[Show abstract][Hide abstract] ABSTRACT: Eighteen protein loci have been electrophoretically analyzed in one population of Podarcis hispanica lizards from the Columbrets Islands and in four others of P. lilfordi from islets close to the Menorca coast. These populations and almost all of those previously studied from Mallorca showed much higher heterozygosities than the average for reptiles. This fact could be tentatively explained by the large population size effects and secondly by possible migration events on the genetic structure of a selectively neutral pool of allozymic alleles. The genetic distances allowed a sharp discrimination between P. hispanica and both Balearic Podarcis, but not between the latter two. Thus, P. pityusensis should be synonymized with P. lilfordi but having a possible taxonomic status of subspecies in agreement with the hybridization results got by other authors. Some possible correspomdances between genetic distances and geographical parameters are also discussed.
[Show abstract][Hide abstract] ABSTRACT: The association between polymorphism at the mc1r locus and colour variation was studied in two wall lizard species (Podarcis lilfordi and P. pityusensis) from the Balearic archipelago. Podarcis lilfordi comprises several deep mitochondrial lineages, the oldest of which originated in the Pliocene, while much shallower mitochondrial lineages are found in P. pityusensis. Here, we examined whether specific substitutions were associated with the melanic colouration found in islet populations of these species. Homologous nuclear sequences covering most of the mc1r gene were obtained from 73 individuals from melanic and non-melanic Podarcis from different populations (the entire gene was also sequenced in six selected individuals). MtDNA gene trees were also constructed and used as a framework to assess mc1r diversity. Mc1r showed greater polymorphism in P. lilfordi than in P. pityusensis. However, we observed no substitutions that were common to all melanic individuals across the two species. Only one significant association was detected in the mc1r partial sequence, but this was a synonymous A/G mutation with A alleles being more abundant in melanic populations. In addition, there were no associations between the main dominant phenotypes (green and brown, blue and yellow spots and ventral colour) and synonymous or non-synonymous substitutions in the mc1r gene. There was no statistical evidence of selection on mc1r. This study suggests no relationship between mc1r polymorphism and colour variation in Balearic Podarcis.
PLoS ONE 09/2013; 8(1):e53088. DOI:10.1371/journal.pone.0053088 · 3.23 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Two monophyletic sister species of wall lizards inhabit the two main groups of Balearic Islands: Podarcis lilfordi from islets and small islands around Mallorca and Menorca and Podarcis pityusensis from Ibiza, Formentera and associated islets. Genetic diversity within the endangered P. lilfordi has been well characterized, but P. pityusensis has not been studied in depth. Here, 2430 bp of mtDNA and 15 microsatellite loci were analysed from P. pityusensis populations from across its natural range. Two main genetic groupings were identified, although geographical structuring differed slightly between the mtDNA and the nuclear loci. In general, individuals from islets/islands adjacent to the main island of Ibiza were genetically distinct from those from Formentera and the associated Freus islands for both mtDNA and the nuclear loci. However, most individuals from the island of Ibiza were grouped with neighbouring islets/islands for nuclear loci, but with Formentera and Freus islands for the mitochondrial locus. A time-calibrated Bayesian tree was constructed for the principal mitochondrial lineages within the Balearics, using the multispecies coalescent model, and provided statistical support for divergence of the two main P. pityusensis lineages 0.111-0.295 Ma. This suggests a mid-late Pleistocene intraspecific divergence, compared with an early Pleistocene divergence in P. lilfordi, and postdates some major increases in sea level between 0.4 and 0.6 Ma, which may have flooded Formentera. The program IMa2 provided a posterior divergence time of 0.089-0.221 Ma, which was similar to the multispecies coalescent tree estimate. More significantly, it indicated low but asymmetric effective gene copy migration rates, with higher migration from Formentera to Ibiza populations. Our findings suggest that much of the present-day diversity may have originated from a late Pleistocene colonization of one island group from the other, followed by allopatric divergence of these populations. Subsequent gene flow between these insular groups seems likely to be explained by recent human introductions. Two evolutionary significant units can be defined for P. pityusensis but these units would need to exclude the populations that have been the subjects of recent admixture.
