Discovery and description of a new species of burrowing frog Sphaerotheca Gnther, 1859 (Anura: Dicroglossidae) from the suburban landscapes of Bengaluru, India.

Recent resolution of prevailing taxonomic ambiguities in the genus Sphaerotheca and new species discoveries from urban/suburban landscapes highlight the need for attention to non-forested habitats for amphibian conservation. In this paper, we review the status of the members of the genus Sphaerotheca and justify the synonymy of Sphaerotheca magadha as a junior synonym of Sphaerotheca swani. The prospects of resurrection of Sphaerotheca swani (herein preliminarily referred to as Sphaerotheca cf. breviceps [swani]) are discussed. In addition, we describe a new species Sphaerotheca varshaabhu sp. nov. from the suburban region of Bengaluru, India. We employ an integrative taxonomic approach to characterize the new species using molecular phylogeny, genetic distance, morphological characters, and geographical isolation as lines of evidence. We also provide a description of vocal repertoire of Sphaerotheca varshaabhu sp. nov. and provide comparative bioacoustics data for four species. This previously undescribed species from the suburban areas of Bengaluru described herein as Sphaerotheca varshaabhu sp. nov. forms a genetically divergent lineage and its genetic distance varied from 3.6% to 12.2% for 16S rRNA with respect to other species of Sphaerotheca. Our phylogenetic analysis for the genus including the new species confirms the synonymy of one recently described species, resulting in 10 valid species in the genus Sphaerotheca. These results emphasize the need for utilizing an integrative taxonomic approach for uncovering hidden diversity of suburban areas. Given these recent discoveries, we advocate for more robust surveys in human dominated areas, so that these amphibians may receive more attention.

Species distribution modelling of Hylarana Species (Anura, Ranidae) and the problem of accurate species identification.

Species of Hylarana Tschudi, 1838 are among the most common amphibian species in the Oriental region. Their poorly known geographical distribution impacts species delimitation and conservation actions. We compiled geographical data from literature, online databases and museum collections for the four currently recognized species, Hylarana erythraea, H. taipehensis, H. tytleri and H. macrodactyla. We obtained 1014 occurrence records and allocated a quality rate to each of these points: A-species identification confirmed by ancillary data; B-species identification unconfirmed; and C-conflict between species name, geographic origin and morphological characters; resulted in exclusion of the latter in final analyses. Based on these different datasets we built species occurrence maps for each of the four species. These maps were compared to already available distribution maps, introducing a more precise distribution for the four species but with persistence of data gaps within less prospected areas. We then built Species Distribution Models (SDMs) based on climatic data. Discrepancies between observed distribution ranges and areas with suitable climatic niches are discussed in a taxonomical context. Hylarana species, although taxonomically still not fully resolved and occupying similar habitats, exhibit differences in distribution pattern. Models also fill data gaps, introducing relevant suitable climatic niches within close range of species' distribution areas. Differences were observed between A and B-quality data models, with high suitable habitats being less present in most cases for B-quality. Our study shows the importance of merging A and B-quality data, represented by A+B dataset, thus creating more accurate models, but also underlines the necessity to have access to correct data to limit bias.

Molecular and morphological identification of frog species collected at Rara Lake in Rara National Park, Nepal.

This study seeks to clarify the taxonomic identity of three adult frogs and two tadpoles from Rara Lake situated in Rara National Park, Mugu district, Western Nepal, using both phenotypic and mitochondrial sequence data (16S rRNA gene). Based on the molecular data, we determined our specimens belonged to two species; Paa cf. ercepeae (Dubois, 1974) and Paa rarica (Dubois, Matsui, and Ohler, 2001), and this assignment is also supported by morphology. Principal component analysis of phenotypic data clustered species into large-bodied (SVL > 55 mm) and small-bodied (SVL < 55 mm) groups. The molecular data suggest that Paa rarica belonged to a group that contains several undescribed lineages and that Paa liebigii (Gnther, 1860) hosts extensive cryptic diversity. Our study also documents that Paa species in the Himalayan region are more widely distributed than previously believed.

ZooNom: an online thesaurus for alleviating ambiguity in the terminology of zoological nomenclature.

