Different Slopes of a Mountain Can Determine the Structure of Ferns and Lycophytes Communities in a Tropical Forest of Brazil
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Anais da Academia Brasileira de Ciências (2014) 86(1): 199-210 (Annals of the Brazilian Academy of Sciences) Printed version ISSN 0001-3765 / Online version ISSN 1678-2690 http://dx.doi.org/10.1590/0001-3765201495912 www.scielo.br/aabc Different slopes of a mountain can determine the structure of ferns and lycophytes communities in a tropical forest of Brazil FELIPE C. NETTESHEIM1, ELAINE R. DAMASCENO2 and LANA S. SYLVESTRE3 1Universidade Federal do Rio de Janeiro, Programa de Pós-graduação em Ecologia, CCS, Ilha do Fundão, 21941-590 Rio de Janeiro, RJ, Brasil 2Universidade Federal do Rio de Janeiro, Programa de Pós-graduação em Botânica, Museu Nacional, Quinta da Boa Vista, s/n, São Cristóvao 20940-040 Rio de Janeiro, RJ, Brasil 3Universidade Federal do Rio de Janeiro. Departamento de Botânica, CCS, Instituto de Biologia, Ilha do Fundão, 21941-902 Rio de Janeiro, RJ, Brasil Manuscript received on Febryary 16, 2012; accepted for publication on March 28, 2013 ABSTRACT A community of Ferns and Lycophytes was investigated by comparing the occurrence of species on different slopes of a paleoisland in Southeastern Brazil. Our goal was to evaluate the hypothesis that slopes with different geographic orientations determine a differentiation of Atlantic Forest ferns and lycophytes community. We recorded these plants at slopes turned towards the continent and at slopes turned towards the open sea. Analysis consisted of a preliminary assessment on fern beta diversity, a Non Metric Multidimensional Scaling (NMDS) and a Student t-test to confirm if sites sampling units ordination was different at each axis. We further used the Pearson coefficient to relate fern species to the differentiation pattern and again Student’s t-test to determine if richness, plant cover and abundance varied between the two sites. There was a relatively low number of shared species between the two sites and ferns and lycophytes community variation was confirmed. Some species were detected as indicators of the community variation but we were unable to detect richness, plant cover or abundance differences. Despite the evidence of this variation between the slopes, further works are needed to evaluate which processes are contributing to determine this pattern. Key words: Atlantic rain forest, topography, pteridophytes, richness, environmental factors. INTRODUCTION such as topography (Vormisto et al. 2004), soil Plant species distribution within tropical forest characteristics (Jones et al. 2006, Tuomisto et al. landscapes has been shown to be generally deter- 2003 b, c), length of the dry period and moisture mined by factors which vary from biogeographic availability (Engelbrecht et al. 2005). Although there to microenvironmental scales (Svenning 1999, Ter is considerable debate as to what degree floristic Steege et al. 2003, Condit et al. 2002). Ranging from distributional patterns depend on environmental local to regional scales, several studies have shown factors relative to other processes (Hubbell 2001, that plant species composition and abundances Dalling et al. 2002, Wyatt and Silman 2004), answer to some common environmental variables findings appear to show that the major role is played by environmental factors (Karst et al. 2005, Jones et Correspondence to: Lana da Silva Sylvestre E-mail: [email protected] al. 2006, 2008, Zuquim et al. 2009). An Acad Bras Cienc (2014) 86 (1) 200 FELIPE C. NETTESHEIM, ELAINE R. DAMASCENO and LANA S. SYLVESTRE The environmental model has provided consis- different scales (Karst et al. 2005, Jones et tently better predictive power for Amazonian ferns al. 2006, 2008, Zuquim et al. 2009). It is thus structure than for trees and palms (Ruokolainen et al. reasonable to think that ferns exposed to the 1997, Pitman et al. 2001, Potts et al. 2002, Tuomisto geomorphologic heterogeneity along the Atlantic et al. 2003a, b, c, Phillips et al. 2003, Vormisto et al. Forest would be influenced by environmental 2004). It is reasonable to associate such a result to variation. This might be especially true where the fact that ferns are plants which present variable topography is highly variable and close to taxon specific establishment success. These plants the ocean, being subjected to very humid cold have adaptations to low levels of light incidence, low fronts and sea wind (Barbosa 2007). Such a evaporative potentials, a unique water dependent geographical situation is likely to cause changes reproductive biology and high sensibility to abiotic in factors that are relevant for fern occurrence, features such as substrate. Therefore ferns and such as moisture and light incidence. Literature lycophytes communities structure are expected