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Pollen Morphology of the Genus Eremanthus Less. (Vernonieae, Asteraceae)

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Pollen Morphology of the Genus Eremanthus Less. (Vernonieae, Asteraceae) Acta Botanica Brasilica 26(1): 46-57. 2012. Pollen morphology of the genus Eremanthus Less. (Vernonieae, Asteraceae) Benoît Loeuille1, Raquel Maria Batista Souza-Souza2, Vanessa Holanda Righetti Abreu2, Cláudia Barbieri Ferreira Mendonça2 and Vania Gonçalves-Esteves2,3 Recebido em 2/10/2011. Aceito em 7/11/2011 RESUMO (Morfologia polínica de espécies do gênero Eremanthus Less. (Vernonieae, Asteraceae)). Com a fi nalidade de avaliar a importância da morfologia polínica para a taxonomia de Eremanthus (Vernonieae, Asteraceae) em nível genérico e infragenérico e fornecer dados adicionais para a sua reconstrução fi logenética, os grãos de pólen de 20 das 23 es- pécies do gênero foram examinados usando a microscopia de luz e eletrônica de varredura. Os grãos de pólen foram acetolisados, medidos, descritos e ilustrados sob microscópio de luz. Para a análise em microscópio eletrônico de varredura foram utilizados grãos de pólen com e/ou sem tratamento químico. As espécies apresentaram grãos de pólen isopolares, oblato-esferoidais, na maioria dos táxons, mais raramente prolato-esferoidais ou suboblatos, âmbito subtriangular, tricolporados e subechinolofados. A variação entre os caracteres quantitativos não se correlacionam com a subdivisão macromorfológica do gênero nem com os limites genéricos ou específi cos. Palavras-chave: Lychnophorinae, Compositae, taxonomia, palinologia ABSTRACT (Pollen morphology of the genus Eremanthus Less. (Vernonieae, Asteraceae)). In order to evaluate the signifi cance of the pollen morphology for generic and infrageneric taxonomy of the genus Eremanthus (Vernonieae, Asteraceae), and to provide additional data for its phylogenetic reconstruction, the pollen of 20 of the 23 species of the genus was examined using light and scanning electron microscopy. Acetolysed pollen grains were measured, described, and illus- trated using light microscopy, while non-acetolysed pollen grains were analyzed using scanning electron microscopy. Pollen grains of these species are isopolar, oblate-spheroidal in most of the species, more rarely prolate spheroidal or suboblate, subtriangular amb, tricolporate and subechinolophate. Th e variation among quantitative characters does not correlate with the macromorphological subdivision of the genus or with the generic or specifi c limits. Key words: Lychnophorinae, Compositae, taxonomy, palynology Introduction hairs, presence of sclerifi ed cells and lack of glands on the anther appendages (Robinson 1992, 1999, 2007; Keeley & Th e tribe Vernonieae Cass. (Asteraceae) contains c. 1500 Robinson 2009). species distributed mostly in the tropical parts of the world Eremanthus, a genus of c. 23 species, endemic to Brazil, with two major centers of diversifi cation: southeastern Brazil is a typical element of the cerrado and campo rupestre fl oras and the central region of Africa (Robinson 1999, 2006/2007; (Keeley & Robinson 2009). Robinson (1980) excluded her- Keeley & Robinson 2009). Th e 129 genera are placed into baceous elements from Eremanthus and MacLeish (1984, 21 subtribes mainly defi ned by their infl orescence pattern, 1987) recircumscribed the genus synonymising Vanillos- pollen morphology, chemical composition and chromo- mopsis (with deciduous inner pappus setae) under Ereman- some number (Keeley & Robinson 2009). Lychnophorinae thus (with persistent inner pappus setae) and transferring is one of the biggest subtribes of Vernonieae, and includes nine species from Eremanthus to Vernonia. Robinson (1999) ca. 100 species (11 genera) from the cerrado and campos restricted the concept of Vernonia to North American taxa, rupestres of Brazil. Th is subtribe is defi ned by the lack of and therefore had to reintegrate fi ve of the latter excluded enlarged nodes or sclerifi ed cells at the base of the styles, species into Eremanthus—E. crotonoides (DC.) MacLeish, usually the extensive presence of a pubescence of T-shaped E. leucodendron Mattf., E. mollis Sch. Bip., E. pabstii G. M. 1 Universidade de São Paulo, Instituto de Biociências, Departamento de Botânica, São Paulo, SP, Brazil 2 Universidade Federal de Rio de Janeiro, Museu Nacional, Departamento de Botânica, Rio de Janeiro, RJ, Brazil 3 Author for correspondence: [email protected] Pollen morphology of the genus Eremanthus Less. (Vernonieae, Asteraceae) Barroso and E. veadeiroensis H. Rob.