Zoological nomenclature is the discipline of taxonomy responsible for regulating the scientific names of animal species. It has its roots in Carolus Linnaeus work and has been governed by an international Code since the turn of the 20th century. Its vocabulary, on the other hand, is not always clear. Various authors have established new terminology in order to reduce ambiguity. To make these new terms, but also the classical terms used by the International Code of Zoological Nomenclature, accessible, an electronic thesaurus (link: https://www.loterre.fr/skosmos/FM8/en/) was created, allowing to compare existing terminologies. This thesaurus is also a tool for reflection and discussion, targeting taxonomists and experts in nomenclature.

A new species of Sphaerotheca Günther, 1859 (Anura: Dicroglossidae) from the degraded urban ecosystems of Bengaluru, Deccan Plateau, India.

The dicroglossid genus Sphaerotheca was erected by Günther in 1859, with its range of distribution in South Asia. Since then, 13 species have been described and 10 species are considered valid. Many of these descriptions were from low to mid-elevation ranges of homestead areas, agroecosystems and degraded landscapes with a few from forested areas. In the present account, a new species of Sphaerotheca, a genetically distinct lineage with a morphological character set distinguishable from its congeners, is described from the surroundings of Bengaluru city, India. Based on evidence for the origin of the original specimen, we also designated a neotype for Rana variegata Gravenhorst, 1829, a junior primary homonym of Rana variegata Linnaeus, 1758, and permanently invalid name, so that it is now a subjective synonym of Rana (Tomopterna) breviceps rolandae Dubois, 1983.

'Barcode fishing' for archival DNA from historical type material overcomes taxonomic hurdles, enabling the description of a new frog species.

Taxonomic progress is often hindered by intrinsic factors, such as morphologically cryptic species that require a broad suite of methods to distinguish, and extrinsic factors, such as uncertainties in the allocation of scientific names to species. These uncertainties can be due to a wide variety of factors, including old and poorly preserved type specimens (which contain only heavily degraded DNA or have lost important diagnostic characters), inappropriately chosen type specimens (e.g. juveniles without diagnostic characters) or poorly documented type specimens (with unprecise, incorrect, or missing locality data). Thanks to modern sequencing technologies it is now possible to overcome many such extrinsic factors by sequencing DNA from name-bearing type specimens of uncertain assignment and assigning these to known genetic lineages. Here, we apply this approach to frogs of the Mantidactylus ambreensis complex, which was recently shown to consist of two genetic lineages supported by concordant differentiation in mitochondrial and nuclear genes. These lineages co-occur on the Montagne d'Ambre Massif in northern Madagascar but appear to have diverged in allopatry. We use a recently published bait set based on three mitochondrial markers from all known Malagasy frog lineages to capture DNA sequences from the 127-year-old holotype of Mantidactylus ambreensis Mocquard, 1895. With the obtained sequences we are able to assign the name M. ambreensis to the lowland lineage, which is rather widespread in the rainforests of northern Madagascar, leaving the microendemic high-elevation lineage on Montagne d'Ambre in north Madagascar in need of description. We describe this species as Mantidactylus ambony sp. nov., differing from M. ambreensis in call parameters and a smaller body size. Thus, using target enrichment to obtain DNA sequence data from this old specimen, we were able to resolve the extrinsic (nomenclatural) hindrances to taxonomic resolution of this complex. We discuss the broad-scale versatility of this 'barcode fishing' approach, which can draw on the enormous success of global DNA barcoding initiatives to quickly and efficiently assign type specimens to lineages.

Sexual Dichromatism Drives Diversification within a Major Radiation of African Amphibians.

Theory predicts that sexually dimorphic traits under strong sexual selection, particularly those involved with intersexual signaling, can accelerate speciation and produce bursts of diversification. Sexual dichromatism (sexual dimorphism in color) is widely used as a proxy for sexual selection and is associated with rapid diversification in several animal groups, yet studies using phylogenetic comparative methods to explicitly test for an association between sexual dichromatism and diversification have produced conflicting results. Sexual dichromatism is rare in frogs, but it is both striking and prevalent in African reed frogs, a major component of the diverse frog radiation termed Afrobatrachia. In contrast to most other vertebrates, reed frogs display female-biased dichromatism in which females undergo color transformation, often resulting in more ornate coloration in females than in males. We produce a robust phylogeny of Afrobatrachia to investigate the evolutionary origins of sexual dichromatism in this radiation and examine whether the presence of dichromatism is associated with increased rates of net diversification. We find that sexual dichromatism evolved once within hyperoliids and was followed by numerous independent reversals to monochromatism. We detect significant diversification rate heterogeneity in Afrobatrachia and find that sexually dichromatic lineages have double the average net diversification rate of monochromatic lineages. By conducting trait simulations on our empirical phylogeny, we demonstrate that our inference of trait-dependent diversification is robust. Although sexual dichromatism in hyperoliid frogs is linked to their rapid diversification and supports macroevolutionary predictions of speciation by sexual selection, the function of dichromatism in reed frogs remains unclear. We propose that reed frogs are a compelling system for studying the roles of natural and sexual selection on the evolution of sexual dichromatism across micro- and macroevolutionary timescales.

