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Areas of Endemism in the Atlantic Forest: Quantitative Biogeography Insights from Orchid Bees (Apidae: Euglossini) André R.S

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Areas of Endemism in the Atlantic Forest: Quantitative Biogeography Insights from Orchid Bees (Apidae: Euglossini) André R.S Areas of endemism in the Atlantic Forest: quantitative biogeography insights from orchid bees (Apidae: Euglossini) André R.S. Garraffoni, Filipe R. Moura, Anete P. Lourenço To cite this version: André R.S. Garraffoni, Filipe R. Moura, Anete P. Lourenço. Areas of endemism in the Atlantic Forest: quantitative biogeography insights from orchid bees (Apidae: Euglossini). Apidologie, Springer Verlag, 2017, 48 (4), pp.513-522. 10.1007/s13592-017-0494-6. hal-01710383 HAL Id: hal-01710383 https://hal.archives-ouvertes.fr/hal-01710383 Submitted on 15 Feb 2018 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie (2017) 48:513–522 Original article * INRA, DIB and Springer-Verlag France, 2017 DOI: 10.1007/s13592-017-0494-6 Areas of endemism in the Atlantic Forest: quantitative biogeography insights from orchid bees (Apidae: Euglossini) 1,2 2 2 André R.S. GARRAFFONI , Filipe R. MOURA , Anete P. LOURENÇO 1Present address: Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, SP, Brazil 2Departamento de Ciências Biológicas, Universidade Federal dos Vales do Jequitinhonha e Mucuri, Diamantina, MG, Brazil Received 27 March 2016 – Revised 9 January 2017 – Accepted 16 January 2017 Abstract – Orchid bees are endemic to the Neotropics and were sampled more intensively in the Atlantic Forest in the last decade than in that of other Brazilian biomes. In this study, we aimed at identifying the main distributional patterns and areas of endemism of Euglossini orchid bee species in the Atlantic Forest using parsimony analysis of endemism and endemicity analysis. The results of these analyses were partly congruent and supported the idea that the distribution of orchid bees is structured into at least five areas of endemism: Pernambuco/coastal Bahia; Espírito Santo/Rio de Janeiro/south of Minas Gerais; north of Minas Gerais/central Bahia; southeast of Minas Gerais/ northeast, central and coast of São Paulo/central and coastal Paraná; and central/coast of Santa Catarina-Rio Grande do Sul. Most of these areas were consistent with other groups of organisms and indicate the existence of real areas of endemism in the Atlantic Rain Forest. distribution pattern / insects / PAE / NDM / orchid bees 1. INTRODUCTION (non-random distributional congruence among distinct taxa—Morrone 1994) or generalized Biogeography investigates the history of or- tracks; and cladistic testing of the previously hy- ganisms in space using comparative biology as a pothesized homology, to find the secondary bio- background (Morrone 1994). This discipline in- geographic homology (Morrone 2001, 2014). volves the following two primary stages: infer- The most common methods for inferring the ence associated with the primary biogeographic primary biogeographic homology are the parsimo- homology based on a common history, which is ny analysis of endemicity (PAE) and endemicity conjectured by identifying areas of endemism analysis (NDM) (Morrone 1994, 2014;Szumik et al. 2002; Szumik and Goloboff 2004;Nihei 2006). PAE and NDM use biota similarity to eluci- Electronic supplementary material The online version of date faunal distribution patterns. Based on species this article (doi:10.1007/s13592-017-0494-6) contains presence–absence data matrices, PAE treats geo- supplementary material, which is available to authorized users. graphic areas as lines (i.e., taxa) in a traditional data Corresponding author: A. Garraffoni, matrix and taxa distribution as characters, similar to [email protected] a cladistic analysis (Nihei 2006; Morrone 2014). Manuscript editor: Marina Meixner NDM identifies areas of endemism by applying an In order to explain the content of the Suppl. Materials, these legends must be included in the article: either here optimality criterion to evaluate congruence among or in an Annex (or Appendix) at the end of the article species distributions, using the endemicity index, to (please see what I suggest in the attached file). reveal distributional patterns (Szumik et al. 2002; 514 A. R.S. Garraffoni et al. Szumik and Goloboff 2004; Ferrari et al. 2010; 2. MATERIALS AND METHODS Carvalho 2011; Casagranda et al. 2012). Based on the adjustment of species distribution to the ana- Distribution data on orchid bees in the Atlantic lyzed grid, NDM uses an optimality criterion to Forest were compiled from the literature (see assign an endemicity score to a given area supplementary material S1 for references). We re- (Escalante et al. 2013). Both PAE and NDM are corded 1630 points of occurrence of Euglossini. used to delineate areas of endemism, using grid- Taxonomic decisions were based on Moure’sBee cells as operational units without inferring relation- Catalogue (Moure et al. 2012). The taxa analyzed ships