Distribution of Satyrini (Lepidoptera, Nymphalidae) in Rio Grande Do Sul State, Southern Brazil
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Ecol Res DOI 10.1007/s11284-013-1035-z ORIGINAL ARTICLE Ana Luiza Gomes Paz • Helena Piccoli Romanowski Ana Beatriz Barros de Morais Distribution of Satyrini (Lepidoptera, Nymphalidae) in Rio Grande do Sul State, southern Brazil Received: 17 May 2012 / Accepted: 27 January 2013 Ó The Ecological Society of Japan 2013 Abstract Knowledge of species distribution is important Keywords Butterflies Æ Pampa biome Æ Satyrinae Æ for effective conservation measures. Considering that Similarity habitat features and abiotic factors can influence local community structure of butterflies, this study aimed to verify the existence of a pattern in the distribution of Sa- Introduction tyrini in southern Brazil. For this objective, we updated the regional Satyrini species list based on field studies and Species respond in different ways to the environment literature, resulting in a total of 54 species. For those and knowledge of their distribution is crucial for studies that presented clearly defined sampling sites, we understanding the dynamics of biological communities analyzed the faunal similarity by site and by phytoeco- (Walla et al. 2004). Thus, knowledge of beta diversity, logical region and verified the influences of environmental which is the variation in species composition among sites (altitude, average annual temperature and precipitation, in a geographical area (Whittaker 1972), is important for and phytoecological region) and spatial variables effective conservation measures (Blackburn and Gaston (PCNM) in species distribution. We analyzed 156 records 1996; Legendre et al. 2005; Ribeiro et al. 2008, 2012). of 40 species at 14 sites and eight phytoecological regions. The beta diversity may be influenced by random pro- Environmental variables—namely, differences in tem- cesses or environmental factors, or both, in different perature and phytoecological region—and spatial posi- proportions (Legendre et al. 2005). Generally, species tion proved to effect the distribution of Satyrini in similarity between sites has a negative relationship with southern Brazil. We thus urge that the future Conserva- geographic distance (Whittaker 1972) because neigh- tion Units should be well spaced and properly distributed boring sites tend to have more similar environmental through environmentally distinct units in the landscape, conditions (Legendre 1993), and also because of the representing different phytoecological regions. These influence of configuration and context of habitat on the conclusions shall provide subsidies to biodiversity con- dispersion of species (Nekola and White 1999; Soininen servation. et al. 2007). If beta diversity is influenced by environ- mental factors, careful studies are important for defining the spatial organization that protect the greater number of species possible (Bridgewater et al. 2004; Legendre A. L. G. Paz (&) Æ A. B. B. de Morais et al. 2005). Programa de Po´ s-Graduac¸ a˜ o em Biodiversidade Animal, The Rio Grande do Sul State (RS), in southern CCNE, Universidade Federal de Santa Maria, Faixa de Camobi, Brazil, comprises two biomes: Atlantic Forest, in the km 9, 97105-900 Santa Maria, RS, Brazil north, and Pampa, in the south of the state (IBGE E-mail: [email protected]ffarroupilha.edu.br Tel.: +55-32-574160 2004). The Atlantic Forest biome has a high environ- Fax: +55-32-571263 mental heterogeneity (Ribeiro et al. 2009) and in southern Brazil comprises a complex set of phytophisi- A. L. G. Paz ognomies with few remaining forest fragments (Leite Instituto Federal Farroupilha Campus Sa˜ o Vicente do Sul, Rua 20 de Setembro s/n, 97420-000 Sa˜ o Vicente do Sul, RS, Brazil 2002), and also includes important grassland formations (Overbeck 2007; Boldrini 2009). Despite being consid- H. P. Romanowski ered one of the world’s biodiversity ‘‘hotspots’’, there Programa de Po´ s-Graduac¸ a˜ o em Biologia Animal, Instituto de remains only 7.39 % of its original formation in the RS Biocieˆ ncias, Universidade Federal do Rio Grande do Sul, State (SOSMA/INPE 2008). The Pampa biome in Brazil Av. Bento Gonc¸ alves 9500, pre´ dio 43435, sala 229, 91501-970 Porto Alegre, RS, Brazil is represented only in the RS State and covers approxi- mately 63 % of its territory (IBGE 2004). Currently, inventories conducted in the RS State. We reviewed all approximately 50 % of the area is being used for agri- studies cited in Teston and Corseuil (2008), and later culture, livestock, and forestry activities, resulting in works that used entomological net as the sampling great biodiversity loss (Overbeck 2007; Roesch et al. methodology and identified individuals to species level 2009). On the other hand, information on native wildlife (Giovenardi et al. 2008; Paz et al. 2008; Sackis