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[Aim]: Discerning the relative role of geographical and ecological factors in promoting diversification is central to our understanding of the origin and maintenance of biodi-versity. We explore the roles of geology and ecological tolerance in the diversification of a group of wingless beetles with low dispersal potential. [Location]: Western Mediterranean (Iberian Peninsula and North Africa)., [Taxon]: Darkling beetles (Tenebrionidae: Misolampus)., [Methods]: We sequenced nine gene fragments from the mitochondrial and nuclear genomes in all extant Misolampus species to reconstruct their phylogeny, evaluate species boundaries and potential contact zones and estimate divergence times. We modelled species distributions for different time periods to infer ecological preferences and assess the effects of climatic changes since the last interglacial. We used a time-stratified process-based biogeographical model to estimate ancestral areas of origin and the evolution of geographical ranges., [Results]: The palaeoclimatic model projections show contractions of favourable areas during the last interglacial period and mid-Holocene, and wide stretches of suitable areas during the last glacial maximum. Analyses of ancestral bioclimatic preferences reveal ecological adaptations in isolated lineages within three species. The phylog-eny of Misolampus is strongly supported and unveils deep divergences within the six species. Two well-supported clades were recovered, one distributed in North Africa- Balearic Islands and another in the Iberian Peninsula. The divergence between the North African and Iberian clades occurred during the early Miocene. The geographical ranges of species are dynamic, shaped by biotic (e.g. life history, demography, species interactions) and abiotic (e.g. tectonic events, climatic variables) factors that vary through space and time. This variation ultimately dictates differential patterns of population viability, connectivity and growth through time in dif-ferent sections of species ranges, with important consequences for species diversification processes (Bellard et al., 2012; Gouveia et al., 2014; Thuiller et al., 2005; Wiens, 2011). Following climatic changes, species can respond by contracting their ranges into cli-matic refugia and/or dispersing to areas where their environmen-tal preferences are maintained (niche tracking), by adapting to new environmental conditions in situ (niche evolution), and often by undergoing extinction in environmentally unsuitable areas (Kozak & Wiens, 2006; Yesson & Culham, 2006a). The fragmentation of ancestral distributions because of tectonic or climatic changes pro-motes allopatric speciation, sometimes associated with niche evolution, whereas demographic expansion of populations tracking climatic changes can lead to range fusion and admixture between formerly isolated population groups. Discerning the relative role of biotic and abiotic factors in promoting diversification is central to our understanding of the origin and maintenance of biodiversity (Wiens & Graham, 2005).The ability of different taxa to track climatic conditions or evolve new adaptations depends on life-history traits like dispersal. Dispersal limitation can result in geographical isolation and ecolog-ical specialization, by promoting species persistence in small areas, which could lead to some resilience to changing climatic conditions and might accelerate the consolidation of a species' ecological niche even at the population level (Baselga et al., 2011). Usually, local adaptation to extrinsic ecological factors is initiated by the presence of a barrier to gene flow. Additionally, dispersal limitation may lead to ecological specialization and reinforce restrictions to gene flow, promoting speciation even in the face of future demographic pro-cesses such as population contractions, fragmentations or expan-sions (Wiens, 2004).Addressing the question of how much lineage diversification is affected by range expansions/contractions through historical biogeography is often tackled by reconstructing ancestral range shifts using phylogenetic analyses (Ree & Smith, 2008 ; Ronquist & Sanmartín,2011). However, the question often remains partially an-swered because of the lack of information on the timing of range shifts regarding the speciation process itself (Recuero et al., 2012; Recuero & García-París, 2011). Robust, accurate estimates of di-vergence times between species and intraspecific lineages are key to consider evolutionary processes at different temporal scales that lead to diversification, which is controlled by multiple extrin-sic ecological factors and intrinsic organismal traits (Wiens, 2004). Therefore, assessing the temporal scale of demographic processes such as range contractions, fragmentations and expansions and their impact on lineage divergence provides decisive evidence to reconstruct the evolutionary history of species. Within a temperate region such as the Western Mediterranean, the influence of demographic processes under climatic oscillations has been well studied, providing evidence for two major evolutionary patterns. On the one hand, allopatric fragmentation, or demographic contraction into refugial areas during glacial periods have allowed species to persist as isolated populations over long peri-ods of time (Gutiérrez-Rodríguez et al., 2017 ; Hewitt, 2004, 2011; Martínez- Freiría et al.,2020; Martínez-Solano et al., 2006; Recuero & García-París, 2011). On the other hand, during interglacial periods many species have expanded their ranges through newly available colonization routes, sometimes forming secondary contact areas with related species or lineages (Branco et al., 2000; Gómez & Lunt, 2007 ; Gonçalves et al., 2009; Miraldo et al., 2011). These two scenarios involve periods without effective contact or gene flow be-tween isolated populations, which has consequences on the process of lineage divergence. For instance, intraspecific divergence could be either indicative of allopatric cladogenetic events (cladogenesis), or ecological specialization (niche differentiation) (Hewitt, 2011).The Western Mediterranean hunchback darkling beetles Misolampus (Tenebrionidae: Stenochiinae: Misolampinii) are well suited for studies on the contribution of geology-driven processes and ecological divergence to diversification in species groups with limited dispersal capacity. These flightless beetles are ecologically linked to woodlands and live mainly beneath the bark or inside dead logs of Pinus, Quercus and other tree species along a broad altitudinal analyses infer an ancestral range including the Iberian, Betic and Rifean Plates, with subsequent splits followed by dispersal events., [Main conclusions]: Our results favour a dual role of vicariance and dispersal in driving the historical biogeography and diversification of Misolampus since the early Miocene. We also found evidence for incipient speciation events, underscoring the role of tectonic events and adaptation to local climatic conditions in the diversification of the group., This study was supported by the Spanish government (MINECO/MIUC/AEI) and the European Fund for Regional Development (FEDER) under grants CGL2015-66571-P (collecting and Museum visits) and PID2019-110243GB-I00/AEI/10.13039/501100011033 (Ministerio de Ciencia, Innovación y Universidades, Spain) (molecular analyses) to MGP. Support of the publication fee was granted by the CSIC Open Access Publication Support Initiative through its Unit of Information Resources for Research (URICI).

