Pollen Morphology of Eight Species of Stemodia (Plantaginaceae) from South America María De Las Mercedes Sosa a , Andrea F
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This article was downloaded by: [Mercedes Sosa] On: 28 October 2011, At: 05:34 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Palynology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tpal20 Pollen morphology of eight species of Stemodia (Plantaginaceae) from South America María de las Mercedes Sosa a , Andrea F. Panseri a & Cristina R. Salgado a a Instituto de Botánica del Nordeste (UNNE-CONICET), Sargento Cabral 2131, Casilla de Correo 209, 3400, Corrientes, Argentina Available online: 25 Oct 2011 To cite this article: María de las Mercedes Sosa, Andrea F. Panseri & Cristina R. Salgado (2011): Pollen morphology of eight species of Stemodia (Plantaginaceae) from South America, Palynology, DOI:10.1080/01916122.2011.572372 To link to this article: http://dx.doi.org/10.1080/01916122.2011.572372 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Palynology iFirst article 2011, 1–9 Pollen morphology of eight species of Stemodia (Plantaginaceae) from South America Marı´a de las Mercedes Sosa*, Andrea F. Panseri and Cristina R. Salgado Instituto de Bota´nica del Nordeste (UNNE-CONICET), Sargento Cabral 2131, Casilla de Correo 209, 3400 Corrientes, Argentina The pollen morphology of eight species of Stemodia L. (Plantaginaceae) was analysed with the aim of refining the taxonomy of this genus. The pollen grains were examined with both light and scanning electron microscopes. The pollen grains are spheroidal, sometimes prolate-spheroidal, small, with a subcircular or subtriangular amb, 3- or 4-colporate and with a long colpus that reaches the poles, defining a small, circular to slightly lalongate apocolpium with irregular margins. The exine is tectate, foveolate or microperforate, 1 mm thick and uniform. The species of Stemodia analysed here exhibit morphological homogeneity, which suggests that the genus is stenopalynous. Keywords: pollen morphology; Stemodia; South America 1. Introduction cytotypes (Sosa and Seijo 2002; Sosa et al. 2009); Stemodia L. (Plantaginaceae) comprises 49 species Stemodia lobelioides Lehm., which has a tetraploid which live in neotropical areas (Dawson 1968; Turner cytotype (Sosa et al. 2009); and Stemodia viscosa, which and Cowan 1993a; Barringer and Burger 2000). A total has tetraploid and hexaploid cytotypes (Raghavan and of 29 species have been recognised for the Americas, 16 Srinivasan 1940; Subramanian and Pondmudi 1987). of which are exclusive to South America (Turner and However, a correlation between size of pollen grains Cowan 1993b). There are 9 species in southernmost and ploidy levels has not yet been determined. South America, but only Stemodia verticillata (Mill.) Palynological studies are a valuable tool for Hassl. has a broader distribution. The remaining 8 taxonomic separation. In this sense, studies on pollen species are typically endemic to Argentina, Paraguay, morphology have allowed the formation of natural Uruguay, southern Bolivia and southern Brazil. The taxonomic groups in the Family Scrophulariaceae Stemodia species are annual or perennial marshy herbs, sensu lato (Hong 1984). Although several palynological common in marshlands and lowlands; only Stemodia studies have been carried out in species from Europe, verticillata grows in drier areas, mostly on clay-rich acid Asia and Africa (Varghese 1968; Inceoglu 1982; Argue soils. 1984, 1986, 1993; Karim and El-Oqlah 1989; Fernan- The genus is a natural, well-delimited group, dez et al. 1997), these types of studies in neotropical although the identification of some taxa may be taxa are relatively scarce (Argue 1980, 1981, 1985; Downloaded by [Mercedes Sosa] at 05:34 28 October 2011 complex. For that reason, the genus has been subject Correa et al. 1995; Santos and Melhem 1999, 2000). to taxonomical, anatomical and cytological studies Morphological analysis of South American species (Raghavan and Srinivasan 1940; Subramanian and of Stemodia (Sosa 2009) has shown that different taxa Pondmudi 1987; Turner and Cowan 1993b; Sosa and are similar in vegetative characters, which hinders their Seijo 2002; Sosa 2005, 2008; Sosa et al. 2009). identification. Additional data on anatomy or palynol- However, studies on pollen morphology have been ogy may provide further information on taxonomy. In made only on Stemodia viscosa Roxb. (Varghese 1968). this paper, the pollen morphology of eight species of The variation of some pollen characters such as Stemodia from southern South America is described