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Geographic Location and Phylogeny Are the Main Determinants of the Size of the Geographical Range in Aquatic Beetles Pedro Abellán1,2* and Ignacio Ribera1

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Geographic Location and Phylogeny Are the Main Determinants of the Size of the Geographical Range in Aquatic Beetles Pedro Abellán1,2* and Ignacio Ribera1 Abellán and Ribera BMC Evolutionary Biology 2011, 11:344 http://www.biomedcentral.com/1471-2148/11/344 RESEARCHARTICLE Open Access Geographic location and phylogeny are the main determinants of the size of the geographical range in aquatic beetles Pedro Abellán1,2* and Ignacio Ribera1 Abstract Background: Why some species are widespread while others are very restricted geographically is one of the most basic questions in biology, although it remains largely unanswered. This is particularly the case for groups of closely related species, which often display large differences in the size of the geographical range despite sharing many other factors due to their common phylogenetic inheritance. We used ten lineages of aquatic Coleoptera from the western Palearctic to test in a comparative framework a broad set of possible determinants of range size: species’ age, differences in ecological tolerance, dispersal ability and geographic location. Results: When all factors were combined in multiple regression models between 60-98% of the variance was explained by geographic location and phylogenetic signal. Maximum latitudinal and longitudinal limits were positively correlated with range size, with species at the most northern latitudes and eastern longitudes displaying the largest ranges. In lineages with lotic and lentic species, the lentic (better dispersers) display larger distributional ranges than the lotic species (worse dispersers). The size of the geographical range was also positively correlated with the extent of the biomes in which the species is found, but we did not find evidence of a clear relationship between range size and age of the species. Conclusions: Our findings show that range size of a species is shaped by an interplay of geographic and ecological factors, with a phylogenetic component affecting both of them. The understanding of the factors that determine the size and geographical location of the distributional range of species is fundamental to the study of the origin and assemblage of the current biota. Our results show that for this purpose the most relevant data may be the phylogenetic history of the species and its geographical location. Background However, there are still fundamental questions unre- Why some species are widespread while others are very solved, best exemplified by the fact that closely related restricted geographically is one of the most basic ques- species often display dramatic differences in range size tions in biology, although it remains basically unan- for largely unknown reasons. Tests of these differences swered, despite the sustained interest from ecologists, remain relatively scarce, have been performed for exam- biogeographers and evolutionary biologists (e.g., [1-5]). ples of very few taxa (usually vertebrates), and generally A range of ecological and evolutionary explanations fail to address the potentially confounding effects of the have been suggested for the observed range size varia- phylogenetic relatedness of species. tion, based on differences in niche breadth or environ- In this work we aim to test some likely determinants mental tolerance, body size, population abundance, of the size of the geographical range in a phylogenetic latitude, environmental variability, colonization and comparative framework. Closely related species are extinction dynamics, and dispersal ability [3,6-8]. expected to show more similarity than those that are distantly related because they share more common evo- * Correspondence: [email protected] lutionary history [9,10]. How range size evolves and the 1Institute of Evolutionary Biology (CSIC-UPF), Passeig Maritim de la extent of heritability of the geographical range sizes of Barceloneta 37, 08003 Barcelona, Spain species has received much attention in the last years Full list of author information is available at the end of the article © 2011 Abellán and Ribera; licensee BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Abellán and Ribera BMC Evolutionary Biology 2011, 11:344 Page 2 of 15 http://www.biomedcentral.com/1471-2148/11/344 [11-14], as evidence for range size heritability would range sizes [14,25], with species with a “privileged” geo- have important implications for ecology, evolution, and graphical position displaying higher range-sizes. For biogeography [15]. Although a number of studies have example, latitudinal gradients in geographic range size investigated the existence of phylogenetic signal in range (Rapoport’s rule) have been extensively studied and size in a variety of clades and from a wide range of ana- documented [6,26,27]. Evidence supporting that range lytical approaches, the patterns found have been mixed sizes increase with latitude in the Palearctic and Nearc- and the “heritability” of range size remains a contentious tic above 40°-50°N has been found in a number of ter- issue [13,14]. Phylogenetic comparative methods applied restrial groups [26], but the extent to which this is a to whole lineages and not only species-pairs (e.g., general pattern remains contentious and has rarely been [16-18]) may provide a more robust and powerful tested in a phylogenetic framework. approach to estimate the phylogenetic signal in range 4) Age and area. We also consider the possible rela- size. tionship between age and area, which requires to be Among the potential determinants of range size we tested in the context of a phylogeny even if not in the include ecological tolerance, dispersal ability, geographic same comparative framework as the previous factors. location, and age of the species. For all of them we test Originally proposed to explain the distribution of the their phylogenetic signal, as well as the phylogenetic sig- endemic flora of some islands [28], in its basic form the nal of the size and location of the range itself. “age and area” hypothesis states that the older a species 1) Ecological tolerance. A broad ecological niche is the more likely it is to have occupied a wider geogra- allows a species to persist in a wide range of different phical area. More precisely, it could be expected that environments, while a narrow niche restricts a species species have a “life cycle” from origin to extinction that to the few places where its niche requirements are met could be described through a variety of simple models [19,20]. Hence, species with broad niches should be dis- (see [26] for a review). Although a number of studies tributed over a wider range of different biomes than have examined the evidence for geographic range size species with narrower ecological requirements, leading changes over evolutionary time across a wide range of to larger geographical ranges [21]. When these biomes clades (but never in insects), no consistent evidence has are distributed equally along environmental gradients emerged to support any particular model [29]. An inher- this poses the problem that species with lager ranges ent limitation of all these studies is that a direct test of will inevitably overlap more different biomes. However, theageandareamodelrequiresthegeographicrange in the western Palaearctic (the centre of distribution of size of a species or a clade to be known throughout its most of our studied lineages, see below) this problem is evolutionary history [3]. This is usually not possible partly alleviated by the very heterogeneous distribution without extensive palaeontological data. Hence, a differ- of environmental gradients. In this case, those species ent approach has to be taken in neontological studies, occurring in large biomes (e.g., [20,21]) would have considering interspecific variation in range sizes of con- large range sizes (as found e.g. by [14]), but they should temporary species as a reflection of the intraspecific not necessarily occupy more biomes than species with relationship [30]. The examination of the interspecies smaller ranges. relationship between geographic range size and age 2) Dispersal ability. As a surrogate measure of disper- could thus be used as a surrogate of the transformation sal ability we use water flow, as previous studies in of range size with age in individual species [29,31]. freshwater invertebrates have established the relation- We use a set of lineages of closely related species of ship between main habitat type (lotic or lentic) and the different families of aquatic Coleoptera to investigate the size of the geographical range [22,23]. Lentic species relative role of different factors on determining the size should have better dispersal abilities due to the shorter of the geographical range of species. The Western geological duration of their habitats, and display on Palearctic water beetle fauna is a suitable model to average larger geographical ranges than the species inha- study range size issues, as water beetles are a rich and biting the more persistent lotic habitats [24]. However, well-known insect group in both Europe and the Medi- the role of habitat constraints in aquatic organisms has terranean Basin, exhibiting a high level of endemism but yet not been assessed from a phylogenetic comparative also with species widely distributed across the Palearctic framework in lineages in which there are species inha- and Holartic regions [32-34]. Spatial determinants of biting both habitat types. range size and temporal patterns of
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