Polypedates smaragdinus Blyth, 1852-a senior subjective synonym of Rhacophorus maximus Günther, 1858.

After more than 160 years, the onomatophores (name-bearing types) of Polypedates smaragdinus Blyth, 1852 were located in the collection of the ZSI (Kolkata, India). This allowed to study the specimens, to identify them and to propose an allocation of this nomen to the taxon Rhacophorus maximus. To stabilize nomenclatural and taxonomic issues, a lectophoront (lectotype) is designated for Polypedates smaragdinus Blyth, 1852 and described. A formal synonymy list is given. The species currently known as Rhacophorus maximus Günther, 1864 should now bear the nomen Rhacophorus smaragdinus (Blyth, 1852).

A new Reed Frog (Hyperoliidae: Hyperolius) from coastal northeastern Mozambique.

A new species of African reed frog (genus Hyperolius Rapp, 1842) is described from the Coastal Forests of the Eastern Africa Biodiversity Hotspot in northeastern Mozambique. It is currently only known from less than ten localities associated with the Mozambican coastal pans system, but may also occur in the southeastern corner of Tanzania. Phylogenetic reconstructions using the mitochondrial 16S marker revealed that it is the sister taxon of Hyperolius mitchelli (>5.6% 16S mtDNA sequence divergence) and forms part of a larger H. mitchelli complex with H. mitchelli and H. rubrovermiculatus. The new species is distinguished from other closely related Hyperolius species by genetic divergence, morphology, vocalisation, and dorsal colouration.

The identity of the South African toad Sclerophrys capensis Tschudi, 1838 (Amphibia, Anura).

The toad species Sclerophrys capensis Tschudi, 1838 was erected for a single specimen from South Africa which has never been properly studied and allocated to a known species. A morphometrical and morphological analysis of this specimen and its comparison with 75 toad specimens referred to five South African toad species allowed to allocate this specimen to the species currently known as Amietophrynus rangeri. In consequence, the nomen Sclerophrys must replace Amietophrynus as the valid nomen of the genus, and capensis as the valid nomen of the species. This work stresses the usefulness of natural history collections for solving taxonomic and nomenclatural problems.

When young are more conspicuous than adults: a new ranid species (Anura: Ranidae) revealed by its tadpole.

Tadpoles of Clinotarsus alticola collected nearby the type locality in Assam, India are barcoded and described. A detailed morphological and morphometrical description of the specimens, along with a study of the anatomy of the buccal cavity are provided. A comparison of these tadpoles with "Clinotarsus alticola" tadpoles from peninsular Thailand and of the genetic variation of a fragment of their mtDNA 16S gene led us to assign the population of peninsular Thailand to a new species, Clinotarsus penelope sp. n. The holotype of the new species is chosen among the tadpole series as no adult could be found in the type locality. Presumed conspecific adults of nearby localities are morphologically described and compared to barcoded adults of Clinotarsus alticola, waiting for further molecular confirmation. The tadpole of the new species differs from that of C. alticola by a much greater size at comparative stages (e.g., 77.7 mm vs. 53.3 mm in stage 36, respectively), a black coloration (vs. a yellow-olive tinge), several ocelli on the tail muscle (vs. only one), a rounded snout (vs. a more pointed snout) and a different Keratodont Row Formula (KRF; nine keratodonts rows maximum on both labia in C. penelope vs. eight maximum in C. alticola). A discussion about the choice of the holotype, the assignment of adult specimens and the future confirmation of this assignment are provided, as well as a comparison with older descriptions of "Clinotarsus alticola" sensu lato tadpoles and with Clinotarsus curtipes tadpoles from Karnataka, India. The lectotype of Clinotarsus alticola is redescribed.

The authorships and dates of the specific nomina Megophrys shuichengensis and Pseudohynobius shuichengensis (Amphibia).

