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Palynological Analysis of the Genus Dryopteris Adans

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Palynological Analysis of the Genus Dryopteris Adans An Acad Bras Cienc (2020) 92(Suppl. 2): e20181052 DOI 10.1590/0001-3765202020181052 Anais da Academia Brasileira de Ciências | Annals of the Brazilian Academy of Sciences Printed ISSN 0001-3765 I Online ISSN 1678-2690 www.scielo.br/aabc | www.fb.com/aabcjournal BIOLOGICAL SCIENCES Palynological analysis of the genus Dryopteris Running title: PALYNOLOGICAL Adans. (Dryopteridaceae) in Argentina ANALYSIS OF Dryopteris IN ARGENTINA DANIEL A. GORRER, JUAN P. RAMOS GIACOSA & GABRIELA E. GIUDICE Academy Section: BIOLOGICAL Abstract: The spore morphology and wall ultrastructure of Dryopteris fi lix-mas, D. patula SCIENCES and D. wallichiana from Argentina were studied using light microscope, scanning and transmission electron microscope. The study was carried out with herbarium material from Argentine institutions. Equatorial diameters, polar diameters and laesura length e20181052 were measured. The spores are monoletes with a rugate ornamentation. The folds are short to long, infl ate, irregular in shape and size, and varying from subglobose to elongate. The perispore surface is rugulate. The exospore of all the species analyzed is two-layered 92 in section. Simple and branched channels are also present. The perispore is composed (Suppl. 2) of two layers, the inner one forms the ornamentation and the outer covers all the outer 92(Suppl. 2) and inner surfaces. Some abnormalities, such as globose, triangular or twisted spores were observed. The morphology and ultrastructure of the species are very similar. The differences observed are related to the length and thickness of the perispore folds. The characteristics of these spores would not provide relevant information to differentiate species or sections within the genus, but can provide information for phylogenetic studies as well as for alterations in the biological cycles. Key words: Argentina, Dryopteris, ornamentation, spore, ultrastructure. INTRODUCTION and D. wallichiana (Spreng.) Hyl., for continental Argentina. The Dryopteridaceae family is one of the most In tropical America, they usually grows in diverse among leptosporangiate ferns, with humid mountain forests, cloud forests and 26 genera and an estimated 2115 species of lowland rain forests. They grow in areas ranging cosmopolitan distribution (PPG I 2016). The from sea level to 4000 m, preferably within the genus Dryopteris Adans. is one of the most 1000 - 2500 m (Tryon & Tryon 1982, Narváez et complex within the family and one of the al. 2008). richest in species, with about 225-300 species Classifi cations within the genus have been throughout the world except Antarctica (Geiger established on the basis of morphology (Sessa et & Ranker 2005, Sessa et al. 2012b). About 160 al. 2012b): Ito (1935, 1936) has treated the species of them are found in Southeast Asia, which of Japan and Taiwan whereas Ching (1938) has seems to be their center of diversity (Sessa et done so with those of China, the Himalayas, al. 2012b), about 15 species are distributed in India and Sri Lanka. Fraser-Jenkins (1986) has Central and South America (Prado et al. 2014) made the most accepted classifi cation which and 3 species are cited by Ponce & Arana (2016): includes 225 species, subdivided in 4 subgenera D. fi lix-mas (L.) Schott, D. patula (Sw.) Underw. and 16 sections, several incertae sedis and about 90 hybrids. Within the last classifi cation, An Acad Bras Cienc (2020) 92(Suppl. 2) DANIEL A.D. GORRER, JUAN P. RAMOS GIACOSA & GABRIELA E. GIUDICE PALYNOLOGICAL ANALYSIS OF Dryopteris IN ARGENTINA the 3 species growing in Argentina belong to the Several authors have reported that subgenus Dryopteris although they are located smallest spores are diploid while the largest in different sections: D. filix-mas correspond to ones are tetraploid and hexaploid (Mehra & Sect. Dryopteris, D. patula to Sect. Cinnamomeae Loyal 1965, Kanamori 1969, Mitui 1972, Tryon & and D. wallichiana to Sect Fibrillosae. Lugardon 1991, Quintanilla & Escudero 2006). The geographic distribution of D. filix-mas Regarding hybrids, their chromosomes tend to corresponds to North America, Europe, Asia, be irregularly distributed so the spores differ Madagascar and it was introduced in South genetically among them and in size (Witthier & America. In Argentina, it was cultivated and Wagner 1971). Thus, differences in spore sizes naturalized in the Patagonian region (Sessa et are related to ploidy level and hybridity (Manton al. 2012a, Ponce & Arana 2016). D. patula grows 1950, Wagner 1966, Kanamori 1971, Nakato & in Central and South America. In Argentina it is Mitui 1979, Moran 1982, Pryer & Britton 1983, found in the Northwestern region (Prado et al. Barrington et al. 1986). 