to has registered that ferns leaf length appears to be very subjective to environmental changes related decrease with increasing aridity while locations to solar irradiance and atmosphere moisture levels with greater moisture normally have greater (Wylie 1948, Barrington 1993, Chazdon et al. 1996, diversity and abundance (Page 2002, Kessler Sonoike 1996, Page 2002). Only recently, studies et al. 2007). Throughout the year, southeastern started addressing such issues in the Atlantic forest Brazilian forests located in northerly slopes have with plant groups. Therefore, knowledge about the greater exposure to sunlight than those located in relative contribution of environmental factors to southerly slopes (Lima 1986, Oliveira et al. 1992, explain floristic composition variation along the Marques et al. 2005). Studies in this region show Atlantic forest is still starting to stack up (Oliveira- that areas turned to the ocean are indeed more Filho and Fontes 2000, Oliveira-Filho et al. 2005, humid due to a greater mean annual rainfall than Paciencia and Prado 2005, Duarte et al. 2010, areas turned inland (Oliveira-Filho and Fontes Machado and Oliveira-Filho 2010, Nettesheim et 2000, Oliveira et al. 2005, Paciencia and Prado al. 2010, Marques et al. 2011, Garbin et al. 2012). 2005, Nettesheim et al. 2010). These efforts The extensive habitat heterogeneity along a wide argue that such patterns are in general agreement range distribuition (Tonhasca Jr 2005) makes this with the Atlantic forest’s evergreen and seasonal forest a good model to test recent findings regarding distribution, apparently limited by annual rain tropical plants distributional patterns (Svenning et and principally by the duration of the dry period. al. 2010, Jones et al. 2011, Kluge and Kessler 2011, Once many efforts have consistently predicted Ricklefs and Renner 2012). ferns and lycophytes floristic variation using A particular group containing good potential environmental models and assuming different slopes for the study of the processes regulating plants near the ocean are likely to exhibit environmental distribution in the Atlantic forest appears to be variation, the aims of this study were: (1) determine the ferns. Not only do they have high richness if, as expected, there is floristic variation of the and endemism levels (Tryon and Tryon 1982), Atlantic forest ferns community between two but they are known to have had a large part of different slopes of a paleoisland (northern-facing its floristic variation consistently predicted by vs. southwestern-facing); (2) considering that environmental variation. Among the array of the southwestern slopes are more humid than the environmental factors, topographic and edaphic northwestern slopes, determine if it has greater fern factors appear to be noticeably important at richness, abundance and plant cover area. An Acad Bras Cienc (2014) 86 (1) SLOPE ORIENTATION AFFECTS PTERIDOPHYTE COMMUNITY 201 MATERIALS AND METHODS A total of thirty 5x5m individual sample units (plots) were arbitrarily established at each of the STUDY SITE two studied sites. Location of the plots was about Marambaia paleoisland is a portion of granite- 4km apart. Plots were scattered to maximize the gneiss crystalline basement land linked to the sampling design reach throughout an altitudinal continent by Marambaia Restinga. It is located range between 150 and 300m within each site. We south of the State of Rio de Janeiro (23°04'37.09"S; established a total of 60 plots (each with 25m2) 43°59'2.15"W), in Sepetiba Bay, on Mangaratiba at Marambaia paleoisland, sampling a total area municipality. The paleoisland together with Serra of 1,500m2. Plot area was measured close to the do Mar Range is a consequence of tectonic activity ground, since inventory considered terrestrial herbs, which dates back to the Jurassic Period (130 million tree ferns, plants growing on fallen trees (including years ago), associated with rupture of ancient in cases where epiphytic species survived the Gondwana supercontinent (Almeida and Carneiro fall), climbers and low trunk epiphytes. We used 1998). Marambaia paleoisland was formerly a tape measure to record width and length of each isolated from the Serra do Mar Range by rising individual. These values were then multiplied by ocean level in the quaternary (Menezes et al. 2005, each other to determine plant cover estimated value. Souza et al. 2005). It is covered with 2,125.43ha Plantlets were not sampled since the identification of Atlantic rain forest, characterized by Veloso et of these individuals are inaccurate. Climbing al. (1991) as a Sub-Montane Dense Ombrophylous individuals were considered only when at least one Forest. The terrain altitude varies from 0 to 641 green frond was present at maximum 2 m above meters. The site is encompassed as macroclimate the ground (Tuomisto et al.