam.)
Baker
B. serrulata 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 ex Malme Heering 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 Americanlineages 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