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Biogeography and Areas of Endemism of Prepops Reuter (Heteroptera: Miridae)

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Biogeography and Areas of Endemism of Prepops Reuter (Heteroptera: Miridae) Biogeography and areas of endemism of Prepops Reuter (Heteroptera: Miridae) Lívia Aguiar Coelho1, 2*, Carlos Molineri3, Daniel Andrés Dos Santos4 & Paulo Sérgio Fiuza Ferreira5 1,5. Universidade Federal de Viçosa, Departamento de Entomologia, Museu Regional de Entomologia, 36570-900, Viçosa, MG, Brazil; [email protected], [email protected] 2. Present Address: Universidade Federal da Grande Dourados, Faculdade de Ciências Biológicas e Ambientais, Programa de Pós-Graduação em Entomologia e Conservação da Biodiversidade, 79804-970, Dourados, MS, Brazil. 3,4. Instituto de Biodiversidad Neotropical, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Tucumán, Horco Molle (4107), Tucumán, Argentina; [email protected], [email protected] * Correspondence Received 26-II-2015. Corrected 25-VII-2015. Accepted 28-VIII-2015. Abstract: The Miridae (Hemiptera: Heteroptera) are one of the most species rich families of insects, with about 11 100 described species. Mirinae is the largest subfamily and its endemism becomes evident at the tribal level and below. Mirines of the tribe Restheniini are found throughout America, with most of the diversity confined to the Neotropics. Prepops Reuter is the largest genus in Restheniini with 198 described species and its geographi- cal range is similar to that of the tribe. The study of the distribution and areas of endemism in Miridae are scarce and non-existent for Prepops. We analyzed all the geographic records of Prepops (707 records for 181 species) in order to identify diversity patterns and areas of endemism, using network analysis. Locality data were assigned using museum specimens and geographic records in the literature. Additionally we used spatial data to propose habitat preferences and geoclimatic variables important for each group of co-occurring species. The results indicate high species richness in subtropical regions, while the tropical belt and high latitudes appear impover- ished. The Neotropical region contains 86% of the species, and the Nearctic about 11 % (but with larger species ranges); the Andean region is sparingly represented. Twenty-three areas of endemism (and two diads), formed by 2 or more endemic species, are distributed in the Nearctic (3), Mexican Transition Zone and Caribbean sub- region (5), South American Transition Zone and Amazonian, Chacoan and Paranaense subregions (15). Overlap between some of the areas indicates regions with high species richness and complex history. The lack of records from the Andean region (except for P. nigrus in Southern Patagonia) and dominance of strictly Neotropical spe- cies (86 %) supports the hypothesis of a Neotropical origin for the tribe Restheniini. The general arrangement of the units of co-occurrence of Prepops species shows a close correlation with known biogeographic regions and subregions. Broad physiographic characteristics most commonly associated with Prepops geographical records are, in order of importance: broad leaf forests (wet and dry), grasslands and xeric habitats. Rev. Biol. Trop. 64 (1): 17-31. Epub 2016 March 01. Key words: Mirinae, Restheniini, plant bugs, spatial analysis, network analysis, diversity gradient, endemic areas. The Miridae (Hemiptera: Heteroptera) are tropical and Mediterranean ecosystems (Cassis one of the most species rich families of insects, & Schuh, 2012). with about 11 100 described species. They are Among the eight recognized subfamilies, found on all continents, except the Antarc- Mirinae is the largest, comprising 419 gen- tic (Wheeler, 2001; Cassis & Schuh, 2012; era in six tribes (Henry, 2009; Schuh, 2013). Schuh, 2013), and are diverse especially in Endemism becomes evident at the tribal level Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 64 (1): 17-31, March 2016 17 and below. Mirines of the tribe Restheniini are biogeographic analysis. Several approaches found throughout America, with most of the have been proposed to achieve these aims but diversity confined to the Neotropics (Wheeler, all have particular methodological problems 2001; Cassis & Schuh, 2012). This aposematic (Crisci et al., 2003; Dos Santos, Fernández, colored tribe includes the largest known mir- Cuezzo, & Domíngues, 2008; Casagranda et ids (15 mm or more) (Schuh & Slater, 1995; al., 2009; Dos Santos, Cuezzo, Reynaga, & Henry, 2009) and much of the work has been Domíngues, 2011; Torres-Miranda, Luna-Veja, descriptive. Host plants are virtually unknown & Oyama, 2013). Dos Santos et al. (2008, (Ferreira, Silva, & Coelho, 2001; Hernández & 2011) proposed to use network analysis based Henry, 2010; Wheeler & Bundy, 2012; Schuh, on sympatry inference as a first step in