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Historical Biogeography of the Land Snail Cornu Aspersum: New Insights Into the Scenario Inferred from Haplotype Distribution in Both Europe and North Africa

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Historical Biogeography of the Land Snail Cornu Aspersum: New Insights Into the Scenario Inferred from Haplotype Distribution in Both Europe and North Africa Historical biogeography of the land snail Cornu aspersum: new insights into the scenario inferred from haplotype distribution in both Europe and North Africa. Annie Guiller, Luc Madec To cite this version: Annie Guiller, Luc Madec. Historical biogeography of the land snail Cornu aspersum: new insights into the scenario inferred from haplotype distribution in both Europe and North Africa.. BMC Evo- lutionary Biology, BioMed Central, 2010, 10, pp.18. 10.1186/1471-2148-10-18. hal-00464987 HAL Id: hal-00464987 https://hal.archives-ouvertes.fr/hal-00464987 Submitted on 18 Mar 2010 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Guiller and Madec BMC Evolutionary Biology 2010, 10:18 http://www.biomedcentral.com/1471-2148/10/18 RESEARCHARTICLE Open Access Historical biogeography of the land snail Cornu aspersum: a new scenario inferred from haplotype distribution in the Western Mediterranean basin Annie Guiller1*, Luc Madec2 Abstract Background: Despite its key location between the rest of the continent and Europe, research on the phylogeography of north African species remains very limited compared to European and North American taxa. The Mediterranean land mollusc Cornu aspersum (= Helix aspersa) is part of the few species widely sampled in north Africa for biogeographical analysis. It then provides an excellent biological model to understand phylogeographical patterns across the Mediterranean basin, and to evaluate hypotheses of population differentiation. We investigated here the phylogeography of this land snail to reassess the evolutionary scenario we previously considered for explaining its scattered distribution in the western Mediterranean, and to help to resolve the question of the direction of its range expansion (from north Africa to Europe or vice versa). By analysing simultaneously individuals from 73 sites sampled in its putative native range, the present work provides the first broad-scale screening of mitochondrial variation (cyt b and 16S rRNA genes) of C. aspersum. Results: Phylogeographical structure mirrored previous patterns inferred from anatomy and nuclear data, since all haplotypes could be ascribed to a B (West) or a C (East) lineage. Alternative migration models tested confirmed that C. aspersum most likely spread from north Africa to Europe. In addition to Kabylia in Algeria, which would have been successively a centre of dispersal and a zone of secondary contacts, we identified an area in Galicia where genetically distinct west and east type populations would have regained contact. Conclusions: Vicariant and dispersal processes are reviewed and discussed in the light of signatures left in the geographical distribution of the genetic variation. In referring to Mediterranean taxa which show similar phylogeographical patterns, we proposed a parsimonious scenario to account for the “east-west” genetic splitting and the northward expansion of the western (B) clade which roughly involves (i) the dispersal of ancestral (eastern) types through Oligocene terranes in the Western Mediterranean (ii) the Tell Atlas orogenesis as gene flow barrier between future west and east populations, (iii) the impact of recurrent climatic fluctuations from mid-Pliocene to the last ice age, (iv) the loss of the eastern lineage during Pleistocene northwards expansion phases. Background processes have especially become a central challenge in The aim of phylogeography is to explain the spatial dis- phylogeography and the recent development of analyti- tribution of genetic lineages within species and highlight cal tools based on coalescent theory is very helpful for the most influential historical factors explaining their investigating the demographic history of populations. distribution [1]. Hypotheses commonly advanced in phy- From information provided by phylogenetic trees and logeography are inherent to dispersal and vicariant the signature in the pattern of DNA substitutions left in events, inferences about which being usually based on the case of historical change in population size, it is pos- estimates of genetic diversity, divergence time and sibletoidentifythedemographicprocesswhichhas demographic history [2]. Inferences on demographic occurred and estimate related parameters [3,4] The wealth of literature dealing with phylogeography and * Correspondence: [email protected] demography in many invertebrate and vertebrate species 1 Laboratoire de Parasitologie Pharmaceutique (CNRS UMR 6553), Faculté des emphasizes the importance of combining both lineage Sciences Pharmaceutiques et Biologiques, 35043 Rennes, France © 2010 Guiller and Madec; 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. Guiller and Madec BMC Evolutionary Biology 2010, 10:18 Page 2 of 20 http://www.biomedcentral.com/1471-2148/10/18 and population components to reveal biogeographical groups of populations (West vs East) between which the histories. Moreover, the comparison of phylogeographic separation occurs in Kabylia (Algeria) is invariably structures across co-distributed taxa allows to find con- observed throughout the north African coastal region; cordant partitions and to infer common historical fac- (ii) almost all European populations clustered with wes- tors of divergence and colonization. Then, in North tern north African ones, with smaller genetic distances America (for review see [5]) as in Europe [6], various than those between western and eastern north Africa species have been studied to determine the climatic and (Fig 2), [9,10]. geological effects on phylogeographic patterns and Two competing biogeographical scenarios based on population structures. More precisely in Europe, the opposite directions of dispersal were considered to Mediterranean basin characterized by an exceptional explain the spatial pattern of variation in C. aspersum level of biodiversity and a complex geological history and especially to determine processes responsible for [7], has been the subject of intensive studies carrying genetic isolation in the eastern part of north Africa (Fig out on genetic diversity and phylogeography. Neverthe- 2) [10]. Both were based on Pliocene geological events less, most of these studies provide insights into the evo- and Quaternary cold periods, but colonization routes lutionary history of southern European species. Despite taken by the species involved migrations either from its key location between the rest of the continent and north Africa to Europe or from western Europe to north Europe, only a few cases concern the north African ter- Africa, with likely secondary contacts in the Kabylian ritory. In addition, investigations including Algerian data area. The doubtful direction of species expansion results sets are relatively rare, and main historical process influ- from discrepancies of spatial structure between allozyme encing the differentiation is generally found in Quatern- and partial mitochondrial data, and from an inconstant ary climatic oscillations, meaning that few studies imply pattern of gene diversity. The previous scenario we pro- events which occurred over times much longer than the posed [9,14], implying a northwards progression of the last glacial period. species (model 1, Fig 2), fits well with published reports Amongst the few species widely sampled in north that place the origin of C. aspersum in north Africa [8]. Africa (i.e. throughout the northern territory from wes- However, mitochondrial gene diversity estimated in tern Morocco to eastern Tunisia) for studying historical north Africa does not corroborate the northward coloni- processes influencing their current distribution, the land zation hypothesis [15]. The value of nucleotide diversity, mollusc Cornu aspersum (= syn. Helix aspersa) provides twice as weak in the east (clade C) as in the west (clade an excellent biological model to understand phylogeo- B) north African mtDNA lineages, together with unre- graphic patterns across north Africa and surrounding solved relationships between eastern populations, is regions of the Western Mediterreanean basin, and eval- rather consistent with southward dispersal either via uate hypotheses leading to population differentiation. Tyrrhenian/Aegean routes or the Strait of Gibraltar This land snail, formerly known as Helix aspersa Müller, (model 2, Fig 2) [15]. Whilst in the first scenario we 1774, is originated from Mediterranean countries. It ascribed the lower diversity in east to the occurrence of comprises a set of north African endemic forms and transitory and prolonged bottlenecks inherent in succes- subspecies that were described at the beginning of the sive migrations that populations experienced from Kaby- 20th century on the basis of shell characteristics [8]. The lia to Tunisia, this may also simply indicate a more most common one, C. a. aspersum (syn. H.
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