Hunchback darkling beetles of the Ibero-Maghrebian genus Misolampus Latreille, 1807 (Tenebrionidae, Stenochiinae) encompass six species: M. gibbulus (Herbst 1799), M. goudotii Guérin-Méneville, 1834, M. lusitanicus Brême, 1842, M. ramburii Brême, 1842, M. scabricollis Graells, 1849, and M. subglaber Rosenhauer, 1856. Previously known distribution ranges of the species were delineated using many old records, the persistence of such populations being questionable under the current situation of global biodiversity loss. Additionally, the status of geographically isolated populations of the genus have been the subject of taxonomic controversy. An exhaustive bibliographical revision and field search was undertaken, and the Misolampus collection of the Museo Nacional de Ciencias Naturales (MNCN-CSIC) was revised. The aims are to (i) provide an updated geographic distribution range for the species of Misolampus; (ii) to determine the taxonomic status of controversial populations; (iii) to provide a catalogue for Misolampus; and (iv) to discuss the conservation status of these saproxylic beetles. As a result, a catalogue including synonymies and type localities, geographical records, diagnoses, and information on natural history for all species of Misolampus is presented. The results reveal that the distribution ranges of the species of Misolampus have not undergone a reduction in the last century, and indicate the presence of the genus in areas where it had never been recorded before. The morphological variability of M. goudotii drove the proposal of different taxa that are here formally synonymised as follows: M. goudotii Guérin-Méneville, 1834 = M. erichsoni Vauloger de Beaupré, 1900, syn. nov. = M. peyerimhoffi Antoine, 1926, syn. nov., PMP is supported by MICINN-FEDER through contract BES-2016-077777. This work was partially funded by the project grant PID2019-110243GB-100 (Ministerio de Ciencia e Innovación-FEDER) of Spain to MG-P., We also acknowledge institutional support from the Unit of Information Resources for Research at the Unit of Information Resources for Research at the "Consejo Superior de Investigaciones Científicas" (CSIC) for the article-processing charges contribution.

Este artículo contiene 29 páginas, 12 figuras, 1 tabla., The terrestrial isopod family Armadillidiidae presents higher diversity in karstic areas, with fewer species present in areas with reduced suitable subterranean habitats, such as siliceous sandy soils. Myrmecophily, although not widespread in the family Armadillidiidae, can help these animals to colonize sandy substrates, as is observed in several populations of myrmecophilous Armadillidiidae species in central and southern Spain. Morphological examination and multilocus phylogenetic analyses, including mitochondrial DNA (Cox1) and nuclear DNA (18S, 28S and H3) markers, indicate that these myrmecophilous populations represent four new taxa: Iberiarmadillidium pinicola gen. & sp. nov., Iberiarmadillidium psammophilum sp. nov., Iberiarmadillidium sakura sp. nov. and Cristarmadillidium myrmecophilum sp. nov. Some of the main diagnostic characters used in the taxonomy of Armadillidiidae are not clearly apomorphic. Among head morphologies, Eluma type seems to be the ancestral state, being typical of several unrelated lineages; duplocarinate and Armadillidium types are derived states observed in unrelated lineages. The presence of a schisma is a convergent character state, because it has been identified in several taxa nested in unrelated clades. The newly described taxa present patterns of morphological stasis and homoplasy, likely to be associated with their shared myrmecophilous habits. The generic taxonomy of the family needs a deep revision including phylogenetic approaches and thorough taxon sampling.

Delimiting species boundaries is a complex challenge usually hindered by overlooked morphological diversification or misinterpretation of geographically structured phenotypic variability. Independent molecular data are extremely useful to characterize and understand such morphological diversity. Morphological and molecular variability of the non-phoretic and apterous, widely distributed, giant blister beetles of the genus Berberomeloe, were investigated within and between lineages across most of the distributional range of the genus. We used two mtDNA gene fragments to characterize genetic variability and to produce a time-calibrated phylogeny of the genus. Our results reveal several mitochondrial lineages, allopatrically, parapatrically and sympatrically distributed. Most clades are not distinguishable between each other based on morphometrics. However, no morphometric overlap is observed between two closely related clades, one of them occurring in sympatry with a distantly congeneric species (B. insignis), suggesting that sympatry could trigger morphological diversification. Although most species share a morphometric space, they can be morphologically identified by a combination of easily observed characteristic qualitative features. Based on the concordance between mtDNA clades and morphological units, we describe six new species of Berberomeloe (B. castuo sp. nov., B. comunero sp. nov., B. indalo sp. nov, B. yebli sp. nov., B. payoyosp. nov. and B. tenebrosus sp. nov.), revalidate two taxa (B. maculifrons comb. nov. and B. laevigatus comb. nov.) and redefine B. majalis., The visits of MGP to the Magyar Természettudományi Múzeum of Budapest (HU-TAF) and to the Muséum National d’Histoire Naturelle of Paris (FR-TAF) were supported by the European Union SYNTHESYS Project (http://www.synthesys.info/) ‘European Community Research Infrastructure Action’ within package FP7 ‘Capacities’. This work has been funded by the Spanish Government project-grant CGL2015-66571-P (Ministerio de Economía y Empresa-Fondo Europeo de Desarrollo Regional) to MGP. The authors declare that there are no conflicts of interest.

El milpiés mexicano Euryurus flavocarinatus Daday, 1889, ha permanecido como incertae sedis por más de un siglo. El examen del holotipo demuestra que la especie pertenece al género Amplinus dentro de la amplia familia Aphelidesmidae, orden Polydesmida. En el presente trabajo presentamos una redescripción completa del holotipo de Amplinus flavocarinatus (Daday, 1889) comb. nov. Por otro lado, Euryurus flavocarinatus Silvestri, 1899 (actualmente cosiderada dentro del género Pycnotropis) es un homónimo primario más reciente de E. flavocarinatus Daday, 1889, por lo que proponemos el nombre Pycnotropis silvestrii Recuero & Aguado- Aranda nom. nov. para sustituir el nombre de Silvestri y resolver la homonimia.

