Molecular Phylogenetics and Evolution 62 (2012) 681–692
Total Page：16
File Type：pdf, Size：1020Kb
Molecular Phylogenetics and Evolution 62 (2012) 681–692 Contents lists available at SciVerse ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev Mitochondrial phylogeny and biogeographic history of the Greek endemic land-snail genus Codringtonia Kobelt 1898 (Gastropoda, Pulmonata, Helicidae) ⇑ Panayiota Kotsakiozi a, Aristeidis Parmakelis b, , Sinos Giokas c, Irene Papanikolaou b, Efstratios D. Valakos a a Department of Animal and Human Physiology, Faculty of Biology, University of Athens, Panepistimioupoli Zografou, GR-15784 Athens, Greece b Department of Ecology and Taxonomy, Faculty of Biology, University of Athens, Panepistimioupoli Zografou, GR-15784 Athens, Greece c Section of Animal Biology, Department of Biology, University of Patras, GR-26500 Patras, Greece article info abstract Article history: The aim of this work was to infer the phylogeny of the Greek endemic land-snail genus Codringtonia Received 22 March 2011 Kobelt 1898, estimate the time frame of the radiation of the genus, and propose a biogeographic scenario Revised 8 November 2011 that could explain the contemporary distribution of Codringtonia lineages. The study took place in the dis- Accepted 15 November 2011 tricts of Peloponnese, Central Greece and Epirus of mainland Greece. Sequence data originating from Available online 28 November 2011 three mtDNA genes (COI, COII, and 16S rDNA) were used to infer the phylogeny of the eight nominal Cod- ringtonia species. Furthermore, the radiation time-frame of extant Codringtonia species was estimated Keywords: using a relaxed molecular clock analysis and mtDNA substitution rates of land snails. The phylogenetic Dispersal analysis supported the existence of six Codringtonia lineages in Greece and indicated that one nominal Greece Land-snails species (Codringtonia neocrassa) might belong to a separate genus distantly related to Codringtonia. The Phylogeny time frame of differentiation of Codringtonia species was placed in the Late Miocene–Pleistocene epoch. Vicariance The dispersal–vicariance analysis performed indicated that most probably Codringtonia exhibited a north-to-south spread with the ancestral area being that of central Greek mainland, accompanied with duplication (speciation) and vicariance events. Ó 2011 Elsevier Inc. All rights reserved. 1. Introduction of the Greek region’’. However, the vast majority of these studies are focused on the insular part of Greece and more specifically in The complex late Tertiary geological history of the Balkan the Aegean Archipelago, whereas comprehensive studies surveying Peninsula and especially of the Greek area (Creutzburg, 1963; taxa of continental Greece are considerably underrepresented (de Dermitzakis, 1990; Meulenkamp, 1985; Steininger and Roegl, Weerd and Gittenberger, 2005; de Weerd et al., 2006, 2004, 1984), located at the margin of the Eurasian and African plates, 2005; Gittenberger et al., 2004; Klossa-Kilia et al., 2006; Parmakelis have contributed to the diversification and distribution of many et al., 2008; Poulakakis and Sfenthourakis, 2008; Sotiropoulos terrestrial animals (Beerli et al., 1996; Bittkau and Comes, 2005; et al., 2007; Thanou et al., 2005; Tryfonopoulos et al., 2008). Never- Papadopoulou et al., 2009; Parmakelis et al., 2005, 2006a,b; theless, the distributional patterns of genetic diversity of certain Poulakakis et al., 2003, 2005a,b). Multiple land connections offered taxa that have been studied from mainland Greece highlight that the opportunities for dispersal in some cases, whereas in others the processes shaping the biodiversity of mainland Greece are quite submergence of land bridges disrupted the distributions of related complex, regionally specific and divergent (Sotiropoulos et al., taxa. In biological terms these geological events resulted in high 2007). This is partly due to the palaeogeographic history of levels of diversity and endemism in the entire mainland and insular mainland Greece that, at a regional level, was even more complex Greece (Medail and Diadema, 2009; Medail and Quezel, 1999; than that of the insular area [see Parmakelis et al. (2006a)]. Addi- Parmakelis et al., 2005; Sfenthourakis and Legakis, 2001). Several tionally, the topographic complexity of mainland Greece is much studies have employed molecular markers to survey the impor- higher compared to that of the islands (with the possible exception tance of vicariance events and dispersal in shaping the distribution of Crete), and regionally fluctuating. Consequently, the regional of genetic diversity for the terrestrial fauna of the Greek territory. character of the processes shaping biodiversity in the mainland is As stated in Triantis and Mylonas (2009) ‘‘over the last 25 years, well justified, and therefore it is of major interest to determine numerous ecological, biogeographical, and evolutionary studies the taxa and region specific biogeographic patterns in order to be have greatly enhanced our knowledge of the biodiversity patterns able to distinguish between idiosyncratic and general evolutionary processes. Here we investigate the evolutionary history of the land-snail ⇑ Corresponding author. Fax: +30 2107274885. genus Codringtonia Kobelt 1898, which is endemic to Greece E-mail address: [email protected] (A. Parmakelis). 