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Coupling Impoverishment Analysis and Partitioning of Beta Diversity Allows a Comprehensive Description of Odonata Biogeography in the Western Mediterranean

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Coupling Impoverishment Analysis and Partitioning of Beta Diversity Allows a Comprehensive Description of Odonata Biogeography in the Western Mediterranean Org Divers Evol (2014) 14:203–214 DOI 10.1007/s13127-013-0161-3 ORIGINAL ARTICLE Coupling impoverishment analysis and partitioning of beta diversity allows a comprehensive description of Odonata biogeography in the Western Mediterranean Markus Heiser & Leonardo Dapporto & Thomas Schmitt Received: 31 March 2013 /Accepted: 24 October 2013 /Published online: 14 November 2013 # Gesellschaft für Biologische Systematik 2013 Abstract Islands host a subset of organisms occurring at their Simpson compared to Sørensen index, and indicator species sources, and these assemblages are usually dominated by the from islands where unpredicted to occur by impoverishment most generalistic and dispersive species. In this study, we aim analysis. This suggests that island species predicted absent to identify which species are missing on islands and which determine most of an island’s turnover pattern, thus ecological traits are responsible for differential occurrence. encompassing fundamental biogeographic information. Due Then, we apply this information to beta diversity analyses. to their absence on nearest sources, they are also at higher risk As a study group and area, we selected the Odonata in the of extinction, and deserving of special conservation effort. Western Mediterranean. Based on the presence/absence of 109 species, we applied a series of analyses at both commu- Keywords Dragonflies . Island biogeography . Community nity and individual species level. The islands of the Balearics, level . Species assemblages . Species impoverishment . Corsica, Sardinia and Malta are highly impoverished, but Dispersal capacity . Species-area relationship Sicily is not. Non-parametric multivariate adaptive regression splines predicted the occurrence of individual species on each island. Principal component analysis recognised differences Introduction between damselflies (Zygoptera) and dragonflies (Anisoptera), but members of the two suborders have similar The equilibrium theory of island biogeography (MacArthur occurrences on islands, and island occurrence is determined and Wilson 1967) is one of the most challenging theories in mostly by species’ frequencies at source and by their degree of ecology. One of this theory’s most important assumptions generalism. Island species predicted correctly to occur on predicts that immigration and extinction reach an equilibrium islands showed opposite characteristics to species unpredicted after an island-specific time of constant environmental condi- to occur and being present. The similarity pattern highlighted tions, thus stabilising species richness. Consequently, mature by turnover (Simpson index) is clearer than that obtained by island communities should host a defined number of species non-partitioned beta diversity (Sørensen index). In fact, indi- as a function of their area and isolation (MacArthur and cator value analyses revealed more indicator species for the Wilson 1967; Emerson and Gillespie 2008; Lomolino et al. 2010; Losos and Ricklefs 2010). Furthermore, the biotic com- position of the neighbouring areas is of major importance Electronic supplementary material The online version of this article because it represents the source of potential immigrants (doi:10.1007/s13127-013-0161-3) contains supplementary material, which is available to authorized users. (Williamson 1981; Dennis and Shreeve 1996; Santos et al. : 2011). M. Heiser T. Schmitt This line of reasoning introduces a corollary of the equilib- Department of Biogeography, Faculty of Geography / Geosciences, Trier University, 54286 Trier, Germany rium theory: due to their isolation, and the limited variety of ecological niches and geographic extent, islands usually host a L. Dapporto (*) limited fraction of their source fauna and flora and are thus Centre for Ecology, Environment and Conservation, Department of “impoverished” compared to their adjoining mainland regions Biological and Medical Sciences, Oxford Brookes University, Headington, Oxford OX3 0BP, UK (Williamson 1981; Whittaker and Fernández-Palacios 2007). e-mail: [email protected] The relationships between numbers of species occurring on 204 M. Heiser et al. islands and island characteristics have been tested in a plethora should be strong relationships between species member- of studies (see Whittaker and Fernández-Palacios 2007; Losos ships and their ecological traits; in such cases, SpaP should and Ricklefs 2010 for recent reviews). However, the “hidden” represent a very small fraction of species. Nevertheless, side of richness (i.e. the quantity and identity of species they usually