Flightless, phoretic insects of the tribe Meloini include several widespread species able to inhabit both mainland and oceanic and continental islands. Relationships between phoretic and non-phoretic species of Meloini are unclear, precluding broad evolutionary and biogeographic analyses within the group. We provide a robust molecular phylogenetic framework for Meloini, with finer sampling in the Western Palaearctic, by analysing two nuclear (Wg, 18S) and two mtDNA (cox1, 16S) gene fragments of 24 species representing eight subgenera of Meloe and Physomeloe. Our phylogenetic analyses revealed that Meloe, as currently recognized, is paraphyletic. Our phylogenetic hypothesis clarifies several controversial relationships within the group. We provide a new taxonomic framework that better reflects the evolutionary relationships of the group by raising to genus all previously considered subgenera: Afromeloe Schmidt, 1913, Alveomeloe Pripisnova, 1987, Chiromeloe Reitter, 1911, Desertimeloe Kaszab, 1964, Eurymeloe Reitter, 1911, Lampromeloe Reitter, 1911, Lasiomeloe Reitter, 1911, Listromeloe Reitter, 1911, Meloegonius Reitter, 1911, Mesomeloe Reitter, 1911, Meloenellus Reitter, 1911, Micromeloe Reitter, 1911, Taphromeloe Reitter, 1911 and Treiodous Dugès, 1869; while Anchomeloe Iablokoff-Khnzorian, 1983, is recovered as subgenus of Meloe. In addition, we conducted an extensive intraspecific sampling for four phoretic taxa (Meloe proscarabaeus, Meloe autumnalis, Eurymeloe mediterraneus and Eurymeloe tuccia) and used mtDNA phylogeographic analyses to unveil patterns of overseas dispersal. Our phylogeographic analyses revealed a high level of geographically unstructured haplotype diversity within taxa, suggesting that transmarine dispersal events have occurred multiple times along the evolutionary history of phoretic species of blister beetles. Larval phoresy may explain the existence of idiosyncratic biogeographic and phylogeographic patterns in these flightless organisms., This work was funded by the Spanish Government project-grant PID2019-110243GB-100 (Ministerio de Ciencia e Innovación-FEDER) to M. García-París.

nostraca are known by their ability for long-distance dispersal, but the existence in several spe-cies of deep, geographically structured mtDNA lineages suggests their populations are subjectedto allopatric differentiation, isolation, and prevalence of local scale dispersion.Tanymastix stagna-lisis one of the most widespread species of Anostraca and previous studies revealed an uncleargeographical pattern of mtDNA genetic diversity. Here, we analyze populations from the Iberianand Italian Peninsulas, Central Europe, and Scandinavia, with the aim to characterize the patternsof genetic diversity in a spatio-temporal framework using mtDNA and nuclear markers to test geneflow among close populations. For these aims we built a time-calibrated phylogeny and carried outBayesian phylogeographic analyses using a continuous diffusion model. Our results indicated thatT. stagnalispresents a deeply structured genetic diversity, including 7 ancient lineages, some ofthem even predating the Pleistocene. The Iberian Peninsula harbors high diversity of lineages, withstrong isolation and recent absence of gene flow between populations. Dispersal at local scaleseems to be the prevailing dispersal mode ofT. stagnalis, which exhibits a pattern of isolation-by-distance in the Iberian Peninsula. We remark the vulnerability of most of these lineages, giventhe limited known geographic distribution of some of them, and the high risk of losing importantevolutionary potential for the species., This work was supported by the Project Grant CGL2015-66571-P (MINECO/FEDER) of Spain.

Millipede diversity in tropical regions, and in Mexico in particular, is still mostly unknown. A modest but recurrent source of new Mexican species is the colonization of exotic species, due to human activity. The invasive speciesCylindrodesmus hirsutusPocock, 1889 has spread from its area of origin in Indonesia or Melanesia and become a virtually pantropical species. Although long known from South and Central America, reports from the Caribbean are sparse and limited to some eastern islands and southern Central America. On 9 March 2016, two adult specimens were found on Cozumel Island, Quintana Roo, in an area of medium semideciduous tropical forest. This paper comprises the first record of this species from Mexico and the northern Caribbean. Given the intense commercial activity in the region, the presence of more populations both in Cozumel Island and in the mainland coast is highly probable.

Este artículo contiene 11 páginas, 5 figuras, 1 tabla., Dolistenus Fanzago, 1874 is a widespread Mediterranean millipede genus of the order Platydesmida, currently including three valid species, each with relict distributions. Here we describe a fourth species of Dolistenus, and characterize it using morphological and molecular (mitochondrial DNA) characters. We provide an updated key to the species of Dolistenus and the first COI barcode sequences for the new species and several other European representatives of the Andrognathidae (Platydesmida).

Facultative or obligate paedomorphosis has evolved several times in Mexican populations of the salamander genus Ambystoma, leading to increased genetic divergence among populations with alternate life histories and contributing to population divergence in this species complex. In the present study, we surveyed the genetic diversity of Ambystoma populations in lakes of the Cuenca Oriental, a high elevation closed drainage basin that encompasses permanent crater lakes harbouring salamander populations. We genotyped individuals from five populations aiming to better understand population dynamics and the evolution of paedomorphosis in this system. Specifically, we tested the hypotheses that the evolution of paedomorphosis in Ambystoma taylori resulted in reduced genetic exchange with populations of Ambystoma velasci in neighbouring lakes. Second, we tested whether the populations in brackish lakes of the Cuenca Oriental, Lake Atexcac, and Lake Alchichica show restricted gene flow across the basin, possibly as a result of local adaptation to those microhabitats. Using various indices of population genetic diversity, Bayesian assignment, and approximate Bayesian computation methods, we show that genetic exchange between brackish lakes and freshwater lakes is negligible, despite continued gene flow among freshwater lakes. We show that the first divergence among populations occurred between Alchichica and the remaining populations and that the evolution of paedomorphosis in A. taylori was likely favoured by local adaptation to saline conditions, thus increasing its genetic isolation. An apparently similar process appears to be in progress independently in lake Atexcac, showing that local adaptation may play an important role in population isolation and, ultimately, in speciation.

