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Baccharis Systematics (Astereae) - Fapesp

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<strong>Baccharis</strong> <strong>Systematics</strong> (<strong>Astereae</strong>)<br />

Gustavo Heiden & José Rubens Pirani<br />

Universidade de São Paulo, Brasil<br />

<strong>Baccharis</strong> macrophylla Dusén


Compositae: 12 subfamilies, more than 40 tribes (Funk et al. 2009, Funk & Bonifacino 2010)<br />

Subfamily Asteroideae<br />

Tribe <strong>Astereae</strong><br />

Monophyletic<br />

(Brouillet et al., 2009; Funk et al., 2005;<br />

Goertzen et al., 2003; Jansen et al., 1991; Kim<br />

et al., 1992; Kim & Jansen, 1995; Lane et al.,<br />

1996; Noyes & Rieseberg, 1999; Panero &<br />

Funk, 2002, 2008; Zhang & Bremer, 1993)


<strong>Astereae</strong>: 18 subtribes, 222 genera, ~3.100 spp. (Brouillet et al., 2009; Nesom & Robinson, 2006)<br />

31 genera and ~740 spp. in South America<br />

Ecaudate and ecalcarate anther bases<br />

Two distinct marginal stigmatic lines and deltate<br />

to triangular or lanceolate style appendages,<br />

glabrous adaxially and with sweeping hairs<br />

abaxially


<strong>Astereae</strong> in Brazil: 8 subtribes, 16 genera, ~220 spp. (120 endemic)<br />

Baccharidinae<br />

<strong>Baccharis</strong> 160 spp.<br />

Machaerantherinae<br />

Grindelia 7 spp.<br />

Asteropsis 1 sp.<br />

Conyzinae<br />

Apopyros 2 spp.<br />

Podocominae<br />

Inulopsis 4 spp.<br />

A.A.Schneider<br />

Conyza 12 spp.<br />

Microgyne 1 sp.<br />

Hysterionica 3 spp.<br />

Podocoma 6 spp.<br />

Leptostelma 4 spp.<br />

Neja 3 spp.<br />

Sommerfeltia 1 sp.<br />

Solidaginae<br />

Solidago 1 sp.<br />

Chrysopsidinae<br />

Noticastrum 8 spp.<br />

Grangeinae<br />

Egletes 1 sp.<br />

Symphyotrichinae<br />

Symphyotrichum 5 spp.<br />

www.cnip.org.br


<strong>Astereae</strong>: Baccharidinae<br />

Variability in the sexual systems<br />

Homogamous or heterogamous capitula, not in the same plant<br />

Archibaccharis Heering<br />

32 spp.<br />

Archibaccharis schiedeana (Benth.) J. Jacks<br />

Monoecious<br />

Flowers hermaphrodite and unisexual<br />

Cypselae 2-ribbed, flattened<br />

www.sbs.utexas.edu<br />

<strong>Baccharis</strong> L.<br />

~400 spp.<br />

Dioecious<br />

Flowers unisexual<br />

Cypselae >5-ribbed, cylindrical<br />

<strong>Baccharis</strong> leucocephala Dusén


<strong>Baccharis</strong> L.<br />

Southeastern Canada (Nova Scotia) and Western USA (Oregon) to<br />

Eastern Brazil and Southern Tierra del Fuego in Argentina and Chile<br />

<strong>Baccharis</strong> salicifolia (Ruiz & Pav.) Pers.<br />

<strong>Baccharis</strong> ciliata Gardner<br />

Several widespread and many narrow endemic species


<strong>Baccharis</strong><br />

>90% of the species occurs in South America<br />

Two mains centers of diversity: Andes and Southeastern South America<br />

Mainly in grasslands, savannas and montane vegetation


<strong>Baccharis</strong><br />

From the sea level to the summits of mountains above 5000 m...<br />

<strong>Baccharis</strong> arenaria Baker <strong>Baccharis</strong> uncinella DC.<br />

...the 400 species of <strong>Baccharis</strong> occur in a great diversity of environments,<br />

playing an important role in several plant communities<br />

E. Meireles


<strong>Baccharis</strong><br />

B. gracilis DC.<br />

The wide range of environments inhabited<br />

reflects in a diverse array of morphological<br />

features within the genus<br />

B. riograndensis Teodoro & Vidal<br />

Herbaceous, subshrubby, shrubby, arboreal and lianescent species<br />

B. granndimucronata Teodoro<br />

B. longiattenuata A.S.Oliveira<br />

<strong>Baccharis</strong> junciformis DC.<br />

B. anomala DC.


<strong>Baccharis</strong><br />

Useful...<br />

B. articulata (Lam.) Pers.<br />

...or weeds<br />

“caused the death of 18 big bulls”<br />

<strong>Baccharis</strong> coridifolia DC.<br />

<strong>Baccharis</strong> dracunculifolia DC.<br />

<strong>Baccharis</strong> coridifolia DC.