or hierarchies between the areas. were represented by 66 Euglossini species, belonging The Atlantic Rain Forest is distributed along to Euglossa (45 spp.), Eufriesea (7 spp.), Eulaema most of the Brazilian coast, having originally oc- (10 spp.), and Exaerete (4 spp.) (supplementary cupied approximately 15% of the Brazilian terri- material S2). As all genera in Euglossini used in the tory. However, the forest has become completely present study have the initial BE^,weusedthe fragmented, now covering less than 8% of its following abbreviations: Eg .forEuglossa ; Ef .for original area (Morellato and Haddad 2000). The Eufriesea ; El .forEulaema ;andEx .forExaerete . Atlantic Forest biome shows great environmental complexity and can support high levels of diver- 2.1. Parsimony analysis of endemicity sity and endemism (Nemésio 2009; DaSilva and Pinto-da-Rocha 2011). This method was originally developed and Although orchid bees (Apidae: Euglossini) implemented by Rosen (1988) and Rosen and now range from the southern USA to southern Smith (1988) and can be used both statically and Brazil and northern Argentina (Nemésio 2009), dynamically (Nihei 2006). Static PAE, which was they are a well-known taxon in the Atlantic Forest applied here, cannot prove or reveal any informa- resulting in numerous sampling sites and a well- tion about processes and is applied to a biota established taxonomy (Nemésio 2009;Nemésio situated within a single geologic age (present), and Vasconcelos 2013). Five genera have been whereas, dynamic PAE is applied to biota from recognized in the classification of Euglossini: two or more separate geological ages (Nihei 2006). Euglossa Latreille 1802, Eufriesea Cockerell Grid-cells are a decisive component in biogeo- 1908, Eulaema Lepeletier 1841, Exaerete graphic analyses (Morrone and Escalante 2002; Hoffmannsegg 1817,andAglae Lepeletier and Contreras-Medina et al. 2007); therefore, the PAE Serville 1825, collectively encompassing more analysis of the entire Atlantic Forest domain was than 250 species (Roubik and Hanson 2004; divided into grid-cells of two different sizes, 1 × 1° Michener 2007;Nemésio2009). Orchid bees are and 3 × 3°. Applying grid-cells of different sizes to commonly known for their metallic hues (differ- a single data set makes it possible to obtain results ent shades of green, blue, violet, and red), medium that are more complete because different quadrant to large body size (8 to 30 mm), and corbiculae sizes can be sensitive to different distributional (Silveira et al. 2002; Michener 2007; Nemésio patterns within the same geographic region (the 2009). use of small grid-cells can produce poorly resolved This study aimed at identifying areas of ende- area cladograms) (Morrone and Escalante 2002; mism in the Atlantic Forest characterized by dis- Contreras-Medina et al. 2007). In addition, a study tributional patterns of orchid bees by means grid- area may contain different levels of endemism in cell analyses (PAE and NDM). Furthermore, it different regions (DaSilva 2008). also complements previous studies that analyzed We only considered an area of the endemism the spatial distribution patterns of orchid bees when the relationship between the grid-cells was along the different subregions of the Neotropics supported by the occurrence of two or more ex- (not only the Atlantic Forest) using ordination clusive species (Bsynapomorphic species^). procedures to describe the groups of assemblages Parsimony analysis was carried out using the according to occurrences of euglossine bees TNT software (Goloboff et al. 2003, 2008). A (Nemésio and Silveira 2007; Sydney et al. 2010). traditional search was executed using 1,000,000 Areas of endemism for euglossine bees 515 replicates and with a tree bisection and reconnec- Alagoas and Pernambuco (areas 7, 8, 14, 15, 16, 17, tion swapping algorithm with five trees saved per 18, 20, and 27), which shared the synapomorphic replicate. The saved trees were analyzed using species Eg . mixta and Eg . milenae. Winclada (Nixon 2002). Strict consensus was The PAE using a 3-degree quadrat matrix re- used to summarize the information obtained from sulted in 45 cladograms with 170 steps. Strict the most parsimonious cladograms. consensus (supplementary material S6)indicated only one trichotomy among the 28 areas and at 2.2. Endemicity analysis least two major clades were observed (Figure 1b). In this case, the following two areas of endemism NDM was originally proposed by Szumik et al. were observed: one major area including the (2002) and Szumik and Goloboff (2004) and was coasts of Paraíba, Pernambuco, Alagoas states, implemented as a computer program by Goloboff the south and southeast coast of Bahia state and (2005). We used the computer program NDM/ the northern coast of Espírito Santo (areas 3, 7, 8, VNDM 2.5 (Goloboff 2005) to test two distinct and 9) that shared the synapomorphic species Eg . grids, with grid-cells of 1 × 1° and 3 × 3°, adiastola , Eg . amazonica , Eg . calycina , Eg .
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