Aim: Phylogeographic studies on a number of Western Palaearctic taxa imply that the southern Caspian Sea region served as a refugium during the Last Glacial Maximum (LGM). Nevertheless, the LGM biogeography of the terrestrial and freshwater organisms in this region is poorly understood. By studying phylogeography of a species of freshwater crab, we unravel the location of probable local refugia and colonization patterns across this topographically complex landscape.Location: Southern Caspian Sea region.Taxon: The freshwater crab Potamon ibericum. Methods:We inferred the demographic history of Potamon ibericum using 163 mtDNA Cox1 sequences from 16 localities. For a subset of specimens, we amplified mtDNA 16S and four nuclear markers (28S, Histone3, Enolase, NaK) to confirm the phylogeographic structure. We tested Pleistocene expansion-contraction dynamics, using species distribution modelling (SDM) and assessed morphological variability by geometric-morphometric methods. Results: Mitochondrial markers revealed the existence of three parapatrically distributed lineages in the western, central and eastern parts of the region. The more conserved nuclear markers did not reflect this. The SDM revealed the fundamental niche expansion of P. ibericum along the southern Caspian Sea during the LGM, agreeing with results from the Cox1 data of demographic expansions. Despite strong morphological resemblance, geometric-morphometrics elucidated clinal carapace shape variation. Main Conclusions: Considering the Caspian Sea level fluctuations during the late Pleistocene, we hypothesize a scenario of demographic contraction during interglacial/warmer conditions, due to the Caspian Sea transgression and latitudinal range shift within the narrow distribution of the species between the southern Caspian Sea and northern slopes of the Alborz Mountains. During the glacial periods, the sea regressed and moist and warm conditions, characterizing glacial refugia, became prevalent in the region and caused isolated populations to expand. Our study confirms the deep transverse phylogeographic break, previously identified for Darevskia lizards in the region, and adds support to the presence of several LGM refugia in the southern Caspian Sea region.

BackgroundRecently, population genetic studies of Mediterranean marine species highlighted patterns of genetic divergence and phylogeographic breaks, due to the interplay between impacts of Pleistocene climate shifts and contemporary hydrographical barriers. These factors markedly shaped the distribution of marine organisms and their genetic makeup. The present study is part of an ongoing effort to understand the phylogeography and evolutionary history of the highly dispersive Mediterranean green crab, Carcinus aestuarii (Nardo, 1847), across the Mediterranean Sea. Recently, marked divergence between two highly separated haplogroups (genetic types I and II) of C. aestuarii was discerned across the Siculo-Tunisian Strait, suggesting an Early Pleistocene vicariant event. In order to better identify phylogeographic patterns in this species, a total of 263 individuals from 22 Mediterranean locations were analysed by comparing a 587 basepair region of the mitochondrial gene Cox1 (cytochrome oxidase subunit 1). The examined dataset is composed of both newly generated sequences (76) and previously investigated ones (187).ResultsOur results unveiled the occurrence of a highly divergent haplogroup (genetic type III) in the most north-eastern part of the Mediterranean Sea. Divergence between the most distinct type III and the common ancestor of both types I and II corresponds to the Early Pleistocene and coincides with the historical episode of separation between types I and II. Our results also revealed strong genetic divergence among adjacent regions (separating the Aegean and Marmara seas from the remaining distribution zone) and confirmed a sharp phylogeographic break across the Eastern Mediterranean. The recorded parapatric genetic divergence, with the potential existence of a contact zone between both groups in the Ionian Sea and notable differences in the demographic history, suggest the likely impact of paleoclimatic events, as well as past and contemporary oceanographic processes, in shaping genetic variability of this species.ConclusionsOur findings not only provide further evidence for the complex evolutionary history of the green crab in the Mediterranean Sea, but also stress the importance of investigating peripheral areas in the species’ distribution zone in order to fully understand the distribution of genetic diversity and unravel hidden genetic units and local patterns of endemism.Electronic supplementary materialThe online version of this article (10.1186/s12862-018-1167-4) contains supplementary material, which is available to authorized users.

Refugia are critical for the maintenance of biodiversity during the periods of Quaternary climatic oscillations. The long‐term persistence of refugial populations in a large continuous refugium has resulted in a homogenous pattern of genetic structure among populations, while highly structured evolutionary lineages characterize the restriction of refugial populations to smaller subrefugia. These mechanisms have resulted in the identification of hot spots of biodiversity within putative glacial refugia. We studied phylogeography of Potamon ibericum (Brachyura: Potamidae) in the drainages of the western Caucasus biodiversity hot spot (i.e., Colchis and the Caucasus) to infer spatial genetic structure and potential refugia for a freshwater crab in this region. These areas have traditionally considered as a refugium due to the presence of Tertiary relict species. We integrated population genetic data and historical demographic analysis from cytochrome oxidase subunit I sequences and paleoclimatic data from species distribution modeling (SDM). The results revealed the lack of phylogeographic structure and provided evidence for demographic expansion. The SDM presented a rather homogenous and large refugium that extended from northeast Turkey to Colchis during the last glacial period. In contrast to these findings, previous phylogeographic study on P. ibericum of the eastern Caucasus biodiversity hot spot (i.e., Hyrcania) identified multiple independent refugia. By combining these results, we explain the significance of this important western Palearctic hot spot of biological diversity in shaping the geographic distribution of intraspecific genetic diversity in a freshwater taxon.

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In the current study, we redescribe and revalidate a freshwater crab species of the genus Potamon Savigny, 1816, from northern Iran. Potamon elbursi Pretzmann, 1962, differs from the species it has been synonymised with, Potamon persicum Pretzmann, 1962, mainly by the shape of the first gonopods in males. Consistent and marked genetic divergence was also recognised in the mitochondrial 16S rRNA and cytochrome oxidase subunit I genes. This study elevates P. elbursi to species rank and therefore increases the number of valid species of the genus Potamon to a total of twenty-two.

Hydrographically

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Many species and subspecies of Potamon have been described from the easternmost distribution of the genus in the western tributaries of the Indus River. Most of them were synonymised subsequently under the two names of currently valid species known from the region: Potamon gedrosianum Alcock, 1909 and Potamon ruttneri Pretzmann, 1962. Genetic and morphological information, based on mitochondrial 16S rDNA and the first male gonopod (G1), were gathered in the course of the present study. The corresponding results suggest the occurrence of four groups and question the taxonomic status of both species. We also revise the distribution range of both species, in particular that of P. gedrosianum, with a new record from Iran. Overall, the study reveals the need for a major revision using further morphological and molecular data. Because of the complexity of this necessary revision and the incomplete sampling, we here refrain from proposing any taxonomic conclusions.