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Pollen Morphology in Brazilian Species of Codonanthe (Mart.) Hanst

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Pollen Morphology in Brazilian Species of Codonanthe (Mart.) Hanst Grana ISSN: 0017-3134 (Print) 1651-2049 (Online) Journal homepage: https://www.tandfonline.com/loi/sgra20 Pollen morphology in Brazilian species of Codonanthe (Mart.) Hanst. and Nematanthus Schrader (Gesneriaceae) Eduardo Custodio Gasparino, Maria Amelia Vitorino Da Cruz-Barros & Alain Chautems To cite this article: Eduardo Custodio Gasparino, Maria Amelia Vitorino Da Cruz- Barros & Alain Chautems (2013) Pollen morphology in Brazilian species of Codonanthe (Mart.) Hanst. and Nematanthus Schrader (Gesneriaceae), Grana, 52:4, 258-274, DOI: 10.1080/00173134.2013.826274 To link to this article: https://doi.org/10.1080/00173134.2013.826274 Published online: 27 Sep 2013. Submit your article to this journal Article views: 434 View related articles Citing articles: 8 View citing articles Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=sgra20 Grana, 2013 Vol. 52, No. 4, 258–274, http://dx.doi.org/10.1080/00173134.2013.826274 Pollen morphology in Brazilian species of Codonanthe (Mart.) Hanst. and Nematanthus Schrader (Gesneriaceae) EDUARDO CUSTODIO GASPARINO1, MARIA AMELIA VITORINO DA CRUZ-BARROS2 &ALAINCHAUTEMS3 1Faculdade de Ciências Agrárias e Veterinárias, UNESP - Universidade Estadual Paulista, Jaboticabal, Brazil, 2Núcleo de Pesquisa em Palinologia, Instituto de Botânica, São Paulo, Brazil, 3Conservatoire et Jardin Botanique de La Ville de Genève, Geneva, Switzerland Abstract Pollen morphology of 24 Brazilian species of Codonanthe and Nematanthus (Gesneriaceae) was investigated by means of light microscopy, scanning and transmission electron microscopy, in search of new characters that might contribute to taxonomic characterisation of the genera. The pollen grains are tricolpate or tricolporate, semitectate, reticulate, with simplicolumellate muri, sometimes with microreticulum and perforations, or duplicolumellate muri ring-shaped around lumina. The species studied were grouped into five pollen types on the basis of aperture characteristics and details of surface ornamentation. Sub-types were defined with reference to presence or absence of microreticulum and perforations around the lumina. Based on analysis of the morphology and measurements of pollen grains, we verified a close relationship among the genera, but can nevertheless discriminate their species. Keywords: Brazil, Episcieae, palynology, pollen grains The large and diverse tribe Episceae (Gesnerioideae, Codonanthe are not monophyletic as pointed out as Gesneriaceae), comprising about 22 genera and over early as Clark and Zimmer (2003). Nematanthus 700 species, is restricted to the Neotropics. The is distributed with 31 species in the Atlantic for- species are characterised by axillary flowers derived est along the Brazilian coast (Chautems, 1988; from a pair-flowered cyme inflorescence by reduc- Chautems et al., 2005). The genus consists of tion, a three-trace trilacunar node with split lateral sub-shrubs, epiphytes or lithophytes, with fleshy bundles, superior ovaries, mostly with a haploid fruits with loculicidally dehiscing capsules, and a chromosome number of n = 9[n = 8inCodonanthe haploid chromosome number of n = 8 (Chautems (Mart.) Hanst. and Nematanthus Schrader] and by & Matsuoka, 2003). The presence or absence of the presence of sepals connate at the base and an resupinate flowers was used by Moore (1973a) urceolate cup or funnel-shaped calyx (Wiehler, 1983; and Chautems (1988) to separate groups of Clark et al., 2006). Molecular data shows that the species within Nematanthus. Thus, floral morphol- tribe represents a monophyletic clade with a hetero- ogy has been used to assist in the classifica- geneous morphology within the Gesneriaceae (Smith tion of Nematanthus. Chautems (1988) recognised & Carroll, 1997; Smith, 2000a, 2000b; Zimmer 26 species and distinguished three groups: species et al., 2002; Clark et al., 2006, 2012). with non-resupinate flowers, with resupinate flowers, According to Clark et al. (2006), some generic lim- and with pendent resupinate flowers. Weber (2004) its of Episcieae are not well-defined, although the defined four types of flowers for the genus: hypocyr- Nematanthus, Codonanthe and Codonanthopsis Mansf. toid non-resupinate, hypocyrtoid resupinate, pen- clade is well-supported, the genera Nematanthus and dent resupinate and large campanulate white flowers. Correspondence: Eduardo C. Gasparino, Departamento de Biologia aplicada à Agropecuária, Faculdade de Ciências Agrárias e Veterinárias de Jaboticabal, Universidade Estadual Paulista, UNESP, via de acesso Prof. Paulo Castellane, s/n, CEP 14884-900, Jaboticabal, SP, Brazil. E-mail: [email protected] (Received 19 March 2013; accepted 1 July 