<strong>Baccharis</strong><br />

Since Candolle (1836), <strong>Baccharis</strong> has never been<br />

revised taxonomically as a whole<br />

Müller (2010)<br />

World checklist of <strong>Baccharis</strong> L.<br />

(Compositae-<strong>Astereae</strong>)<br />

Ir. Teodoro Luis / Malagarriga Heras<br />

(1977)<br />

Nomenclator Baccharidinarum<br />

omnium


<strong>Baccharis</strong><br />

Most of the modern taxonomic studies on the genus are geographically limited<br />

Colombia<br />

Cuatrecasas 1967, 1969<br />

Bolivia<br />

Müller 2006<br />

Chile<br />

Heering 1905<br />

Hellwig 1990<br />

Venezuela<br />

Steyermark 1953<br />

Aristeguieta 1964<br />

Brazil<br />

Barroso 1976<br />

Oliveira et. al. 2006<br />

Heiden & Schneider 2010, 2011<br />

Uruguay<br />

Arechavaleta 1906-1908<br />

Argentina<br />

Giuliano & Ariza Espinar 1999<br />

Giuliano 2000, 2005


<strong>Baccharis</strong> s.s.: Hellwig (1993, 1996)<br />

<strong>Baccharis</strong> L.<br />

Pingraea Cass.<br />

Segregation into new, reinstated or recircumscribed smaller genera<br />

Pingraea anomala (DC.) F.H.Hellw.<br />

B. magellanica (Lam.) Pers.<br />

Pingraea articulata (Lam.) F.H.Hellw.<br />

B. linearis (Ruiz & Pav.) Pers.<br />

Neomolina F.H.Hellw.<br />

Stephananthus Lehm.<br />

Stephananthus junceus Lehm.<br />

Neomolina racemosa (Hook. & Arn.) F.H.Hellw.


<strong>Baccharis</strong>: traditional circumscription<br />

<strong>Baccharis</strong> (~400 spp.)<br />

Baccharidastrum (2 spp.)<br />

?<br />

Heterothalamus (3 spp.)<br />

Baccharidiopsis (1 sp.)


<strong>Baccharis</strong> fide Nesom (1988)<br />

<strong>Baccharis</strong> (~400 spp.)<br />

<strong>Baccharis</strong> sect. Baccharidastrum (2 spp.)<br />

?<br />

Heterothalamus (3 spp.)<br />

Baccharidiopsis (1 sp.)


<strong>Baccharis</strong> s.l.: Müller (2006)<br />

<strong>Baccharis</strong> (~400 spp.)<br />

<strong>Baccharis</strong> subgen. Molina sect. Baccharidastrum (2 spp.)<br />

<strong>Baccharis</strong> subgen. Molina sect. Heterothalamus (3 spp.)<br />

?<br />

<strong>Baccharis</strong> subgen. Pteronioides [incl. Baccharidiopsis = B. hirta]


<strong>Baccharis</strong> s.l.: what support the adoption of a “broad” circumscription?<br />

<strong>Baccharis</strong> macrophylla Dusén<br />

Tufted indumentum<br />

with adjoining<br />

basal cells<br />

Volkens (1890)<br />

Florets functionally unisexual


<strong>Baccharis</strong> s.l.: infrageneric relationships?<br />

?<br />

Given the great diversity and wide geographic distribution of <strong>Baccharis</strong>,<br />

the recognition of smaller and morphologically well-circumscribed<br />

infrageneric groups is critical to achieve strong advances in taxonomy


<strong>Baccharis</strong> s.l.: infrageneric taxonomy<br />

Candolle (1836): 8 sections<br />

�Followed by Weddel (1855), Baker (1882) and Hoffmann (1890)<br />

Gray (1886): 4 informal groups � Influenced Heering (1903, 1904a, 1904b): 5 subgenera, several<br />

sections and subsections<br />

Cuatrecasas (1967, 1982): circumscription of 24 sections based mainly on Colombian species<br />

�Influenced Ariza (1973), Nesom (1988, 1990), Giuliano (2001, 2005) and Giuliano & Nesom (2003)<br />

Barroso (1976): 15 informal groups, similar to the sections of Cuatrecasas<br />

Müller (2006): 4 subgenera with several informal groups and 3 groups not ascertain to a subgenus<br />

