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Apidae: Euglossini) View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Biblioteca Digital da Produção Intelectual da Universidade de São Paulo (BDPI/USP) Universidade de São Paulo Biblioteca Digital da Produção Intelectual - BDPI Sem comunidade WoS 2012 The Ecological Basis for Biogeographic Classification: an Example in Orchid Bees (Apidae: Euglossini) NEOTROPICAL ENTOMOLOGY, LONDRINA,, v. 41, n. 6, supl. 4, Part 1-2, pp. 442-449, DEC, 2012 http://www.producao.usp.br/handle/BDPI/37124 Downloaded from: Biblioteca Digital da Produção Intelectual - BDPI, Universidade de São Paulo Neotrop Entomol (2012) 41:442–449 DOI 10.1007/s13744-012-0069-1 ECOLOGY, BEHAVIOR AND BIONOMICS The Ecological Basis for Biogeographic Classification: an Example in Orchid Bees (Apidae: Euglossini) 1,2 2 APARRA-H ,GNATES-PARRA 1Depto de Biologia, Fac de Filosofia Ciências e Letras de Ribeirão Preto, Univ de São Paulo, Ribeirão Preto, SP, Brasil 2Lab de Investigaciones en Abejas-LABUN, Depto de Biología, Fac de Ciencias, Univ Nacional de Colombia, Bogotá, Colombia Keywords Abstract Beta diversity, biogeography, community Biogeography has been difficult to apply as a methodological approach ecology, correspondence analysis, euglossine because organismic biology is incomplete at levels where the process of bees, SHE analysis formulating comparisons and analogies is complex. The study of insect biogeography became necessary because insects possess numerous Correspondence evolutionary traits and play an important role as pollinators. Among A Parra-H, Depto de Biologia, Fac de insects, the euglossine bees, or orchid bees, attract interest because the Filosofia Ciências e Letras de Ribeirão Preto, Univ de São Paulo-USP, Av. Bandeirantes, study of their biology allows us to explain important steps in the 3900, Bairro Monte Alegre, CEP evolution of social behavior and many other adaptive tradeoffs. We 14040-901 Ribeirão Preto, SP, Brasil; analyzed the distribution of morphological characteristics in Colombian [email protected] orchid bees from an ecological perspective. The aim of this study was to Edited by Fernando B Noll – UNESP observe the distribution of these attributes on a regional basis. Data Received 13 March 2012 and accepted 26 corresponding to Colombian euglossine species were ordered with a June 2012 correspondence analysis and with subsequent hierarchical clustering. Published online 24 August 2012 Later, and based on community proprieties, we compared the resulting hierarchical model with the collection localities to seek to identify a * Sociedade Entomológica do Brasil 2012 biogeographic classification pattern. From this analysis, we derived a model that classifies the territory of Colombia into 11 biogeographic units or natural clusters. Ecological assumptions in concordance with the derived classification levels suggest that species characteristics as- sociated with flight performance, nectar uptake, and social behavior are the factors that served to produce the current geographical structure. Introduction Seberg 1986,Mackeyet al 2008). Biogeographical approaches have a strong ecological background be- As an empirical, multidisciplinary, and theoretical ap- cause trends linking organisms with their environment proach, biogeography aims to explain the conditions are the principal modulators of such patterns (Pianka and natural phenomena affecting the distribution of 1994,Thompson2005). organisms (Craw 1983,Wiley1988, Noonan 1988, Insects represent 79% of the known global fauna Wilson 1991, Crisci & Morrone 1992, Morrone 2001, (Noonan 1988). Among insects, bees (Hymenoptera: García-Barros et al 2002,Mackeyet al 2008). Apoidea) include the most important pollinator taxa Different perspectives have been applied to explain (Danforth et al 2006, Michener 2007). Bees probably orig- such patterns. It is probable that Darwin (1859)made inated ca. 125 my ago in extratropical and open areas the first contributions to biogeography with his disper- during the Middle Cretaceous (Roubik 1989), diversifying sionist theory (Wilson 1991). Due to the lack of an 125 to 90 my ago in the tropics as they simultaneously adequate ecological, taxonomic, and systematic consen- tracked the radiation of the angiosperms (Michener 1979). sus, biogeography remains inconsistent (Craw 1983, A valid theory explains that bees originated from a wasp- Ecological Basis for Biogeographic Classification 443 like ancestor (Engel 2000, Poinar & Danforth 2006, differences within the tribe has produced discussions Michener 2007), subsequently abandoning their predatory about the relationships among the species (Cameron habits to become pollen and nectar feeders as they ex- 2004, Michel-Salzat et al 2004, Michener 2007). panded to temperate regions (Silveira