Two amphibian species from China are designated by the specific nomen shuichengensis, which refers to the Shuicheng County (26°34'N, 104°51'E), south of the city of Liupanshui in the province of Guizhou: Megophrys shuichengensis (Amphibia, Anura) and Pseudohynobius shuichengensis (Amphibia, Urodela). The holotypes (holophoronts) of both species were deposited in Department of Biology of the Liupanshui Teachers Higher College (LTHC below). Both species share the particularity of having been described as new twice, at different dates, in different journals and with different authorships. Although this has been acknowledged for the salamander, it has not yet been so for the frog.

Improving biodiversity assessment of anuran amphibians using DNA barcoding of tadpoles. Case studies from Southeast Asia.

Amphibian populations are dramatically declining, while their inventory is far from being achieved. Tadpoles are usually overlooked from biodiversity survey, whereas their consideration will optimize species counts and knowledge of their ecological and developmental requirements is essential in conservation planning. Two mitochondrial markers, 16S (397 new sequences obtained) and COI (343 new sequences obtained), are used to test DNA barcoding on a set of larval and adult Asian amphibians represented by 83 recognized species from 65 sites. The advantages and drawbacks of each marker are assessed, COI barcoding being advocated for global DNA barcoding, whereas 16S suits for taxonomically or geographically restricted DNA barcoding. About half of the collected tadpoles were badly identified or incompletely named in the field. All tadpole sequences (except one case of probable introgressive hybridization) were correctly assigned to their respective species. Finally six clusters of tadpole sequences without conspecific adults were revealed, stressing the importance of collecting and taking into account tadpoles in biodiversity survey and conservation planning.

Does counting species count as taxonomy? On misrepresenting systematics, yet again.

Recent commentary by Costello and collaborators on the current state of the global taxonomic enterprise attempts to demonstrate that taxonomy is not in decline as feared by taxonomists, but rather is increasing by virtue of the rate at which new species are formally named. Having supported their views with data that clearly indicate as much, Costello et al. make recommendations to increase the rate of new species descriptions even more. However, their views appear to rely on the perception of species as static and numerically if not historically equivalent entities whose value lie in their roles as "metrics". As such, their one-dimensional portrayal of the discipline, as concerned solely with the creation of new species names, fails to take into account both the conceptual and epistemological foundations of systematics. We refute the end-user view that taxonomy is on the rise simply because more new species are being described compared with earlier decades, and that, by implication, taxonomic practice is a formality whose pace can be streamlined without considerable resources, intellectual or otherwise. Rather, we defend the opposite viewpoint that professional taxonomy is in decline relative to the immediacy of the extinction crisis, and that this decline threatens not just the empirical science of phylogenetic systematics, but also the foundations of comparative biology on which other fields rely. The allocation of space in top-ranked journals to propagate views such as those of Costello et al. lends superficial credence to the unsupportive mindset of many of those in charge of the institutional fate of taxonomy. We emphasize that taxonomy and the description of new species are dependent upon, and only make sense in light of, empirically based classifications that reflect evolutionary history; homology assessments are at the centre of these endeavours, such that the biological sciences cannot afford to have professional taxonomists sacrifice the comparative and historical depth of their hypotheses in order to accelerate new species descriptions.

Radically different phylogeographies and patterns of genetic variation in two European brown frogs, genus Rana.

We reconstruct range-wide phylogeographies of two widespread and largely co-occurring Western Palearctic frogs, Rana temporaria and R. dalmatina. Based on tissue or saliva samples of over 1000 individuals, we compare a variety of genetic marker systems, including mitochondrial DNA, single-copy protein-coding nuclear genes, microsatellite loci, and single nucleotide polymorphisms (SNPs) of transcriptomes of both species. The two focal species differ radically in their phylogeographic structure, with R. temporaria being strongly variable among and within populations, and R. dalmatina homogeneous across Europe with a single strongly differentiated population in southern Italy. These differences were observed across the various markers studied, including microsatellites and SNP density, but especially in protein-coding nuclear genes where R. dalmatina had extremely low heterozygosity values across its range, including potential refugial areas. On the contrary, R. temporaria had comparably high range-wide values, including many areas of probable postglacial colonization. A phylogeny of R. temporaria based on various concatenated mtDNA genes revealed that two haplotype clades endemic to Iberia form a paraphyletic group at the base of the cladogram, and all other haplotypes form a monophyletic group, in agreement with an Iberian origin of the species. Demographic analysis suggests that R. temporaria and R. dalmatina have genealogies of roughly the same time to coalescence (TMRCA ~3.5 mya for both species), but R. temporaria might have been characterized by larger ancestral and current effective population sizes than R. dalmatina. The high genetic variation in R. temporaria can therefore be explained by its early range expansion out of Iberia, with subsequent cycles of differentiation in cryptic glacial refugial areas followed by admixture, while the range expansion of R. dalmatina into central Europe is a probably more recent event.