[Aim]: To reconstruct the historical biogeography of Hyla molleri, a tree frog endemic to the Eurosiberian and Mediterranean bioclimatic zones in the Iberian Peninsula. Location: Iberian Peninsula., [Methods]: We used molecular data (mtDNA and species-specific, polymorphic microsatellite loci) and species distribution modelling (SDM) from the Last Interglacial (LIG) to the present to characterize spatial patterns of genetic diversity in this species and assess their relationship with climatically favourable areas through time., [Results]: Genetic diversity is heterogeneously distributed across the range of H. molleri, with two main genetic reservoirs located in (a) central and southern Portugal and (b) a fragmented area encompassing mountainous areas in northern Spain. According to SDM, the Iberian Peninsula has experienced a progressive and continuous decrease in climatically favourable areas for H. molleri since the LIG, especially in southern and eastern Iberia, where the species is currently absent. However, we found no correlation between areas that have remained climatically favourable since the LIG and current genetic diversity., [Main conclusions]: Our results suggest that the demographic history of H. molleri since the Pleistocene has been characterized by relative stability, contrasting with the large-scale cycles of extinction-recolonization inferred for other more thermophilous, co-distributed amphibian species in Iberia. Accounting for discordant demographic responses to climatic changes across syntopic species provides new insights about the evolutionary history of amphibian communities in southern Europe., This research was funded by grants CGL2008‐04271‐C02‐01/BOS and CGL2011‐28300 (Ministerio de Ciencia e Innovación, Ministerio de Economía y Competitividad, Spain and FEDER) to I.M.S., who was supported by funding from the Spanish Ramón y Cajal (RYC‐2007‐01668) and Severo Ochoa (SEV‐2012‐0262) programmes. A.M.B. was supported by the Portuguese “Investigador FCT” programme through contract and exploratory project IF/00266/2013/CP1168/CT0001.

Currently with 85 named species, the family Platyrhacidae in America is still poorly studied, particularly regarding Barydesmus Cook, 1896, the most diversified among the American genera. The center of diversification of Barydesmus lies in the northern third of the Andes Mountains, a megadiverse region where surely more species of these large, conspicuous diplopods await description. Here we present a new species, Barydesmus nangaritza sp. nov., easily diagnosable by the shape of the paranota, unique among the American Platyrhacidae. We provide also an updated bibliographical, annotated checklist of the whole family in America, with the aim of encouraging further studies in the group. The checklist includes the following new combinations under the genus Barydesmus: Barydesmus acanthopleurus (Hoffman, 1960) comb. nov., B. acanthosternus (Brölemann, 1900) comb. nov., B. acompus (Chamberlin, 1941) comb. nov., B. aequinoctius (Attems, 1914) comb. nov., B. affinis (Brölemann, 1919) comb. nov., B. andinus (Cook, 1896) comb. nov., B. azulae (Kraus, 1956) comb. nov., B. balsapuertus (Chamberlin, 1941) comb. nov., B. bifasciatus (Silvestri, 1897) comb. nov., B. bombonus (Chamberlin, 1941) comb. nov., B. broelemanni (Attems, 1914) comb. nov., B. brunnior (Chamberlin, 1941) comb. nov., B. celinus (Chamberlin, 1941) comb. nov., B. chuncho (Chamberlin, 1941) comb. nov., B. clathratus (Gervais, 1847) comb. nov., B. comptus Cook, 1896 comb. nov., B. contayus (Chamberlin, 1941) comb. nov., B. dunalii (Gervais, 1847) comb. nov., B. exsul (Cook, 1896) comb. nov., B. festae (Silvestri, 1897) comb. nov., B. fuscatus (Chamberlin, 1941) comb. nov., B. helophorus (Attems, 1899) comb. nov., B. incus (Chamberlin, 1941) comb. nov., B. javarynus (Schubart, 1950) comb. nov., B. leucus (Chamberlin, 1941) comb. nov., B. loretus (Chamberlin, 1941) comb. nov., B. manserichus (Chamberlin, 1941) comb. nov., B. medius (Chamberlin, 1941) comb. nov., B. obscurus (Kraus, 1955) comb. nov., B. orellanus (Chamberlin, 1941) comb. nov., B. pococki (Brölemann, 1911) comb. nov., B. retentus (Chamberlin, 1941) comb. nov., B. riparius (Carl, 1902) comb. nov., B. rufipes (Koch, 1847) comb. nov., B. scaber (Koch, 1847) comb. nov., B. socius (Chamberlin, 1941) comb. nov., B. strenuus (Silvestri, 1897) comb. nov., B. tambonus (Chamberlin, 1952) comb. nov., B. tapichus (Chamberlin, 1941) comb. nov., B. trichotypus (Chamberlin, 1941) comb. nov., B. utoquinius (Chamberlin, 1941) comb. nov. and B. zygethus (Chamberlin, 1941) comb. nov.

Author(s): Rovito, SM; Parra-Olea, G; Recuero, E; Wake, DB | Abstract: The Neotropical bolitoglossine salamanders represent an impressive adaptive radiation, comprising roughly 40% of global salamander species diversity. Despite decades of morphological studies and molecular work, a robust multilocus phylogenetic hypothesis based on DNA sequence data is lacking for the group. We estimated species trees based on multilocus nuclear and mitochondrial data for all major lineages within the bolitoglossines, and used our new phylogenetic hypothesis to test traditional biogeographical scenarios and hypotheses of morphological evolution in the group. In contrast to previous phylogenies, our results place all Central American endemic genera in a single clade and suggest that Central America played a critical role in the early biogeographical history of the group. The large, predominantly Mexican genus Pseudoeurycea is paraphyletic, and analyses of the nuclear data place two lineages of Pseudoeurycea as the sister group of Bolitoglossa. Our phylogeny reveals extensive homoplasy in morphological characters, which may be the result of truncation or alteration of a shared developmental trajectory. We used our phylogenetic results to revise the taxonomy of the genus Pseudoeurycea.