and and the current state of conservation of the biome are Morais 2008; Bonfantti et al. 2009; Iserhard et al. 2010; scarce, making it difficult to identify priority areas that Ritter et al. 2011; Rosa et al. 2011; Paz et al. 2012, deserve to be preserved (Morais et al. 2007; Santos et al. unpublished data). We updated the taxonomy following 2008; Roesch et al. 2009). Lamas (2004) and Wahlberg et al. (2009). The tribe Satyrini comprises 80 % of the species of The database sites (Fig. 1) and their respective biome the subfamily Satyrinae (Nymphalidae), which contains and phytoecological regional classifications, as well as approximately 2,200 species (Pen˜ a and Wahlberg 2008; abiotic variables and sampling efforts (net-hour), are Marı´ n et al. 2011). The Satyrini caterpillars feed mostly listed in Appendix 1. The classification of phytoecolog- on grasses (Poaceae) and the adults feed on fermented ical regions followed Cordeiro and Hasenack (2009). fruit, nectar, plant sap, and animal excrement (De Vries Abiotic data covering average annual temperature and 1987; Freitas and Brown 2004; Beccaloni et al. 2008; precipitation (Maluf 2000), altitude, latitude and longi- Pen˜ a and Wahlberg 2008). The adult butterflies of this tude (UTM) were obtained for each site. tribe usually have pale or brown colors and there are many studies including taxonomic (Pen˜ a and Lamas 2005; Pyrcz and Fratello 2005; Pyrcz et al. 2006; Freitas Statistical analysis et al. 2011), systematic (Pen˜ a et al. 2006; Marı´ n et al. 2011) and diversity aspects (Pyrcz and Rodrı´ guez 2007; For the statistical analyses, we used only those studies Pyrcz et al. 2011; Pyrcz and Garlacz 2012) of some that had sampling sites that were clearly defined and a species, tribes or sub-tribes in the Neotropical region. richness of at least five species (Romanowski et al. 2001, The richness of Satyrini in Brazil has not been com- unpublished data; Kru¨ ger and Silva 2003; Iserhard and pletely determined, and new species have recently been Romanowski 2004; Marchiori and Romanowski 2006a, b; described (Freitas et al. 2011). In RS State, the majority Dessuy and Morais 2007; Giovenardi et al. 2008; Paz of studies on the group were inventories of the total et al. 2008; Sackis and Morais 2008; Bonfantti et al. fauna of butterflies and descriptions of lists of associated 2009; Iserhard et al. 2010; Ritter et al. 2011). This gen- host plants (Silva et al. 1968; Biezanko et al. 1974). erated a difference between the number of species Teston and Corseuil (2008) compiled 48 species of this effectively used in the analyses and the general list of tribe from data sourced from literature reviews, field species. To test the differences of total sampling efforts sampling, and examinations of collections and museum between biomes, we run a test of equal or given pro- specimens. A more recent research review of southern portions with R package QuikPCNM v.7.7.1. We also South America cited two Satyrini, Hermeuptychia her- assessed whether there was association between sam- mes (Fabricius 1775) and Paryphthimoides phronius pling effort and species richness (Spearman correlation). (Godart 1824), among the most abundant and com- To test whether the distribution of species was different monly found butterfly species in fields and pastures of from chance, the null model SIM 2 was applied (fixed the RS State (Morais et al. 2007). The same authors lines and equiprobable columns), with 5,000 simulated highlight two other characteristic species of preserved null communities by sequential replacement using environments: Pampasatyrus periphas (Godart 1824) the program ECOSIM (Gotelli and Entsminger 2003, and P. ocelloides (Schaus 1902). 2011). Given that the habitat features and abiotic factors The Jaccard Index, using data of presence and absence can influence local structure of butterfly communities of species, was calculated for analyses of butterfly fauna (Brown and Freitas 2000a, b; Ribeiro et al. 2008, similarities, between sites and between phytoecological 2012; Dover and Settele 2009), this study aimed to regions. Thereafter, for the characterization of these verify the existence of a pattern in the Satyrini dis- similarities, we made an agglomerative classification tribution in the RS State. The influences of environ- analysis (Unweighted Pair-Group Method—UPGM) mental and spatial variables were tested. For this combining the fauna by sites and also by phytoecological purpose, we used a database assembled from literature region. UPGM clustering has high stability and maxi- and field sampling. mizes the cophenetic correlation coefficient (Rohlf and Sokal 1981). These procedures were made with NTSYSpc 2.10s software (Rohlf 2000). Materials and methods Canonical correspondence analysis (CCA) was used to assess whether the species composition of sites was related Satyrini Database