—without specifi c des DC.) to his category VI: grains spherical, tricolporate, morphological investigations. Th e narrower concept of micropunctate, subechinolophate tending to echinate. Eremanthus, i.e., including Vanillosmopsis but not Paralych- Peçanha et al. (2008) described pollen grains of Paralych- nophora (MacLeish 1987, Hind 2000), is here followed (for a nophora bicolor (DC.) MacLeish (as E. bicolor (DC.) Baker) diff erent point of view, see Robinson 1997, 1999, 2006/2007). and E. erythropappus (DC.) MacLeish (as Vanillosmopsis Pollen characters of Asteraceae have been shown to erythropappa (DC.) Baker). Th is study revealed some in- be particularly variable and form useful patterns in the teresting variation of the exine pattern at the generic level: context of phylogenies. Such patterns could be used to Paralychnophora having a sublophate exine and Eremanthus support hypotheses of relationships, or provide diagnostic a subechinolophate one. Peçanha et al. (2008) concluded characters for groups at diff erent levels (Wortley et al. 2007, that their data do not support the inclusion of Vanillosmopsis Wortley et al. 2008, Blackmore et al. 2009). Blackmore et in Eremanthus (inclusion made by MacLeish in 1987). Such al. (2009) noticed that Vernonieae provides an example conclusion however cannot be considered valid, since they where a large number of pollen characters are congruent actually compared pollen grains of Paralychnophora with with the relationships presented in the Asteraceae supertree Eremanthus subg. Vanillosmopsis MacLeish, and not the (Funk et al. 2005). Blackmore et al. (2009) provide puta- latter with E. subg. Eremanthus Sch. Bip. as intended. tive synapomorphies for groups, most of them concern- Th e present study aimed to conduct a comprehensive ing African clades. Vernonieae pollen grains are usually examination of the pollen morphology in the genus Er- lophate or sublophate, tricolporate or triporate, with either emanthus, using LM and SEM, with almost complete spe- a continuous or discontinuous punctate or micropunctate cies coverage, in order to evaluate the signifi cance of this tectum, and are with or without spines (Keeley & Robinson structure for generic and infrageneric taxonomy and to 2009; for a detailed description see Blackmore et al. 2009). provide additional data for the phylogenetic reconstruction. Th e large pollen diversity of the tribe has been used exten- sively in taxonomic delimitations at generic and subtribal levels (Jones 1979, 1981, Bolick & Keeley 1994; Keeley & Materials and methods Robinson 2009). Kingham (1976) created six pollen Types Pollen grains of 20 species of Eremanthus were examined (I–VI) for the tribe, based on the study of 85 species (mostly (see Appendix). Th e samples were obtained from anthers of African). Keeley and Jones (1977, 1979) also identifi ed six fl ower buds from specimen kept at the herbaria ESA, HRB, basic Types (A–F) to which additional variations have been HUEFS, IBGE, MBM, SP, SPF, UB and UEC; acronyms ac- added (Robinson 1999). Currently, ten pollen Types are cording to Holmgren et al. (1990). recognised for the tribes (Dematteis & Pire 2008). Using Th e terminology used for descriptions follows Punt et mainly the classifi cation of Keeley and Jones, several authors al. (2007), taking into consideration size, form, number have analysed and described pollen grains of Vernonieae, of apertures and the ornamentation of the sexine. For the mostly of Vernonia s.l. (Keeley & Jones 1977, 1979; Jones light microscopy (LM) study, the pollen grains were pho- 1979, 1981; Robinson 1987a,b,c, 1988a,b,c, 1990; Skvarla et tomicrographed with a Sony Cyber-shot DSC-W7 digital al. 2005; Mendonça & Gonçalves-Esteves 2000; Mendonça camera coupled with a Zeiss Axiostar Plus microscope. et al. 2007a,b,c, 2009, 2010; Dematteis & Pire 2008; Angulo For scanning electron microscopy (SEM), acetolysed and & Dematteis 2010). non-acetolysed pollen samples were placed on aluminium Th e pollen grains of Lychnophorinae are tricolporate, stubs covered with carbon tape and sputter-coated with a echinate, sublophate with a perforated tectum continuous thin layer of gold palladium (c. 150 Å). Th e samples were between colpi (Type “A”) (Robinson 1992, 1999; Keeley analysed using a ZEISS DS M960 microscope at Laboratório & Robinson 2009). Th e endoaperture is usually lalongate de Ultraestrutura Celular Hertha Meyer, Inst. de Biofísica (Peçanha et al. 2008). Th e Type “A” pollen is considered (UFRJ) and a JEOL JSM-5800 microscope at Departamento a reversion from more strongly lophate ancestors in the de Invertebrados do Museu Nacional (UFRJ). Vernonieae, but it is nevertheless one of the most common Pollen grains were prepared for light microscopy by the forms in the tribe and is consistent in many groups such acetolysis method of Erdtman (1952), modifi ed by Melhem as the Piptocarphinae, Vernonia s.s. and Vernonanthura et al. (2003). Measurements in equatorial view (PD= polar (Robinson 1990, 1992, 1999). Robinson (1999) suggested diameter and ED=equatorial diameter) were randomly that due to common reversion of lophate pollen to Type “A”, taken on 25 pollen grains per sample. Measurements
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