shape and size can be related to ploidy level (Stebbins for the first time. The results are discussed in relation 1971; Soltis and Soltis 1999). In that sense, some authors to the taxonomy of the genus. have observed a positive correlation between the size of pollen grains and ploidy levels (Stebbins 1971; Oliveira et al. 2004; Almada et al. 2006). Polyploidy has been 2. Materials and methods reported in three species: Stemodia hyptoides Cham. & Twenty specimens belonging to eight species of Schltdl., which has diploid, tetraploid and hexaploid Stemodia were studied (Plates 1, 2). Pollen samples *Corresponding author. Email: [email protected] ISSN 0191-6122 print/ISSN 1558-9188 online Ó 2011 AASP – The Palynological Society http://dx.doi.org/10.1080/01916122.2011.572372 http://www.tandfonline.com 2 M. de las Mercedes Sosa et al. were obtained by removing one or two flowers from 7 km from the city, 23-XI-2005, Sosa et al. 213 (CTES, specimens of each species deposited in the Herbarium SP, G). PAL-CTES 2534. of the Instituto de Bota´nica del Nordeste (CTES). In some cases, flower buds were obtained from the plants growing in greenhouse conditions; these were pre- Stemodia hyptoides Cham. & Schltdl. viously fixed in formaldehyde-acetic acid-alcohol Argentina. Corrientes: Dept. Mburucuya´, Paso (FAA) for 24 hours then preserved in 70% alcohol. Aguirre, 6-V-2001, Sosa 43 (CTES). PAL-CTES Pollen grains were acetolysed using the technique of 2544. Dept. Mercedes, El Socorro, 24-II-2006, Sosa Erdtman (1960). For light microscopy (LM) the pollen et al. 245 (CTES) PAL-CTES 2536. Dept. Saladas, samples were mounted in glycerin jelly on glass slides, Paso Naranjo, 17-I-2004, Sosa 128 (CTES) PAL- sealed with paraffin wax and examined with a Wild M CTES 2545. Misiones: Dept. Capital, Posadas, Zaima´n 20 microscope. Permanent slides were deposited at the stream, 16-II-2005, Sosa and Rodrı´guez 138 (CTES) Palynological Laboratory of the Universidad Nacional PAL-CTES 2530. Dept. Montecarlo, Puerto Piray, del Nordeste (PAL-CTES). 12-IV-2006, Sosa and Rodrı´guez 249 (CTES) PAL- For observation with scanning electron microscopy CTES 2535. Dept. San Martı´n, Puerto Rico, 12-IV- (SEM), acetolysed pollen grains were washed in 968 2006, Sosa and Rodrı´guez 251 (CTES) PAL-CTES alcohol which was followed by washing in absolute 2546. alcohol. The pollen grains were then air-dried on aluminium plates and sputtered with gold palladium and studied with a JEOL 5800 LV SEM. The description Stemodia lanceolata Benth. of pollen grains follows the terminology of Punt et al. Argentina. Chaco: Dept. 18 de Mayo, Colonia Benı´tez (1994, 2007). The length of the polar axis (P) and the 19-IV-2001, Sosa and Loizaga de Castro 41 (CTES, equatorial diameter axis (E) were measured from 30 pollen SPF) PAL-CTES 2543. Salta: Dept. Ora´n, Tabacal, grains per specimen using LM. Perforations and thickness 15-XII-2005, Sosa et al. 228 (CTES, ESA) PAL-CTES of the pollen wall were measured with the SEM. 2531. The Shapiro–Wilk test was used to evaluate the normal distribution of the measurements. Prior to statistical analyses, data with non-normal distribution Stemodia lobelioides Lehm. (W ¼ 0.891, p 5 0.0001) were log-transformed. The Argentina. Corrientes: Dept. Monte Caseros, Uruguay measurements of pollen size for diploid species were river, 6-V-2005, Sosa and Schinini 210 (CTES) analysed using a multivariate analysis of variance PAL-CTES 2540. Uruguay. Dept. Artigas: Bella (MANOVA); when differences were significant, se- Unio´n, 25-XI-2001, Sosa et al. 70 (CTES, HUEFS). paration of means was performed using the Bonferro- PAL-CTES 2532. ni’s a posteriori tests (p 5 0.05). Since we have previously reported three cytotypes in Stemodia hyptoides (2n ¼ 2x ¼ 22, 2n ¼ 4x ¼ 44 and 2n ¼ 6x ¼ 66) Stemodia palustris A. St.-Hil. (Sosa and Seijo 2002; Sosa et al. 2009), we used Uruguay. Dept. Artigas, near Yacutuya´Minı´stream, Downloaded by [Mercedes Sosa] at 05:34 28 October 2011 MANOVA to analyse the pollen size between these 25-XI-2001, Sosa et al. 71 (CTES) PAL-CTES 2537; three cytotypes. We also used correspondence analysis around Guabiyu´stream, 5-V-2005, Sosa and Schinini to explore the relationship between grain pollen shape 209 (CTES) PAL-CTES 2541. (categorical data) and species. All analyses were performed using the Infostat program (version 2010). Stemodia stricta Cham. & Schltdl. Argentina. Jujuy: Dept. Ledesma, around Zapla river, 2.1. Specimens examined 16-XII-2005, Sosa et al. 230 (CTES, ESA) PAL- Stemodia ericifolia (Kuntze) K. Schum. CTES 2539. Salta: Dept. San Martı´n, near Carapaı´ Argentina. Formosa: Dept.