[Show abstract][Hide abstract] ABSTRACT: Podarcis filfolensis is an endemic lizard from the Maltese archipelago. There is evidence of human-mediated decline and even extirpation of some insular populations of this species. However, information about the intraspecific genetic diversity and phylogeographic patterns of this species is limited. Here we analyze genetic markers from a multi-locus dataset (mtDNA, 2,533 bp; nuclear c-mos gene, 353 bp; 11 microsatellites) for individuals from extant populations of P. filfolensis. Despite generally low genetic variability, two main mitochondrial groupings were clearly identified. In general, individuals from the main island of Malta were genetically distinct from those from Gozo, Comino, Cominotto and Small Blue Lagoon Rock, and also from Linosa and Lampione individuals. Three genetic clusters were detected based on microsatellite data: one was found at higher frequency on Malta, while the other two included samples from the remaining islands, showing some concordance with the mtDNA pattern. A time-calibrated Bayesian tree for the principal mitochondrial lineages indicated strong statistical support for two P. filfolensis lineages that originated in the Pleistocene (105.4–869 Ka). We show that these lineages largely meet the criteria for recognition as evolutionary significant units despite some recent admixture (possibly due to recent translocations between islands). Human disturbance, low genetic variability, evidence of bottlenecks and extirpation on one island indicate that a thorough review of the current conservation status of P. filfolensis would be timely.
[Show abstract][Hide abstract] ABSTRACT: Fifteen microsatellite loci are described for the lizard Podarcis lilfordi from the Balearic Islands. Loci were isolated from partial genomic libraries that had been enriched for AAAG and ACAG repeat
sequence. All loci were highly polymorphic (six to 19 alleles) in a sample of 26 individuals from the island of Dragonera.
Observed and expected heterozygosities ranged from 0.46 to 0.92 and 0.78 to 0.95, respectively. These loci will be used to
study the evolutionary history of different forms, the occurrence of cross-islet introductions, and the validity of subspecific
designations in this species.
[Show abstract][Hide abstract] ABSTRACT: Podarcis lilfordi is an endemic lizard found on islands and islets off the coast of Mallorca and Menorca (including the Cabrera archipelago) in the Balearic Islands, which is extinct on the two main islands themselves. Analyses of mtDNA revealed four unconnected parsimony networks. These represented (I) all Menorcan populations, (II) Dragonera, Malgrats and Toro islands (Western Mallorca), (III and IV) the remaining populations from Cabrera and Mallorca. We applied NCPA to evaluate its utility in understanding the genetic sub-structure. Within network I, contiguous range expansion was inferred for both main clades. Populations from western Mallorca (II) have undergone historical allopatric fragmentation events following isolation around the start of the Pleistocene. Clades within networks III and IV showed some allopatric fragmentation and restricted gene flow (isolation by distance) among islands from Cabrera, as well as long distance dispersal between the Cabrera archipelago and southern Mallorca. The NCPA strongly supports the importance of allopatric fragmentation, with some additional support for range expansion. The primary advantage of the NCPA approach was to provide statistical support for these inferences.
[Show abstract][Hide abstract] ABSTRACT: Aim To describe and analyse phylogeographical patterns in the endangered endemic lizard Podarcis lilfordi from across its remaining range and thereby establish baseline information on genetic diversity that will help determine conservation priorities and assist future reintroduction programs.
Location Balearic Islands, Spain.
Methods We analysed mitochondrial DNA (2382 bp sequence from eight genes) from 118 individuals and characterized the relationships among haplotypes using parsimony networks, as well as phylogenetic inference. Analyses of historical gene flow and population growth were used to provide further insights into population histories.
Results Four unconnected parsimony networks were obtained that mirrored the main clades in the phylogenetic tree: (I) all Menorcan populations, (II) Dragonera, Malgrats and Toro islands (Western Mallorca) (III and IV) and the remaining populations from Cabrera and Mallorca. Two major haplotype groups were detected in Menorca (I) and these provided signatures of a demographic expansion and asymmetrical historical gene flow, respectively, concordant with the expected direction of colonization from south to north of the island. Populations from western Mallorca (II) showed evidence of historical allopatric fragmentation events following isolation around the start of the Pleistocene. In networks III and IV, Cabreran populations appear to have become isolated from north and south Mallorca quite recently, with asymmetric gene flow indicating a northwards dispersal direction.