2013) © 2013 Collegium Palynologicum Scandinavicum Codonanthe and Nematanthus pollen morphology 259 However, Clark et al. (2006) followed Chautems morphological features, pollen grains allow the dif- (1998) arrangement and treated the fourth group as ferentiation of genera and even species. white campanulate non-resupinate flowers. The aim of this study was to characterise the pollen Codonanthe has 18 species, growing epiphytically grains of native Brazilian species of Codonanthe and or lithophytically, with corollas tubular-campanulate Nematanthus (Gesneriaceae), in attempt to recognise to slightly ventricose, and as in Nematanthus,ahap- qualitative and quantitative characteristics that can loid chromosome number of n = 8 (Moore, 1973b; contribute to useful taxonomic characterisation for Wiehler & Chautems, 1990; Chautems & Matsuoka, these genera. 2003; Weber, 2004). The Amazonian and Central American species of the genus present extrafloral nectaries on the abaxial leaf side and, according to Material and methods Prance (1973), ants can assume an important role in seed dispersal. Fritsch (1894) used an anther feature, Pollen from dried herbarium specimens was supplied i.e. the broad connective causing a spatial separa- by the following herbaria: Universidade Estadual de tion of the pairs of thecae, to separate the species Campinas, Campinas, Brazil (UEC), Instituto de of Codonanthe from other genera of the subtribe Botânica, São Paulo, Brazil (SP) and Universidade Codonanthinae. de São Paulo, São Paulo, Brazil (SPF). Five out Currently, based on the fruit type, the genus is of 19 described species of Codonanthe,17out divided into two subgenera, Codonanthella (includ- of 31 species of Nematanthus and two hybrids ing species distributed in Central America and the from a total of 35 specimens were studied by Caribbean with fleshy capsules, as in Nematanthus) transmitted light microscopy (LM), scanning elec- and Codonanthe (with species that occur in Brazil tron microscopy (SEM) and transmission electron and the Amazon Basin). The species of the sub- microscopy (TEM). Pollen of 16 of these species genus Codonanthe are divided into two sections: had not been studied previously. The pollen sam- Spathuliformae occurring in eastern and north- ples were acetolysed according to the method of ern Brazil, Ecuador and Central America, and Erdtman (1960), with modifications by Melhem Codonanthe including the endemic species of the et al. (2003), and measured within seven days after Atlantic forest sympatric with species of Nematanthus their preparation (Salgado-Labouriau et al., 1965). (Chautems, 1988; Wiehler & Chautems, 1990). Permanent light microscopy slide preparations are There are only a few studies on pollen morphol- held in the pollen reference collection of the Instituto ogy of species of New World Gesneriaceae, and de Botânica, São Paulo, Brazil. papers dealing with Brazilian species are restricted to The polar axis (P) and equatorial diameter (E)of the studies of Campos (1962), Melhem and Mauro 25 pollen grains were measured per sample; other (1973), Melhem et al. (2003), Fourny et al. (2010) measurements (apocolpia, sexine, nexine thickness, and Gasparino et al. (2011). According to these lumina, muri, apertures and ornamentation) were taken from ten pollen grains per specimen. Statistical analyses were conducted to obtain the means, stan- Table I. Pollen grain metric variables used in PCA and cluster dard deviations, confidence intervals (95%), ampli- += = analysis ( present; – absent). tudes (minimum and maximum values) and variabil- Pollen grains ity coefficients (Vieira, 1981;Zar,1996). A principal Variables Tricolpate Tricolporate component analysis (PCA) was performed to ver- ify whether the pollen data permitted the grouping APOC (Apocolpium) ++of the species included here. This analysis was per- CLEN (Length of colpus) ++formed using the programs FITOPAC (Shepherd, CWID (Width of colpus) ++ EDEV (Equatorial diameter in ++1996) and PC-ORD (McCune & Mefford, 1999). equatorial view) To determine whether the pollen characteristics pro- EDPV (Equatorial diameter in ++vided additional discrimination among the analysed polar view) Codonanthe and Nemantanthus species, we compared PDEV (Polar diameter in ++the obtained pollen grains data by a cluster analy- equatorial view) ELEN (Length of endoaperture) −+sis (UPGMA and Euclidean distance) using version EWID (Width of endoaperture) −+7.0 of the STATISTICA software package (StatSoft. EXIN (Exine) ++Inc., Tulsa, OK, USA). For these analyses, we used SHAP (Shape) ++13 metric variables of the pollen grains (Table I). NEXI (Nexine) ++To calculate the width index of colpi, we divided the SEXI (Sexine) ++ TECT (Tectum) ++equatorial diameter by the width of the colpi. Indices below 5.0 were obtained for very large colpi, between 260 E.
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