West Africa - a safe haven for frogs? A sub-continental assessment of the chytrid fungus (Batrachochytrium dendrobatidis).

A putative driver of global amphibian decline is the panzootic chytrid fungus Batrachochytrium dendrobatidis (Bd). While Bd has been documented across continental Africa, its distribution in West Africa remains ambiguous. We tested 793 West African amphibians (one caecilian and 61 anuran species) for the presence of Bd. The samples originated from seven West African countries - Bénin, Burkina Faso, Côte d'Ivoire, Ghana, Guinea, Liberia, Sierra Leone - and were collected from a variety of habitats, ranging from lowland rainforests to montane forests, montane grasslands to humid and dry lowland savannahs. The species investigated comprised various life-history strategies, but we focused particularly on aquatic and riparian species. We used diagnostic PCR to screen 656 specimen swabs and histology to analyse 137 specimen toe tips. All samples tested negative for Bd, including a widespread habitat generalist Hoplobatrachus occipitalis which is intensively traded on the West African food market and thus could be a potential dispersal agent for Bd. Continental fine-grained (30 arc seconds) environmental niche models suggest that Bd should have a broad distribution across West Africa that includes most of the regions and habitats that we surveyed. The surprising apparent absence of Bd in West Africa indicates that the Dahomey Gap may have acted as a natural barrier. Herein we highlight the importance of this Bd-free region of the African continent - especially for the long-term conservation of several threatened species depending on fast flowing forest streams (Conraua alleni ("Vulnerable") and Petropedetes natator ("Near Threatened")) as well as the "Critically Endangered" viviparous toad endemic to the montane grasslands of Mount Nimba (Nimbaphrynoides occidentalis).

Nomenclatural and taxonomic problems related to the electronic publication of new nomina and nomenclatural acts in zoology, with brief comments on optical discs and on the situation in botany.

In zoological nomenclature, to be potentially valid, nomenclatural novelties (i.e., new nomina and nomenclatural acts) need first to be made available, that is, published in works qualifying as publications as defined by the International Code of zoological Nomenclature ("the Code"). In September 2012, the Code was amended in order to allow the recognition of works electronically published online after 2011 as publications available for the purpose of zoological nomenclature, provided they meet several conditions, notably a preregistration of the work in ZooBank. Despite these new Rules, several of the long-discussed problems concerning the electronic publication of new nomina and nomenclatural acts have not been resolved. The publication of this amendment provides an opportunity to discuss some of these in detail. It is important to note that: (1) all works published only online before 2012 are nomenclaturally unavailable; (2) printed copies of the PDFs of works which do not have their own ISSN or ISBN, and which are not obtainable free of charge or by purchase, do not qualify as publications but must be seen as facsimiles of unavailable works and are unable to provide nomenclatural availability to any nomenclatural novelties they may contain; (3) prepublications online of later released online publications are unavailable, i.e., they do not advance the date of publication; (4) the publication dates of works for which online prepublications had been released are not those of these prepublications and it is critical that the real release date of such works appear on the actual final electronic publication, but this is not currently the case in electronic periodicals that distribute such online prepublications and which still indicate on their websites and PDFs the date of release of prepublication as that of publication of the work; (5) supplementary online materials and subsequent formal corrections of either paper or electronic publications distributed only online are nomenclaturally unavailable; (6) nomenclatural information provided on online websites that do not have a fixed content and format, with ISSN or ISBN, is unavailable. We give precise examples of many of these nomenclatural problems. Several of them, when they arise, are due to the fact that the availability of nomenclatural novelties now depends on information that will have to be sought not from the work itself but from extrinsic evidence. As shown by several examples discussed here, an electronic document can be modified while keeping the same DOI and publication date, which is not compatible with the requirements of zoological nomenclature. Therefore, another system of registration of electronic documents as permanent and inalterable will have to be devised. ZooBank also clearly needs to be improved in several respects. Mention in a work of its registration number (LSID) in ZooBank would seem to be possible only if this registration has occurred previously, but some works that have purportedly been registered in ZooBank are in fact missing on this web application. In conclusion, we offer recommendations to authors, referees, editors, publishers, libraries and the International Commission on Zoological Nomenclature, in the hope that such problems can be limited along with the potential chaos in zoological nomenclature that could result, if careful attention is not paid to the problems we highlight here, from a somewhat misplaced, and perhaps now widespread, understanding that electronic publication of nomenclatural novelties is now allowed and straightforward. We suggest that, as long as the problematic points linked to the new amendment and to electronic publication as a whole are not resolved, nomenclatural novelties continue to be published in paper-printed journals that have so far shown editorial competence regarding taxonomy and nomenclature, which is not the case of several recent electronic-only published journals.

