Ecological Patterns Strongly Impact the Biogeography of Western Palaearctic Longhorn Beetles (Coleoptera: Cerambycoidea)
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Org Divers Evol (2017) 17:163–180 DOI 10.1007/s13127-016-0290-6 ORIGINAL ARTICLE Ecological patterns strongly impact the biogeography of western Palaearctic longhorn beetles (Coleoptera: Cerambycoidea) Francesco Vitali1,2 & Thomas Schmitt2,3,4 Received: 30 December 2015 /Accepted: 17 May 2016 /Published online: 6 June 2016 # Gesellschaft für Biologische Systematik 2016 Abstract We aim to unravel the biogeographic structuring of the Mediterranean region and underline the importance of western Palaearctic longhorn beetles with focus on the loca- Provence, Crimea and Crete as such refugia. Crete even might tion of different refugia, barriers to dispersal and postglacial be an area of old endemism. The Atlanto- and the Ponto- range expansions with their particular filters. The interaction Mediterranean regions are more strongly structured than as- of different ecological features with these structures is sumed in classical biogeography. Mediterranean assemblages analysed. The western Palaearctic was divided into 95 geo- are mostly composed of non-flying species, root feeders and graphic entities. We produced presence-only matrices for all species with small distributions not found outside their glacial 955 Cerambycoidea species autochthonous to this area and refugia. Tree feeders left their glacial retreats with their host derived species richness distributions and extracted faunal re- plants. These range dynamics result in biogeographic struc- gions and faunal elements by cluster analyses and principal tures with several dispersal barriers and filters composed of component analyses. Similar analyses were performed for mountains, sea straits and climatic conditions. sub-families and ecological groups. Longhorn beetles show a strong biogeographic structuring in the western Palaearctic. Keywords Biogeographic filters . Ecological features . Species numbers strongly decrease to the north and west. Less Glacial refugia . Island biogeography . Postglacial mobile species and root feeders mostly contribute to the fauna colonisation . Species diversity of the Mediterranean region, whilst mobile species are more widespread. Feeders on broad-leaved trees dominate in west- ern Europe, whilst feeders on coniferous trees are most impor- Introduction tant in northern Europe. Our results support multiple refugia in Since the early 1990s, considerable improvements in statisti- Electronic supplementary material The online version of this article cal methods and computer capacities have led to remarkable (doi:10.1007/s13127-016-0290-6) contains supplementary material, progress in classical biogeographic research. The possibility which is available to authorized users. of applying advanced statistical analyses to large groups of animals or plants has led to a better understanding of the * Francesco Vitali biogeographic history of large geographic entities, their faunal [email protected] elements and faunal regions (Podani 1997; Baselga 2010, 2012; Dobrovolski et al. 2012;Dapportoetal.2013, 2015; 1 National Museum of Natural History, rue Münster 24, Baselga and Leprieur 2015). Consequently, a better under- 2160 Luxembourg, Luxembourg standing of spatiotemporal range dynamics over entire conti- 2 Biogeography, Faculty of Regional and Environmental Sciences, nents has resulted (e.g. pioneer work by Dennis et al. 1991, Trier University, 54286 Trier, Germany 1995, 1998). 3 Senckenberg German Entomological Institute, Eberswalder Straße Up to now, such analyses based on advanced statistical 90, 15374 Müncheberg, Germany methods have been conducted on a variety of different groups 4 Zoology, Institute of Biology, Faculty of Natural Sciences I, Martin of vertebrates, e.g. birds (Hagemeier and Blair 1997), mam- Luther University Halle-Wittenberg, 06099 Halle, Saale, Germany mals (Baquero and Telleria 2001), freshwater fishes (Reyjol 164 F. Vitali, T. Schmitt et al. 2007), coral reef fishes (Mouillot et al. 2013), amphib- new features, such as their multifarious relations with plants ians and reptiles (Araújo et al. 2008) or even on a combination and the resulting highly differentiated use of habitat structures, of different groups including vascular plants (Holt et al. 2013). as well as the existence of winged and wingless species, which However, proper biogeographic analyses on invertebrates are entail significant differences in dispersal capacities. The aim much more scarce and comprise mainly studies on butterflies of this study is therefore to obtain a comprehensive under- (Dennis et al. 1991, 1995, 1998; Dapporto et al. 2012, 2014, standing of the biogeography of the longhorn beetles of the 2015;Rákosyetal.2013) but also dragonflies and damselflies western