Main conclusions P. lilfordi is a genetically diverse species that shows substantial mtDNA structuring both between regions and, at a finer scale, between some islet populations within regions. The precarious state of some islet populations shown here to be quite divergent (e.g. Toro island in western Mallorca) means that conservation of this intraspecific biodiversity requires urgent action.
[Show abstract][Hide abstract] ABSTRACT: Phylogenetic relationships and timings of major cladogenesis events are investigated in the Balearic Island lizards Podarcislilfordi and P.pityusensis using 2675bp of mitochondrial and nuclear DNA sequences. Partitioned Bayesian and Maximum Parsimony analyses provided a well-resolved phylogeny with high node-support values. Bayesian MCMC estimation of node dates was investigated by comparing means of posterior distributions from different subsets of the sequence against the most robust analysis which used multiple partitions and allowed for rate heterogeneity among branches under a rate-drift model. Evolutionary rates were systematically underestimated and thus divergence times overestimated when sequences containing lower numbers of variable sites were used (based on ingroup node constraints). The following analyses allowed the best recovery of node times under the constant-rate (i.e., perfect clock) model: (i) all cytochrome b sequence (partitioned by codon position), (ii) cytochrome b (codon position 3 alone), (iii) NADH dehydrogenase (subunits 1 and 2; partitioned by codon position), (iv) cytochrome b and NADH dehydrogenase sequence together (six gene-codon partitions), (v) all unpartitioned sequence, (vi) a full multipartition analysis (nine partitions). Of these, only (iv) and (vi) performed well under the rate-drift model. These findings have significant implications for dating of recent divergence times in other taxa. The earliest P.lilfordi cladogenesis event (divergence of Menorcan populations), occurred before the end of the Pliocene, some 2.6Ma. Subsequent events led to a West Mallorcan lineage (2.0Ma ago), followed 1.2Ma ago by divergence of populations from the southern part of the Cabrera archipelago from a widely-distributed group from north Cabrera, northern and southern Mallorcan islets. Divergence within P.pityusensis is more recent with the main Ibiza and Formentera clades sharing a common ancestor at about 1.0Ma ago. Climatic and sea level changes are likely to have initiated cladogenesis, with lineages making secondary contact during periodic landbridge formation. This oscillating cross-archipelago pattern in which ancient divergence is followed by repeated contact resembles that seen between East-West refugia populations from mainland Europe.
[Show abstract][Hide abstract] ABSTRACT: Sequences of the cytochrome b gene were analysed for 47 samples of two species of Podarcis from the Balearic Islands (P. pityusensis from Pityuses and P. lilfordi, from Gymnesies archipelago). The average uncorrected distance between the two species studied was 9.7%. The sampled individuals from each species form reciprocally monophyletic units. Assuming an overall rate of change for cytochrome b of 2% per million years the nucleotide divergence of 9.7% ± 1.9% between P. lilfordi and P. pityusensis, could correspond to a divergence time of 4.95 ± 0.95 million years. The separation time between Gymnesies and Pityuses archipelagos according to geological hypotheses is around 5 million years, thus our results suggest the ancestors of the two present forms became isolated during this event. The most parsimonious networks suggest that currently accepted subspecies do not form monophyletic groups, and so should be reassessed. Despite our limited sampling the level of variability is much higher in P. lilfordi than P. pityusensis. Since they are sister taxa the relative age of each species is equal, therefore the differences might be due to historical population structure differences. However it seems that the forms on the islands are not genetically distinct units, implying that morphological differences are recent adaptations to their environments.
[Show abstract][Hide abstract] ABSTRACT: Eighteen protein loci have been electrophoretically analyzed in one population of Podarcis hispanica lizards from the Columbrets Islands and in four others of P. lilfordi from the islets close to the Menorca coast. These populations and almost all of those previously studied from Mallorca showed much higher mean heterozygosities than the average for reptiles. This fact could be tentatively explained firstly by possible migration events on the genetic structure of a selectively neutral pool of allozymic alleles. The genetic distances allowed a sharp discrimination between P. hispanica and both Balearic Podarcis, but not the latter two. Thus P. pityusensis should be synonymized with P. lilfordi but having a possible taxonomic status of subspecies in agreement with the hybridization results got by other authors. Some possible correspondences between genetic distances and geographical parameters are also discussed.