Giuliano (2011): followed Müller subgenera, described 15 new sections + 3 new combinations to<br />

section level<br />

***several sections and groups are monotypic***<br />

Search for all infrageneric names published<br />

Provisional placement into the described subgenera, sections,<br />

subsections, series and informal groups for further testing of<br />

monophyly of the proposed groups


<strong>Baccharis</strong> subgen. <strong>Baccharis</strong><br />

• 14 to 20 sections<br />

• ~200 species<br />

• USA (Oregon) and Canada (Nova Scotia) to<br />

Patagonia<br />

B. itatiaiae Wawra<br />

B. uncinella DC.<br />

B. brevifolia DC.<br />

�Male floret with style apex bearing<br />

sweeping hairs of unequal size<br />

�Female corolla 5-dentate<br />

�Female pappus multiseriate and<br />

deciduous<br />

�Cypselae glabrous<br />

Müller 2007


<strong>Baccharis</strong> subgen. Molina Heering<br />

B. glaziovii Baker<br />

• 16 to 18 sections<br />

• ~100 species<br />

• Southern USA to Patagonia<br />

B. wagenitzii (F.H.Hellw.) Joch. Mull.<br />

B. serrulata (Lam.) Pers.<br />

�Male floret with style apex bearing<br />

sweeping hairs of equal size<br />

�Female corolla truncate or up to 5-dentate<br />

�Female pappus uniseriate and not<br />

acrescent<br />

�Cypselae glabrous or with twin hairs<br />

Müller 2007


<strong>Baccharis</strong> subgen. Pteronioides Heering<br />

• 8 to 10 sections<br />

• ~60 species<br />

• Southern USA to Patagonia<br />

<strong>Baccharis</strong> maxima Baker<br />

<strong>Baccharis</strong> gracilis DC.<br />

�Male floret with style apex bearing<br />

sweeping hairs of equal size<br />

�Female pappus multiseriate, acrescent<br />

and persistent<br />

�Cypselae large, brownish to dark, with<br />

uniseriate hairs<br />

Müller 2006


<strong>Baccharis</strong> subgen. Stephananthus (Lehm.) Heering<br />

• 2 sections<br />

• 4 species<br />

• Southern Peru to Southern Brazil and Patagonia<br />

<strong>Baccharis</strong> arenaria Baker<br />

<strong>Baccharis</strong> juncea (Lehm.) Desf.<br />

�Leaves fleshy<br />

�Male floret with style apex bearing<br />

sweeping hairs of equal size<br />

�Female pappus multiseriate, acrescent<br />

and persistent<br />

�Cypselae with twin hairs<br />

Müller 2006


<strong>Baccharis</strong> subgen. Tarchonanthoides Heering<br />

• 4 to 6 sections<br />

• ~20 species<br />

• Bolivia and Southeastern Brazil to Central<br />

Argentina<br />

<strong>Baccharis</strong> curytibensis Heering ex Malme<br />

<strong>Baccharis</strong> gibertii Baker<br />

<strong>Baccharis</strong> artemisioides Hook. & Arn.<br />

�Cup-shaped involucre of male capitula<br />

�Female floret with apically broadened and<br />

5-dentate corolla<br />

�Female pappus multiseriate, acrescent<br />

and persistent<br />

�Cypselae with twin hairs<br />

Müller 2006


Baccharidinae and <strong>Baccharis</strong> phylogenetic relationships<br />

Zanowiak (1991): cpDNA RFLP markers


Baccharidinae and <strong>Baccharis</strong> phylogenetic relationships<br />

Lane et al. (1996): cpDNA RFLP markers


Baccharidinae and <strong>Baccharis</strong> phylogenetic relationships<br />

Karaman-Castro & Urbatsch (2009): ITS


Baccharidinae and <strong>Baccharis</strong> phylogenetic relationships<br />

Brouillet et al. (2009): ITS<br />

Two of five subgenera and 1.5% of the<br />

<strong>Baccharis</strong> species sampled<br />

Current knowledge based on a limited<br />

dataset does not provide enough data<br />

on the intergeneric relationships of<br />

genus <strong>Baccharis</strong>, its monophyly and the<br />

relationships of its infrageneric groups<br />

Hence, a comprehensive phylogenetic<br />

study within the genus is in need to<br />

refuse or corroborate its alleged<br />

monophyly<br />

South American lineages


Prospects<br />

�Include more outgroups<br />

�Sample at least one species per infrageneric<br />

group, preferably the type one<br />

�Reduce the geographic bias of sampling<br />

�Search for more informative sequences<br />

�Consider including morphological data<br />

The future recognition of smaller monophyletic and morphologically well<br />

circumscribed groups will make possible the advance of taxonomic knowledge<br />

on the genus, free of geographic limitations<br />

Assembling the big picture of the <strong>Baccharis</strong> phylogeny will provide answers on the<br />

diversification, the wide morphological diversity and the evolution of sexual<br />

expression within one of the largest genera of Compositae<br />

<strong>Baccharis</strong> pseudovaccinioides Teodoro

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