Abstract

BackgroundSpecies thermal requirements are one of the principal determinants of their ecology and biogeography, although our understanding of the interplay between these factors is limited by the paucity of integrative empirical studies. Here we use empirically collected thermal tolerance data in combination with molecular phylogenetics/phylogeography and ecological niche modelling to study the evolution of a clade of three western Mediterranean diving beetles, the Agabus brunneus complex.ResultsThe preferred mitochondrial DNA topology recovered A. ramblae (North Africa, east Iberia and Balearic islands) as paraphyletic, with A. brunneus (widespread in the southwestern Mediterranean) and A. rufulus (Corsica and Sardinia) nested within it, with an estimated origin between 0.60-0.25 Ma. All three species were, however, recovered as monophyletic using nuclear DNA markers. A Bayesian skyline plot suggested demographic expansion in the clade at the onset of the last glacial cycle. The species thermal tolerances differ significantly, with A. brunneus able to tolerate lower temperatures than the other taxa. The climatic niche of the three species also differs, with A. ramblae occupying more arid and seasonal areas, with a higher minimum temperature in the coldest month. The estimated potential distribution for both A. brunneus and A. ramblae was most restricted in the last interglacial, becoming increasingly wider through the last glacial and the Holocene.ConclusionsThe A. brunneus complex diversified in the late Pleistocene, most likely in south Iberia after colonization from Morocco. Insular forms did not differentiate substantially in morphology or ecology, but A. brunneus evolved a wider tolerance to cold, which appeared to have facilitated its geographic expansion. Both A. brunneus and A. ramblae expanded their ranges during the last glacial, although they have not occupied areas beyond their LGM potential distribution except for isolated populations of A. brunneus in France and England. On the islands and possibly Tunisia secondary contact between A. brunneus and A. ramblae or A. rufulus has resulted in introgression. Our work highlights the complex dynamics of speciation and range expansions within southern areas during the last glacial cycle, and points to the often neglected role of North Africa as a source of European biodiversity.

Highlights

  • Species thermal requirements are one of the principal determinants of their ecology and biogeography, our understanding of the interplay between these factors is limited by the paucity of integrative empirical studies

  • Morphology The three species differed in body size, as measured with BL and BW (MANOVA, Roy’s greatest root F = 292.134, P < 0.001), with pairwise differences being significant for all comparisons (Table 1)

  • Agabus ramblae and A. brunneus were fully separated by aedeagus in ventral view (AD), with no intermediate specimens, whilst the specimens identified as A. rufulus had an intermediate, overlapping shape (Additional file 10: Figure S3)

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Summary

Introduction

Species thermal requirements are one of the principal determinants of their ecology and biogeography, our understanding of the interplay between these factors is limited by the paucity of integrative empirical studies. Ambient temperature affects all biological processes [1,2], especially in ectotherms [3], and is usually assumed to be one of the main determinants We attempt such an integrative approach in a clade of diving beetles that has diversified and expanded its range in the western Mediterranean region during the late Pleistocene, the Agabus brunneus complex [16]. In most cases, unglaciated areas are seen as refugia for northern species, little attention being paid to evolutionary and biogeographical processes in them, other than those which affected these species [21,22] In contrast to this view, the current diversity of the Mediterranean area is increasingly seen to result from processes which are not directly related to the range movements of northern species during glacial-interglacial cycles [23,24,25,26], but our understanding of its origin remains fragmentary, for highly speciose groups such as most insects

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