Palaearctic, to delimit the faunal regions and (Heiser and Schmitt 2010, 2013; Rákosy et al. 2013;Heiser localisation of faunal elements in our study region, and to et al. 2014), darkling beetles (Fattorini and Ulrich 2012; analyse the influence of different ecological groups and dif- Fattorini 2014), and spiders (Ysnel et al. 2008); the latter more ferent dispersal capacities on the resulting biogeographic pat- in a conservation context. Preliminary data on the country terns. Finally, we discuss these patterns in comparison with level have also been published for longhorn beetles (Baselga those already obtained by analysis of other groups. Ultimately, 2008). These analyses have considerably enhanced our bio- the study enhances the general understanding of the biogeog- geographic understanding of the western Palaearctic. They raphy of the western Palaearctic realm. reveal the geographic location of centres of endemism and faunal elements in the Mediterranean refugial regions. Faunal regions are characterised by coherent species assem- Materials and methods blages within a region that differ from those of other regions. Such differences have often resulted from range expansion Study group and taxonomy applied (e.g. out of one of the Mediterranean refugia) or are due to a consistent filtering during the colonisation process of an entire A large number of species of cerambycids occur in the western region, as for example the colonisation of the British Isles by a Palaearctic. However, several taxonomic problems are still characteristically impoverished fauna (Dennis et al. 1991; under debate, especially with regard to the more eastern re- Heiser and Schmitt 2010). gions. To achieve a consistent basis for the taxonomic setting Most of the examples mentioned above involve animal in this contribution, we adopted the classification presented in groups with moderate to very high mobility of most of their the catalogue by Löbl and Smetana (2010) as our baseline. representatives. Therefore, it should be highly interesting to However, to reflect the numerous corrections have already gain a better understanding of the biogeographic structures of been made to this (e.g. Danilevsky 2010a, 2010b, 2012a, taxonomic groups most of whose members are less mobile, 2012b, 2012c, 2012d, 2013; Özdikmen 2010, 2011a, b;Löbl such as insects with restricted dispersal capacity. Hence, the and Smetana 2011;Miroshnikov2011), we adjusted our tax- larger beetle families seem to represent suitable target groups onomic system to the quickly modifiable online catalogue for such analyses. However, despite their popularity amongst available at www.biolib.cz. professionals and amateurs, beetles have rarely been investi- Taxa of doubtful status as valid species (e.g. Agapanthia gated in this way (but see Baselga 2008; Fattorini and Ulrich intermedia, A. suturalis) and recently described sibling spe- 2012;Fattorini2014). cies (e.g. Leiopus linnei) were excluded from the analyses For these reasons, we performed an in-depth analysis of the because distribution data on these taxa are still largely uncer- biogeographic structures of the longhorn beetles of the west- tain, incomplete, or largely missing. Furthermore, a large per- ern Palaearctic. The cerambycids (superfamily centage of longhorn beetle species (37.2 %) belong to just Cerambycoidea) of this region represent a suitable model three not xylophagous genera feeding on herbaceous plants group for this analysis for several reasons. The number of (i.e. Dorcadion, Agapanthia and Phytoecia). Species in these autochthonous species (i.e. 955) is relatively high, allowing genera have often been described on the basis of only few reliable analysis, but is still manageable; the group is popular specimens or using doubtful characters. The revisions of amongst entomologists, facilitating the derivation of detailed Spanish (Vives 1976, 1983, 2000) and Greek Dorcadion spe- distribution data for the construction of plausible regionalised cies (Pesarini and Sabbadini 2004, 2007, 2008, 2010)have maps. It consists of several ecological groups having trophic greatly reduced their numbers. Consequently, the number of interactions with different taxonomic groups of plants and cerambycid species in Turkey is likely to be overestimated. further specific requirements on their use, allowing an inter- Özdikmen (2010), for example, listed 200 Dorcadion species, pretation of the impact of ecological specialisation on distri- whilst Löbl and Smetana (2010) mentioned only 150. bution patterns, and whilst some groups are apterous, others Nevertheless, even if the taxonomy of the latter authors is are relatively good flyers so that different dispersal capacities applied, Turkey still shows an extraordinarily high diversity can be compared. of species and genera. Thus, cerambycids as a group not only possess character-