European tree frogs ( Hyla ) characterized by short temporal parameters of the advertisement call form six genetically differentiated but morphologically cryptic taxa, H. arborea sensu stricto, H. orientalis and H. molleri from across Europe to western Asia (together referred to as H. arborea sensu lato), two putative taxa within H. intermedia (Northern and Southern) from the Italian Peninsula and Sicily, and H. sarda from Sardinia and Corsica. Here, we assess species limits and phylogenetic relationships within these ‘short-call tree frogs’ based on mitochondrial DNA and nuclear protein-coding markers. The mitochondrial and nuclear genes show partly incongruent phylogeographic patterns, which point to a complex history of gene flow across taxa, particularly in the Balkans. To test the species limits in the short-call tree frogs and to infer the species tree, we used coalescent-based approaches. The monophyly of H. arborea sensu lato is supported by the mtDNA as well as by the all-gene species tree. The Northern and Southern lineages of H. intermedia have been connected by nuclear gene flow (despite their deep mtDNA divergence) and should be treated as conspecific. On the contrary, the parapatric taxa within H. arborea sensu lato should be considered distinct species ( H. arborea , H. orientalis , H. molleri ) based on the coalescent analysis, although signs of hybridization were detected between them ( H. arborea × H. orientalis ; H. arborea × H. molleri ). A mitochondrial capture upon secondary contact appears to explain the close mtDNA relationship between the geographically remote Iberian H. molleri and H. orientalis from around the Black Sea. Introgressive hybridization occurred also between the Balkan H. arborea and northern Italian H. intermedia , and between the Minor Asiatic H. orientalis and Arabian H. felixarabica (the latter belonging to a different acoustic group/clade). Our results shed light on the species limits in the European short-call tree frogs and show that introgression played an important role in the evolutionary history of the short-call tree frogs and occurred even between taxa supported as distinct species.

Genetic characterization of species using phylogeographic approaches represents a basic reference to understand their evolutionary history as well as to identify conservation priorities to protect areas of particular interest regarding evolutionary potential. Even in well-studied regions such information is lacking for the majority of species, including many endemic species with reduced distribution ranges. We investigate the phylogeographic pattern of the Iberian frog Rana iberica, an endemic amphibian restricted to Central and North-Western Iberian Peninsula. Using mitochondrial sequences, we reconstruct the phylogeographic history of the species to test the effect of Quaternary climate changes on the evolutionary diversification of lineages, that is, the differentiation of mitochondrial lineages and the formation of genetic diversity melting pots, and integrate phylogeographic evidence for future conservation planning. Our results indicate the existence of 3 main mitochondrial lineages differentiated during the Upper Pleistocene. Both historical demographic analyses and climatic niche modeling show a strong effect of glacial climate changes, suggesting recurrent range contractions and expansions. Under such circumstances, differentiation took place most likely by isolation in allopatric interglacial refugia. Secondary lineage admixture in northern Portugal generated a broad mixed zone with highest nucleotide diversity. Given its particular evolutionary potential, its reduced distribution and eventual threats under current climate change scenario, conservation priorities should focus on the isolated lineage from Sierra de Guadalupe., This work was supported by the Portuguese Foundation for Science and Technology (FCT) through post doc grants (SFRH/BPD/27173/2006 to J.T.) and (SFRH/BPD/26555/2006 to H.G.). Currently, H.G. is supported by a postdoctoral Grant from FCT (SFRH/BPD/102966/2014). Additional support was provided from project CGL2015-66571-P by Spanish Ministerio de Economía y Competitividad (MINECO/FEDER).

Genetic characterization of species using phylogeographic approaches represents a basic reference to understand their evolutionary history as well as to identify conservation priorities to protect areas of particular interest regarding evolutionary potential. Even in well-studied regions such information is lacking for the majority of species, including many endemic species with reduced distribution ranges. We investigate the phylogeographic pattern of the Iberian frog Rana iberica, an endemic amphibian restricted to Central and North-Western Iberian Peninsula. Using mitochondrial sequences, we reconstruct the phylogeographic history of the species to test the effect of Quaternary climate changes on the evolutionary diversification of lineages, that is, the differentiation of mitochondrial lineages and the formation of genetic diversity melting pots, and integrate phylogeographic evidence for future conservation planning. Our results indicate the existence of 3 main mitochondrial lineages differentiated during the Upper Pleistocene. Both historical demographic analyses and climatic niche modeling show a strong effect of glacial climate changes, suggesting recurrent range contractions and expansions. Under such circumstances, differentiation took place most likely by isolation in allopatric interglacial refugia. Secondary lineage admixture in northern Portugal generated a broad mixed zone with highest nucleotide diversity. Given its particular evolutionary potential, its reduced distribution and eventual threats under current climate change scenario, conservation priorities should focus on the isolated lineage from Sierra de Guadalupe.

El isópodo terrestre Armadillidium mateui Vandel, 1953 constituye un endemismo de la Península Ibérica poco común y prácticamente desconocido. La especie presenta una distribución fuertemente disyunta, con solo ocho registros publicados de tres regiones diferentes en el sureste, este y norte de España. Viendo este extraño patrón de distribución, la identidad de parte de la información publicada ha sido cuestionada. En este trabajo presentamos nuevos registros en la península ibérica y discutimos sobre la distribución conocida y potencial del isópodo endémico A. mateui, aparentemente mayor de lo previamente esperado.

The Common toad Bufo bufo sensu lato is a widespread, morphologically conserved taxon. Recent studies have uncovered deep genetic differentiation between population groups, highlighting the need to revise the current taxonomy of the group and recognize additional species. Here we investigate patterns of variation in molecular (a mitochondrial DNA restriction enzyme assay and sequence data for two nuclear DNA fragments totalling 979 bp) and 17 morphological variables in Northern France where two of these groups meet (B. bufo sensu stricto and B. spinosus), in order to delineate their contact zone and uncover characters that would allow discrimination of the two taxa. Mitochondrial DNA data show an abrupt transition from areas where B. bufo is present to those inhabited by B. spinosus, with a narrow area of overlap east of the city of Caen. Morphometric characters, particularly those related to the positioning of the parotoid glands and metatarsal tubercle shape and size, proved useful in discriminating between species (AUC ≥ 0.97, kappa ≥ 0.79). We then used the differentiating character states to allocate over 300 museum specimens from Western Europe to either species with consistent results, including comparable values of AUC and kappa of the identification models, indicating that models could successfully be applied across datasets. We summarize available evidence relevant to the delineation of the distribution of B. bufo and B. spinosus in France and discuss the characters differentiating both species in an evolutionary context. In view of the observed morphological and genetic differentiation and the absence of unequivocal evidence for widespread hybridization we support the view that B. bufo and B. spinosus are best considered different species. Finally, we propose that ‘parotoids in parallel position’ and a thin and smooth skin are derived character states for B. bufo over the northern part of its range.