2014). D. wallichiana is a species that grows in the The morphology of the spores of several Himalayas, Africa, India, China, Japan, Malaysia Dryopteris species has been examined in and America, along the Andes, from Mexico to worldwide in various places such as Africa, Asia, the northwest of Argentina (Tryon & Lugardon Europe and, North and Central America (Crane 1991, Narváez et al. 2008, Ponce & Martínez 2012, 1953, 1955, 1956, 1960, Nayar & Devi 1964, Britton Prado et al. 2014, Ponce & Arana 2016). 1972a, b, Mitui 1972, Belling & Heusser 1974, The Dryopteris genus presents one of the Britton & Jermy 1974, Tryon & Tryon 1982, Tryon most difficult fern complexes in America due to & Lugardon 1991, Lee & Park 2014). its tendency to hybridize (Walker 1959, Whittier Crane was the first who studied spores of the & Wagner 1971, Hoshizaki & Wilson 1999) thus Dryopteris genus from North America (1953, 1955, creating problems in the definition of species in 1956, 1960). He made a key for North American the group (Barrington et al. 1989). species based on the characters of the spores. In addition to its tendency to hybridize, the Some of these characters, such as the presence family often has apogamic species (Nayar & or absence of spines, spore size and the size Kaur 1971), including D. wallichiana (Loyal 1959, and number of folds are very valuable, since Fraser-Jenkins 1986, 1989, Pérez-García et al. they help to easily differentiate some species 2001, Geiger & Ranker 2005, Narváez et al. 2008, from others. Britton & Jermy (1974), on the other Sessa et al. 2012b). hand, showed that the spores of D. filix-mas As well as hybridity, the ploidy level has from the United States, Canada and Mexico have also been studied by different authors in the a background pattern of the perispore which is following species: D. wallichiana has been anastomosed or reticuled. registered as diploid, apogamic triploid and Only a few studies using SEM have been hexaploid (Tryon & Tryon 1982, Sessa et al. carried out on the spores of the genus in South 2012a, 2015); D. filix-mas is considered a sexual America. The spores of D. filix-mas have only allotetraploid (Manton 1950, Manton & Walker been illustrated from Argentina by de la Sota 1954, Fraser-Jenkins 1976, Lovis 1977, Xiang et al. et al. (1998) and they have been characterized 2006) and D. patula is a diploid species (Tryon as greenish brown and with crestate perispore. and Tryon 1982, Sessa et al. 2012a). The spores of D. patula have been described by Prado et al. (2014) with material from Brazil as An Acad Bras Cienc (2020) 92(Suppl. 2) e20181052 2 | 12 DANIEL A.D. GORRER, JUAN P. RAMOS GIACOSA & GABRIELA E. GIUDICE PALYNOLOGICAL ANALYSIS OF Dryopteris IN ARGENTINA ellipsoidal, rugose and having long broad folds. For the analysis with LM, the material was Tryon & Lugardon (1991) have observed the acetolized according to the method of Erdtman spores of this species with material from Mexico (1960). For the study with SEM, the spores without as inflated tubercles with fine, superficial ridges. treatment were placed into stubs with adhesive The spores of D. wallichiana from Mexico and double-faced tape and coated with gold. For the Peru have been described by Tryon & Lugardon TEM analysis, the material was treated following (1991) as long inflated folds, forming the rugate the technique of Rowley & Nilsson (1972): it was surface. Narváez et al. (2008) have mentioned rehydrated with 0.1 M buffer and with 1% Alcian the spores of this species from Argentina as Blue (AB) for 2 hours; then the material was fixed dark brown and rugulate, having wide rounded with 1% glutaraldehyde + 1% AB in phosphate usually anastomosed folds and forming short buffer for 12 hours; then it was washed for 15 rounded reticles. minutes in phosphate buffer and postfixed with However, TEM studies of the genus have not 1% Osmium tetraoxide in water + AB for 2 hours. been carried out in America yet. The spores of The spores were dehydrated in an acetone series D. filix-mas from France have been studied with (30-100%) and subsequently embedded in a TEM by Tryon & Lugardon (1991). These authors mixture of Spurr resin. The semi-thin sections have described the spores as having a plain 3 μm thick were stained with Toluidine blue exospore and a perispore of one cavate layer and observed with LM. The ultrathin sections usually showing short scales below. were stained with Uranyl acetate for 15 minutes The spores of the species growing in followed by Lead citrate for 3 minutes. Argentina have scarcely been studied with The observations were made with a Zeiss SEM and have not been studied with TEM yet. EM 109T with Gatan ES1000W digital camera Besides, the sporoderm demands for a thorough from the Instituto de Biología Celular, Facultad study in order to define its ultrastructure and de Medicina, Universidad de Buenos Aires, a complexity. The aim of this work was to analyze JEOL JSMT-100 SEM from the Museo de Ciencias the morphology and ultrastructure of the spores Naturales de La Plata and a Nikon E200 from of the genus Dryopteris in continental Argentina Cátedra de Morfología Vegetal, Facultad de with LM, SEM and TEM. Ciencias Naturales and Museo, Universidad Nacional de La Plata.
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