occur, and have been mostly explained, as lacking on an island as a consequence of its geographic and relict island populations whose occurrence cannot be ex- ecological characteristics) has so far been mostly neglected. plainedonthebasisofmainlanddistribution(Dapportoand The amount of missing species can be assessed with a rela- Dennis 2009, 2010; Dapporto 2011). tively simple comparison of richness between islands and The results obtained for three groups of Lepidoptera neighbouring mainland areas, which are assumed not to be (Papilionidea, Hesperidae and Zygaena moths, Dapporto impoverished (Rosenzweig 1995; Whittaker and Fernández- and Dennis 2009, 2010;Dapporto2011)showedsomesimi- Palacios 2007; Dapporto and Dennis 2009). larities among these groups, but also effects deriving from Source species do not have the same probability to enter an different dispersal ability. Thus, the Zygaena moths, with their island fauna since they differ in dispersal and colonization poor dispersal, showed a more impoverished pattern and a abilities. In this context, larger and less isolated islands offer higher frequency of relict populations than members of other chances also for less opportunistic species (Diamond 1975; groups. Santos et al. 2011;Dennisetal.2012). Deviations from this Moreover, a recent debate on partitioning beta diversity expected pattern occur when an island has been colonised provided new insights into the possibility of separating the from different source assemblages. Such island faunas are effects of differences in richness/nestedness patterns and dif- largely the consequence of stochastic events of immigration ferences due to species replacement (turnover) in determining and extinction, and the spatial complexity deriving from these inter-area dissimilarities (Koleff et al. 2003;Baselga2010; historic phenomena. Long-term processes can also produce Carvalho et al. 2012). The turnover component of diversity unexpected patterns, and species may survive on islands lon- can represent a fundamental and unbiased component for ger than on the mainland, thus producing unexpected distri- identifying biogeographic regionalisation (Kreft and Jetz butions (Masini et al. 2008; Médail and Diadema 2009; 2010); indeed, it encompasses the decisive role of endemic Dapporto et al. 2011, 2012; Dapporto and Bruschini 2012), and non-widespread species (those replacing others in restrict- or they can evolve into endemic taxa (Whittaker and ed island areas). Fernández-Palacios 2007;Kudrnaetal.2011). Endemic and From this perspective, there is strong potential that SpaP relict species represent primary targets for biodiversity con- species, occurring on islands but most likely absent in the servation, and their identification represents an important nearest and most similar mainland units, account for most challenge. However, biogeographic information retained by island turnover patterns. In this study, we couple, for the first the reduced set of endemic or subendemic species can be time, the Dapporto and Dennis (2009) method with hidden by the high frequency of widespread species when partitioning of biodiversity as introduced by Baselga (2010). diversity is examined at the community level. We aim to identify specific island and mainland species re- Therefore, Dapporto and Dennis (2009) provided a sponsible for the turnover component and to recognise their method for (1) the evaluation of island impoverishment, putative dynamics responsible for island occurrence by im- (2) the identification of species predicted to be missing on poverishment analysis. islands, (3) the detection of ecological traits of species In doing so, we selected Odonata, an insect order responsible for differential occurrence and, finally, (4) the characterised by two suborders: the well dispersing dragon- demarcation of potential relict populations. In practice, flies (Anisoptera) and the poorly dispersing damselflies impoverishment analysis distinguishes between species (Zygoptera) (Corbet 2004). As study region, we selected the predicted to be present (Spp) and species predicted to be Western Mediterranean, which was also used in previous absent (Spa) at each island. Spp are subsequently classified studies, so as to allow a direct comparison with Lepidoptera. in those actually present (SppP) and those unexpectedly The distribution of the Odonata is well assessed at the regional absent (SppA), while Spa can be absent (SPAA) or unex- scale in the study area, thus allowing almost unbiased pectedly present (SpaP). SPAA represents species whose modelling. distributions do not fit the geographic characteristics of islands, and actually are not found there. From this perspec- tive, they have no biogeographic relevance. Distinctions Material and methods among the remaining three groups (SppP, SppA, SpaP) are fundamental keys to the recognition of deterministic Study area and study group factors in island occurrences. SppA represents determinis-
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