the 2013). Prepops Reuter is the largest genus in search for areas of endemism. Biogeographic Restheniini with 198 described species (Schuh, analysis typically focuses on sympatry patterns 2013; Coelho, Ferreira and Costa, 2012); the among species. Sympatric species lives in the geographical range is similar to that of the same local community close enough to inter- tribe and research is also required about their act, and generally have broadly overlapping host-plant relationships and habits (Ferreira et geographic distributions (Crisci et al., 2003). al., 2001; Wheeler, 2001; Hernández & Henry, Dos Santos et al. (2008, 2011) proposed a new 2010; Wheeler & Bundy, 2012; Schuh, 2013). operational definition of sympatry: two or more The study of the distribution and areas species are sympatric when there are interpen- of endemism in Miridae is scarce (Wheeler, etration and relative proximity among their 2001); and available information is mostly records. Coincidence of two or more species in focused on the study of particular regions or the same locality is the maximum expression of specific taxa (e.g. Schuh, 1974, 1984, 1991, these properties. 2006; Ribes, 1984; Schuh & Stonedahl, 1986; Network analysis has proven to be a pow- Wheeler & Henry, 1992; Lu & Zheng, 1998; erful tool for studying different aspects of Williams, 2002; Forero & Schwartz, 2009; biological systems (Proulx, Promislow, & Phil- Hernández & Henry, 2010) without using lips, 2005). The major difference between quantitative methods for the biogeographi- conventional distributional data and network cal analysis. Until now, studies that mention the subfamily Mirinae are scarce (Williams, data analysis is that the first focuses on areas 2002; Cassis & Schuh, 2012), and do not treat and their attributes (species), whereas net- the Restheniini. work data analysis focuses on species and Following Hovenkamp (1997), historical their sympatric relationships (Dos Santos et biogeography has two main goals: to recon- al., 2008, 2011). So, species groups satisfying struct the distributional history of individual the requirement of within-group sympatry and groups (taxon biogeography), and to recon- between-group allopatry will conform to natu- struct the history of areas of endemism or the ral units of co-occurrence (UCs). A sympatry history of Earth (the search for general area network may contain dense groups of species relationships). Areas of endemism are tradi- (UCs) connected through intermediary species. tional units for historical biogeography (Crisci, These dense groups will be evident after the Katinas, & Posadas, 2003) and many defini- intermediary species are removed. The spatial tions have been proposed for it (e.g. Platnick, expression of the resulting UCs will be candi- 1991; Harold & Mooi, 1994; Morrone, 1994; dates for areas of endemism, with the species Humphries & Parenti, 1999; Szumik & Golo- belonging to each UC strictly endemic. Addi- boff, 2004). tional information about these candidate areas Identifying and delimiting these basic of endemism can be gathered studying the phy- units of analysis are the fundamental steps, logenetic relationships of the involved taxa or although the most problematical, in a historical ecological variables of the points of occurrence 18 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 64 (1): 17-31, March 2016 (Humphries & Parenti, 1999; Dos Santos et al., NAM analysis: The Prepops distribu- 2008, 2011; Torres-Miranda et al., 2013). tional patterns were studied through NAM The goal of this paper is to explore the based in sympatric inference (Dos Santos et al., diversity of the genus Prepops together with its 2011). The NAM analysis was implemented spatial expression in order to identify areas of using the software R (“R Development Core endemism and areas of high species richness Team”, 2011) through the package SyNet (Dos in the Americas. Additionally, we used spatial Santos, 2011) which is used for network analy- data to propose habitat preferences and geocli- sis. NAM estimates the minimum spanning matic variables, important for each group of tree (MST) for each species and calculates the co-occurring species. orthodromic distances (distances calculated over Earth surface), resulting in two matrices of special association. The inference instance MATERIAL AND METHODS yielded weighted matrices of spatial associa- tion between species, the ACSH matrix and the Compiling records: Each species of Pre- topological resemblance matrix. These weight- pops was assigned at least with one spatial ed matrices were dichotomized using a thresh- point based on locality data associated with old to generate a binary matrix corresponding museum specimens and geographic records to the basal network to be analyzed by NAM. in the literature. When this information was NAM was oriented to identify
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