We investigated the recent evolutionary history of the Danube crested newt, Triturus dobrogicus through reconstructions of: (1) the number and position of refugia at the last glacial maximum, (2) the role of major central European rivers in pattern of post-glacial dispersal, and (3) the present-day distribution pattern. We analysed sequences of mitochondrial DNA (ND2, 1065 bp) and six microsatellite loci in 363 T. dobrogicus individuals from 58 populations covering the range of the species. Our analyses suggested that T. dobrogicus survived the last glacial maximum in two separate refugia positioned in northwestern Pannonia and in Southern Pannonia from where its range expanded along the Danube and Tisza Rivers. Our findings also confirmed that rivers played an important role in shaping the evolutionary history of amphibian species in Central Europe. We compared the T. dobrogicus range with another lowland amphibian, the fire-bellied toad Bombina bombina, using species distribution modelling. In line with these models, the isolated mountains inside Pannonia are occupied not by B. bombina, but by its mountain-dwelling sister-species B. variegata. However, in contrast to the model, crested newts (including T. dobrogicus) are absent from these mountains. We attribute this biogeographical discrepancy to the positioning of the species’ refugia at the last glacial maximum., JV was supported by the project Faunagenezis of the National Office for Research and Technology, Hungary (NKFP 3B-02304), the Hungarian Scientific Research Fund (OTKA K84071), and the Bolyai János Research Scholarship of the Hungarian Academy of Sciences. ER was supported by a DGAPA-UNAM postdoctoral fellowship. A research permit was provided by the Ministry of Environment and Water, Hungary (KJHF-848/3/2005).

Niche conservatism has been proposed as the mechanism driving speciation in temperate montane clades through range fragmentation during climatic oscillations. Thus, a negative relationship between speciation rates and niche width is expected. Here, we test this prediction using American zopherine beetles. Our phylogenetic analyses recovered two clades in addition to that of the genus Zopherus: the genera Verodes and Phloeodes, which originated most likely in the Eocene, and diversified during the Miocene and the Pliocene. The assessment of clade niche width in relation to clade diversity supported the proposition of narrow niches leading to a higher probability of range fragmentation during climatic oscillations, thus increasing speciation. Additionally, almost all current populations of Phloeodes and Verodes are located within regions that retained favourable climatic conditions across warm and cold Pleistocene periods, suggesting that dispersal limitation is a strong factor controlling clade distribution. In sum, our results suggest that (i) niche width is a major determinant of the probability of speciation in temperate montane clades, by controlling the probability of potential range fragmentation and (ii) dispersal limitation is also a major determinant of the speciation process, by increasing the fragmentation of realized ranges even when potential distributions are cyclically fused during climatic oscillations. When dispersal limitation is extreme, as in zopherine beetles, populations persist just in those areas that have retained suitable conditions during extremes of past climatic oscillations. Paradoxically, this relict condition confers zopherine beetles great resilience for facing future climate change.

The loss of genetic diversity in small or isolated populations can increase inbreeding, decrease fitness and adaptive potential and increase a species’ probability of extinction. In species with life histories that naturally result in small populations and/or low levels of gene flow, patterns of anthropogenically induced genetic erosion can be obscured by evolutionary history; yet these species may still be susceptible to genetic loss. We assess genetic diversity among populations of Ambystoma salamanders endemic to Mexico, including populations that are facultatively or obligately paedomorphic, to test whether paedomorphic lineages have lower genetic diversity than metamorphic ones, and whether gene flow contributes to the maintenance of diversity in divergent forms with either life history. We also test the utility of our markers in assigning illegally harvested individuals to populations of origin. We found reduced genetic diversity in paedomorphic compared with some, but not all, metamorphic populations. Populations of both forms showed genetic signatures of bottlenecks, underscoring that factors other than paedomorphosis contribute to historical reductions in population size. In general, Ambystoma populations have low interpopulation gene flow and admixture, but paedomorphic populations have higher within-population relatedness than most metamorphic populations. We discuss historical and current landscape attributes that impact populations and their connectivity, the implications of our findings for ongoing captive propagation programs and the prospects for continued genetic health of Ambystoma in Mexico.

The Pleistocene was characterized by climatic changes that greatly altered the distribution of organisms. Population extinctions, bottlenecks, isolation, range expansions and contractions were often associated with glaciations, leaving signatures in the spatial patterns of genetic diversity across species. Lissotriton helveticus belongs to a Pan-European lineage of newts that were strongly affected by glaciations and represent an excellent model to analyse the effect of generalized climatic changes in phylogeographic patterns. We studied the genetic diversity of the species using data from two mitochondrial and three nuclear genes analyzed in a Bayesian phylogenetic framework to investigate the historical processes shaping spatial patterns of genetic diversity. Mitochondrial haplotypes cluster in four different groups present in the Iberian Peninsula and of Pleistocene origin, probably by allopatric fragmentation. Nuclear genes present no obvious geographic structure patterns, suggesting gene flow and generalized incomplete lineage sorting. Populations north of the Pyrenees are closely related to those from northeastern Iberia, suggesting recent range expansion from this region. Historical demographic analyses indicate a demographic expansion starting about 100,000 years ago and more recent population declines. Compared to other Lissotriton species, L. helveticus includes only relatively young genetic lineages, suggesting a Central European pre-Pleistocene distribution followed by complete extirpation of the species during glaciations in that area. Historical demographic trends in the Iberian Peninsula are reversed with respect to the more Mediterranean species Lissotriton boscai , indicating different responses of both species to climate changes. Diversity patterns among Lissotriton species seem to be defined by four main factors: ancestral distributions, colonization capabilities, interactions with other species and effective population sizes. Differences in these factors define two types of species, referred to as “R” (refugia) and “S” (sanctuaries) that explain part of the diversity in patterns of genetic diversity created by glaciations in Western Europe.