et al 2002, Poinar & Although the Euglossini are accurately characterized as Danforth 2006). corbiculate (i.e., the hypothesis of monophyly is sup- The distribution of bees is frequently analyzed according ported), the biology of the tribe differs from that of its to distinct regional or vegetation terms that are otherwise relatives in aspects such as the lack of true social behavior difficult to define (Roubik & Hanson 2004), and few em- (Noll 2002, Cameron 2004, Nates-Parra 2005) and sub- pirical studies on the geographic distribution of euglossines stantial morphological variability (Michener 2007). Such have been performed (Dick et al 2004, Nemésio 2007b, dissimilarity should contribute to the evolutionary success Ramírez et al 2010). of the Euglossini (Roubik & Hanson 2004) as well as to Of an estimated 20,000–30,000 species worldwide, their habitat use (Otero & Sandino 2003, Sandino 2004, only 2% to 5% of bees display true social behavior Uehara-Prado & Garófalo 2006, Parra-H & Nates-Parra (Michener 2007). Approximately 500 eusocial bee species 2007). are reported in the neotropical region, and 240 occur in Adaptation can be linked to certain morphological and Colombia (Nates-Parra 2005, Michener 2007). This neo- behavioral features. In groups that use traplining foraging tropical diversity includes the orchid bees (Euglossini). strategies, for example, tongue length is related to the These bees are predominant in lowland rain forests preference for the nectar of certain species (Ackerman et (Roubik & Hanson 2004) and are found in habitats from al 1982, Ackerman 1985, Kato et al 1992, Roubik & Hanson sea level to 2,000 m asl (Ramírez et al 2002, Roubik & 2004). In addition, the atmospheric pressure influences the Hanson 2004, Nates-Parra et al 2006). It is commonly process of nectar intake when the tongue is extruded assumed that orchid bees originated between the Andean making feeding efficiency (in relation to viscosity) environ- foothills and the Amazon region more than 20 my ago with mentally dependent (Borrell 2004, 2007a, b, Borrell & a significant radiation of the group occurred in the Krenn 2006). Because nectar resources are equally Miocene/Pleistocene (Engel 1999, Dick et al 2004, Roubik exploited by males and females (Ackerman 1985), intraspe- and Hanson 2004). Recent evidence suggests that euglos- cific variation for tongue length should not be expected sines date from the Eocene and that it is not necessarily between the sexes (Roubik personal communication). clear that they originated in South America (Ramírez et al Body size is involved in thermoregulatory capacity and in 2010). The current distribution of euglossine bees should the optimization of the strength of vertical flight. By in- reflect geological, environmental, and climatic factors in creasing the flux of hemolymph from the thorax to the addition to the occurrence of resource competition, para- abdomen, large bees can release heat and improve their sites and predators (Nemésio & Silveira 2006, Roubik & flight capacity. This ability can be used, for example, for Hanson 2004). flight across open areas exposed to solar radiation (Inouye Given the relatively recent and widespread distribution 1975,Armbruster&Berg1994,Dudley1995,Borrell& of orchid bees (Dick et al 2004), the ancestors of the group Medeiros 2004). Dwarf forms have also been reported in should occur at the time of the uplift of the Andean the Euglossini (Roubik 2004). The frequency of these forms cordillera and at a level of uplift that reached no greater is unclear, and their effect on interspecific variability can- than 40% of the current altitude of the cordillera (Ramírez not be assessed. et al 2010). Bee community structures did not appear in Social behavior significantly contributes to defense association with the uplift (Gregory-Wodzicki 2000, Dick et against parasites and to resource optimization (Roubik & al 2004, Michener 2007). In contrast to the cordillera, the Hanson 2004), and it occurs in each genus of euglossines. Amazon and part of Orinoquia were regions where no Eufriesea Cockerell is predominantly solitary, although co- important geological activity occurred. In these regions, operative forms are reported (Kimsey 1982, Eberhard 1989, the most relevant source of variation was the vegetation Garófalo 1994, Soucy et al 2003). More complex levels of changes caused by glacial cycles (Colinvaux et al 2000). The cooperation exist in Eulaema Lepeletier and Euglossa Caribbean was influenced by volcanic activity (most likely in Latreille (Bennett 1965, Santos & Garófalo 1994, Augusto the Antilles). In this region, tectonic movements prevailed & Garófalo 2004, 2009, 2010). Exaerete Hoffmannsegg and in the Late Cretaceous. The ocean dynamics should affect
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