The origin of modern frogs (Neobatrachia) was accompanied by acceleration in mitochondrial and nuclear substitution rates.

BACKGROUND: Understanding the causes underlying heterogeneity of molecular evolutionary rates among lineages is a long-standing and central question in evolutionary biology. Although several earlier studies showed that modern frogs (Neobatrachia) experienced an acceleration of mitochondrial gene substitution rates compared to non-neobatrachian relatives, no further characterization of this phenomenon was attempted. To gain new insights on this topic, we sequenced the complete mitochondrial genomes and nine nuclear loci of one pelobatoid (Pelodytes punctatus) and five neobatrachians, Heleophryne regis (Heleophrynidae), Lechriodus melanopyga (Limnodynastidae), Calyptocephalella gayi (Calyptocephalellidae), Telmatobius bolivianus (Ceratophryidae), and Sooglossus thomasseti (Sooglossidae). These represent major clades not included in previous mitogenomic analyses, and most of them are remarkably species-poor compared to other neobatrachians. RESULTS: We reconstructed a fully resolved and robust phylogeny of extant frogs based on the new mitochondrial and nuclear sequence data, and dated major cladogenetic events. The reconstructed tree recovered Heleophryne as sister group to all other neobatrachians, the Australasian Lechriodus and the South American Calyptocephalella formed a clade that was the sister group to Nobleobatrachia, and the Seychellois Sooglossus was recovered as the sister group of Ranoides. We used relative-rate tests and direct comparison of branch lengths from mitochondrial and nuclear-based trees to demonstrate that both mitochondrial and nuclear evolutionary rates are significantly higher in all neobatrachians compared to their non-neobatrachian relatives, and that such rate acceleration started at the origin of Neobatrachia. CONCLUSIONS: Through the analysis of the selection coefficient (ω) in different branches of the tree, we found compelling evidence of relaxation of purifying selection in neobatrachians, which could (at least in part) explain the observed higher mitochondrial and nuclear substitution rates in this clade. Our analyses allowed us to discard that changes in substitution rates could be correlated with increased mitochondrial genome rearrangement or diversification rates observed in different lineages of neobatrachians.

Discordant patterns of nuclear and mitochondrial introgression in Iberian populations of the European common frog (Rana temporaria).

Amphibians often show complex histories of intraspecific and interspecific genetic introgression, which might differ in mitochondrial and nuclear genes. In our study of the genetic differentiation of the European common frog, Rana temporaria (159 specimens from 23 populations were analyzed for 24 presumptive allozyme loci; 82 specimens were sequenced for a 540-bp fragment of the mitochondrial 16S rRNA gene), multilocus correspondence analysis (CA) and Bayesian assignment tests of the nuclear data were concordant in identifying 2 population groups corresponding to 1) the Pyrenees in the east and 2) the Galicia and Asturias regions in the west, the latter corresponding to the subspecies R. temporaria parvipalmata. Geographically intermediate populations were genetically intermediate in the allozyme CA and, less clearly in the Bayesian assignment, with mitochondrial haplotypes exclusively belonging to the parvipalmata group. This indicates different degrees of introgression in the mitochondrial and nuclear genomes. Although Pyrenean high-altitude populations are morphologically distinct from low-altitude populations, these 2 groups were not separate clusters in any analysis. This suggests that the morphological differences may be due to fast adaptation to elevational gradients, likely under maintenance of gene flow, and that the underlying genetic changes are not detectable by the analyzed markers. We argue that a parsimonious explanation for the observed pattern along the east-west axis in northern Spain may be competition between invading and resident populations, with no need to invoke selection. However, in order to conclusively rule out selective processes, additional and finer scale data are required to test for asymmetric mating preference/behaviour, sex-biased gene flow, or sex-biased survival of potential hybrids.