Species with highly restricted distributions are vulnerable to extinction, and modification of natural habitats within their small ranges is a primary threat to their persistence. Expansion of urban development significantly impacts natural habitats and, therefore, threatens local diversity. The Mexican axolotl, Ambystoma mexicanum, is a strictly aquatic species that persists currently in two highly threatened and isolated populations. The current habitat remaining for these species are remnants of a historically extensive lacustrine system that occupied the entire Valley of Mexico, but has been destroyed by the growth of Mexico City. Unexpectedly, a third viable population of axolotls has been found in Chapultepec Park, a public recreational area in the heart of Mexico City. Phylogenetic and haplotype network analyses of mitochondrial DNA sequences confirmed low genetic differentiation and a recurrent lack of monophyly in many of the taxa belonging to the Ambystoma tigrinum species group, including A. me...

13 páginas, 3 figuras, 2 tables et al.., Aim To study the patterns of genetic variation and the historical events and processes that influenced the distribution and intraspecific diversity in Hyla meridionalis Boettger, 1874. Location Hyla meridionalis is restricted to the western part of the Mediterranean region. In northern Africa it is present in Tunisia, Algeria and Morocco. In southwestern Europe it is found in the south of France, north-western Italy and northeastern and south-western Iberian Peninsula. There are also insular populations, as in the Canaries and Menorca. Methods Sampling included 112 individuals from 36 populations covering the range of the species. We used sequences of mitochondrial DNA Cytochrome Oxidase I (COI) for the phylogeographical analysis (841 bp) and COI plus a fragment including part of tRNA lysine, ATP synthase subunits 6 and 8 and part of Cytochrome Oxidase III for phylogenetic analyses (2441 bp). Phylogenetic analyses were performed with paup*4.0b10 (maximum likelihood, maximum parsimony) and MrBayes 3.0 (Bayesian analysis). Nested clade analysis was performed using tcs 1.18 and GeoDis 2.2. A dispersal-vicariant analysis was performed with diva 1.0 to generate hypotheses about the geographical distribution of ancestors. Results We found little genetic diversity within samples from Morocco, southwestern Europe and the Canary Islands, with three well-differentiated clades. One is distributed in south-western Iberia and the High Atlas, Anti-Atlas and Massa River in Morocco. The second is restricted to the Medium Atlas Mountains. The third one is present in northern Morocco, north-eastern Iberia, southern France and the Canaries. These three groups are also represented in the nested clade analysis. Sequences from Tunisian specimens are highly divergent from sequences of all other populations, suggesting that the split between the two lineages is ancient. diva analysis suggests that the ancestral distribution of the different lineages was restricted to Africa, and that an explanation of current distribution of the species requires three different dispersal events. Main conclusions Our results support the idea of a very recent colonization of south-western Europe and the Canary Islands from Morocco. South-western Europe has been colonized at least twice: once from northern Morocco probably to the Mediterranean coast of France and once from the western coast of Morocco to southern Iberia. Human transport is a likely explanation for at least one of these events. Within Morocco, the pattern of diversity is consistent with a model of mountain refugia during hyperarid periods within the Pleistocene. Evaluation of the phylogenetic relationships of Tunisian haplotypes will require an approach involving the other related hylid taxa in the area., Support during part of the development of this project for A.I. was provided by an Aranzadi fellowship and for E.R. by an I3P grant of the CSIC.

Widespread species with morphologically and ecologically differentiated populations are key to understand speciation because they allow investigating the different stages of the continuous process of population divergence. The alpine newt, Ichthyosaura alpestris, with a range that covers a large part of Central Europe as well as isolated regions in all three European Mediterranean peninsulas, and with strong ecological and life-history differences among populations, is an excellent system for such studies. We sampled individuals across most of the range of the species, and analyzed mitochondrial (1442 bp) and nuclear (two nuclear genes -1554 bp- and 35 allozyme loci) markers to produce a time-calibrated phylogeny and reconstruct the historical biogeography of the species. Phylogenetic analyses of mtDNA data produced a fully resolved topology, with an endemic, Balkan clade (Vlasina) which is sister to a clade comprising an eastern and a western group. Within the former, one clade (subspecies I. a. veluchiensis) is sister to a clade containing subspecies I. a. montenegrina and I. a. serdara as well as samples from southern Romania, Bosnia-Herzegovina, Serbia and Bulgaria (subspecies I. a. reiseri and part of I. a. alpestris). Within the western group, populations from the Italian peninsula (subspecies I. a. apuana and I. a. inexpectata) are sister to a clade containing samples from the Iberian Peninsula (subspecies I. a. cyreni) and the remainder of the samples from subspecies I. a. alpestris (populations from Hungary, Austria, Poland, France, Germany and the larger part of Romania). Results of (∗)BEAST analyses on a combined mtDNA and nDNA dataset consistently recovered with high statistical support four lineages with unresolved inter-relationships: (1) subspecies I. a. veluchiensis; (2) subspecies I. a. apuana+I. a. inexpectata; (3) subspecies I. a. cyreni+part of subspecies I. a. alpestris (the westernmost populations, plus most Romanian populations); and (4) the remaining populations, including subspecies I. a. serdara, I. a. reiseri and I. a. montenegrina and part of subspecies I. a. alpestris, plus samples from Vlasina. Our time estimates are consistent with ages based on the fossil record and suggest a widespread distribution for the I. alpestris ancestor, with the split of the major eastern and western lineages during the Miocene, in the Tortonian. Our study provides a solid, comprehensive background on the evolutionary history of the species based on the most complete combined (mtDNA+nDNA+allozymes) dataset to date. The combination of the historical perspective provided by coalescent-based analyses of mitochondrial and nuclear DNA variation with individual-based multilocus assignment methods based on multiple nuclear markers (allozymes) also allowed identification of instances of discordance across markers that highlight the complexity and dynamism of past and ongoing evolutionary processes in the species.

We screened a partial genomic library enriched for microsatellites and characterized nine loci for the Mexican species of Ambystoma for studies of population structure. We tested marker variability in two metamorphic ( A. granulosum , A. altamirani ) and two paedomorphic ( A. andersoni , A. mexicanum ) species of the A. tigrinum complex. Our microsatellites were developed from pooled genomic DNA from three species, and may work on all species in the A. tigrinum complex in Mexico. These markers will be important for studies of conservation genetics in this radiation.