1055-7903/$ - see front matter Ó 2011 Elsevier Inc. All rights reserved. doi:10.1016/j.ympev.2011.11.012 682 P. Kotsakiozi et al. / Molecular Phylogenetics and Evolution 62 (2012) 681–692 (Subai, 2005) and has a restricted and mosaic distribution (Fig. 1) the genus. Codringtonia species are found at various altitudes, in the continental area. Characters of the shell and of the distal living in crevices on rocky terrain within maquis and coniferous genitalia anatomy are used for species identification (Subai, (except pines) or mixed (deciduous–coniferous) forests (Giokas 2005). However, the actual number of Codringtonia species distrib- et al., 2007). Due to their very limited distributional range, all spe- uted in Greece is not clear and a resolved phylogeny is missing. cies occurring in Greece (except C. neocrassa) have been registered Subai (2005) concluded that Codringtonia is represented by eight in the Red Data Book of threatened animals of Greece as vulnerable species in Greece, whereas prior to Subai’ s review, the number (Legakis and Maragou, 2009). of Codringtonia species and subspecies reported from Greece was Therefore, our main goals are (1) to infer the evolutionary rela- 12 (Hadjicharalambous, 1996). Four (Codringtonia codringtonii, tionships of the Codringtonia species of Greece, using sequence Codringtonia elisabethae, Codringtonia gittenbergeri and Codringtonia data originating from three mtDNA genes, (2) to estimate the radi- helenae) out of the eight currently recognized Codringtonia species ation time-frame of the Codringtonia species, applying a relaxed (Subai, 2005) can be found only in particular areas of the Pelopon- molecular clock strategy and using mtDNA substitution rates of nese peninsula; a single species (Codringtonia neocrassa) is found in land snails, and (3) to propose, within this phylogenetic frame- some areas of the district of Epirus (northwestern Greece) and in work, a biogeographic scenario accounting for the current Codring- the nearby Albanian territory and another one (Codringtonia tonia lineages’ distribution. parnassia) is found exclusively in Central Greece. Finally, there are two species (Codringtonia eucineta and Codringtonia intuspli- 2. Materials and methods cata) that are present both in Central Greece and Peloponnese pen- insula. Between the district of Epirus and Central Greece no 2.1. Specimens, DNA extraction, amplification and sequence Codringtonia species occur (Subai, 2005). Regarding its insular determination distribution a single species (C. neocrassa) is present only on the island of Kerkira (Corfu) (Fig. 1). The discontinuous and patchy Total genomic DNA was isolated from foot muscle of specimens distribution of the genus is puzzling itself. This is claimed since that were either frozen (À20 °C) or (absolute) ethanol preserved. Codringtonia species are missing from potentially suitable regions To overcome problems of polymerase chain reaction (PCR) inhibi- that are located in between the present day distribution areas of tion by mucopolysaccharides, DNA was extracted using the CTAB BULGARIA FYROM ALBANIA 20 Aegean Sea GREECE 23 24 Epirus 21 22 TURKEY 17 9 18 15 7 19 13 14 6 16 4 5 Peloponnese Central Greece 3 10 11 1 2 8 12 C. codringtonii C. gittenbergeri C. elisabethae C. intusplicata C. eucineta C. parnassia C. helenae C. neocrassa N 100km CRETE Fig. 1. Distribution map of Codringtonia species (redrawn from Subai, 2005) and sampling localities of Codringtonia populations included in the study. Numbers of localities correspond to the map codes reported in Table 1. P. Kotsakiozi et al. / Molecular Phylogenetics and Evolution 62 (2012) 681–692 683 Table 1 Species and populations included in the study, geographic data of the sampling localities, sample sizes (N) of populations and accession numbers of sequences. Map codes refer to Figure 2. Map Sampling District Latitude Longitude Collection Nominal Sample code Accession code Locality (N) Date species numbers COI/ COII/16S 1 Chrysokelaria Peloponnese 36.79154 21.88558 30/11/ C. CcHr1 JQ239920/ Monastery (1) 2008 codringtonii JQ240005/ JQ240089 2 Mathias (1) Peloponnese 36.92928 21.88341 30/11/ C. CcMath1 JQ239921/ 2008 codringtonii JQ240006/ JQ240090 3 Navarino (2) Peloponnese