Species delineation remains one of the most challenging tasks in the study of biodiversity, mostly owing to the application of different species concepts, which results in contrasting taxonomic arrangements. This has important practical consequences, since species are basic units in fields like ecology and conservation biology. We here review molecular genetic evidence relevant to the systematics of toads in the Bufo bufo species group (Anura, Bufonidae). Two studies recently published in this journal (Recuero et al., MPE 62: 71-86 and García-Porta et al., MPE 63: 113-130) addressed this issue but reached opposing conclusions on the taxonomy of the group (four versus two species). In particular, allozyme data in the latter paper were interpreted as evidence for hybridization across species (between B. bufo-B. spinosus and B. bufo-B. verrucosissimus). We tested claims for hybridization through re-analysis of allozyme data for individuals instead of populations, to be able to distinguish between sympatry with and without admixture, and found no evidence of hybridization across taxa. We propose alternative explanations for the observed patterns that García-Porta et al. (2012) failed to consider. In the absence of unequivocal evidence for hybridization and introgression, we reject the proposal to downgrade Bufo spinosus and Bufo verrucosissimus to the subspecies level., IMS is a "Ramón y Cajal" postdoctoral fellow supported by the Spanish Ministerio de Ciencia e Innovación and the Universidad de Castilla la Mancha. Partial funds for this work were provided by grants CGL2008-04271-C02-01/BOS and CGL2011-28300 (Ministerio de Ciencia e Innovación, Ministerio de Economía y Competitividad, FEDER) and PPII10-0097-4200 (Junta de Comunidades de Castilla la Mancha) to IMS.

12 páginas, 5 figuras y 2 tables.., The Baja California populations of Pseudacris regilla, a widespread species in Western North America ranging from British Columbia to southern Baja California, are characterized by extensive geographic fragmentation. We performed phylogeographic and historical demographic analyses on 609 bp of the cytochrome b mitochondrial gene of 110 individuals representing 28 populations to determine the relative influences of current and historical processes in shaping the present distribution of genetic diversity on the Baja California Peninsula. Haplotypes from this area were nested in a clade with three well-differentiated groups. Two of these groups are from Baja California Sur and another is from California and Baja California. The estimated date for the split of these groups, between 0.9–1 Ma, fits with previously proposed hypotheses of vicariance due to diVerent transpeninsular seaways, although successive population fragmentation and expansion due to climatic oscillations during Pleistocene glaciations cannot be discarded. Historical demographic analyses detected signs of past population expansions, especially in the southernmost group. With respect to populations north of this region, two older clades were identified, one with haplotypes mainly distributed in central California, and the other corresponding to the northern half of the species range, in what apparently is a recurrent pattern in the Pacific coast of North America. Based on the concordance between mt-DNA and available allozyme data indicating that these species have a long independent evolutionary history, we propose to consider the three major clades as distinct species: P. regilla, P. pacifica, and P. hypochondriaca., We thank D. Buckley, S. Crews, and A. Leaché for helpful comments on the manuscript. Thanks to the Museum of Vertebrate Zoology, University of California, Berkeley for samples from the Tissue Collection. E.R. is sponsored by a fellowship CSIC-postgrado (Ref.: I3P-BPG2004) and I.M.-S. is funded by a postdoctoral Grant from the Spanish Ministerio de Educación y Ciencia (Ref. EX2004-0921). SEMARNAT provided collecting permits. Studies were financed in part by the Grants UC_Mexus-Conacyt CN-02-74 and PAPIIT-UNAM No. IN226605 to G.P.-O. and by a travel grant (“Marina Bueno”) to I.M.-S. This research has also benefited upon the resources provided by the Grant CGL2004-04680- C10-10/BOS (Ministerio de Educación y Ciencia, Spain).

Este trabajo se centra en el estudio de la diversidad de isópodos terrestres (Isopoda, Oniscidea) en dos microhábitats saproxílicos: oquedades de árboles maduros y madera muerta en suelo, así como en la distribución espacial y temporal de las especies registradas. El estudio se realizó en tres áreas naturales del sur de la provincia de Salamanca dominadas por Quercus pyrenaica Willd. Se dispusieron trampas de emergencia en los dos microhábitats saproxílicos revisándose mensualmente durante un año completo. En total se recolectaron 3692 individuos identificados en 5 especies. La madera muerta en suelo presentó una riqueza de especies superior a las oquedades, mientras que estas últimas registraron una mayor abundancia. Porcellio scaber fue la especie más abundante en las 3 áreas y en los dos microhábitats, si bien mostró preferencia por las oquedades para hibernar. Las Sierras de Béjar y Francia resultó el área natural con mayor número de especies. El estudio de la distribución temporal de las 2 especies principales, P. scaber y P. ingenuus, indicó que no existe solapamiento entre los máximos de actividad de ambas especies. Las dos especies ocuparon preferentemente microhábitats diferentes (P. scaber oquedades y P. ingenuus madera muerta en suelo) como refugio de invernada en los meses de otoño e invierno emergiendo a principios de primavera en el caso de P. ingenuus (abril) y final de primavera y principios de verano en P. scaber (mayo-julio). This work focuses on the study of woodlice diversity (Isopoda, Oniscidea) in two saproxylic microhabitats: hollows of mature trees and downed deadwood, as well as the spatial and temporal distribution of recorded species. The study was carried out in three natural areas in the south of the province of Salamanca dominated by Quercus pyrenaica Willd. Emergence traps were placed in the saproxylic microhabitats, collecting monthly for a year. In total, 3692 individuals, identified in 5 species, were collected. Downed deadwood presented a higher species richness than tree hollows, while tree hollows recorded a greater abundance. Porcellio scaber was the most abundant species in all 3 areas and two microhabitats, although it showed a preference for tree hollows to hibernate. Las Sierras de Béjar y Francia was the natural area with the highest number of species. The temporal distribution of the 2 main species, P. scaber and P. ingenuus, did not show overlapping between the maximum activity of both species. The two species preferably occupied different microhabitats (P. scaber tree hollows and P. ingenuus downed deadwood) as a wintering shelter in autumn and winter, emerging in early spring in the case of P. ingenuus (April) and late spring and early summer in P. scaber (May-July). Ministerio de Economía y Competitividad (proyectos CGL 2011-23658 y CGL2016-78181-R).