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Evolutionary processes behind lineage divergence often involve multidimensional differentiation. However, in the context of recent divergences, the signals exhibited by each dimension may not converge. In such scenarios, incomplete lineage sorting, gene flow, and scarce phenotypic differentiation are pervasive. Here, we integrated genomic (RAD loci of 90 individuals), phenotypic (linear and geometric traits of 823 and 411 individuals, respectively), spatial, and climatic data to reconstruct the evolutionary history of a speciation continuum of liolaemid lizards (Liolaemus kingii group). Specifically, we (a) inferred the population structure of the group and contrasted it with the phenotypic variability; (b) assessed the role of postdivergence gene flow in shaping phylogeographic and phenotypic patterns; and (c) explored ecogeographic drivers of diversification across time and space. We inferred eight genomic clusters exhibiting leaky genetic borders coincident with geographic transitions. We also found evidence of postdivergence gene flow resulting in transgressive phenotypic evolution in one species. Predicted ancestral niches unveiled suitable areas in southern and eastern Patagonia during glacial and interglacial periods. Our study underscores integrating different data and model-based approaches to determine the underlying causes of diversification, a challenge faced in the study of recently diverged groups. We also highlight Liolaemus as a model system for phylogeographic and broader evolutionary studies.

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We describe Liolaemus attenboroughi sp. nov., a lizard distributed in the northwestern Patagonian Steppe of Chubut province (Argentina) previously confused with L. kingii (Bell 1843). Recent studies based on molecular evidence supports its evolutionary independence. Here we provide a morphological diagnosis of this lineage, comparisons between three molecular species delimitation methods, and an updated phylogeny of the L. kingii group. Based on current knowledge of its distribution, this new species is allopatric with geographically close species of the L. kingii group.

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In this study we detangled the evolutionary history of the Patagonian lizard clade Liolaemus kingii, coupling dense geographic sampling and novel computational analytical approaches. We analyzed nuclear and mitochondrial data (restriction site-associated DNA sequencing and cytochrome b) to hypothesize and evaluate species limits, phylogenetic relationships, and demographic histories. We complemented these analyses with posterior predictive simulations to assess the fit of the genomic data to the multispecies coalescent model. We also employed a novel approach to time-calibrate a phylogenetic network. Our results show several instances of mito-nuclear discordance and consistent support for a reticulated history, supporting the view that the complex evolutionary history of the kingii clade is characterized by extensive gene flow and rapid diversification events. We discuss our findings in the contexts of the "gray zone" of speciation, phylogeographic patterns in the Patagonian region, and taxonomic outcomes. [Model adequacy; multispecies coalescent; multispecies network coalescent; phylogenomics; species delimitation.].

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MOTIVATION: Transposable elements (TEs) are ubiquitous in genomes and many remain active. TEs comprise an important fraction of the transcriptomes with potential effects on the host genome, either by generating deleterious mutations or promoting evolutionary novelties. However, their functional study is limited by the difficulty in their identification and quantification, particularly in non-model organisms. RESULTS: We developed a new pipeline [explore active transposable elements (ExplorATE)] implemented in R and bash that allows the quantification of active TEs in both model and non-model organisms. ExplorATE creates TE-specific indexes and uses the Selective Alignment (SA) to filter out co-transcribed transposons within genes based on alignment scores. Moreover, our software incorporates a Wicker-like criteria to refine a set of target TEs and avoid spurious mapping. Based on simulated and real data, we show that the SA strategy adopted by ExplorATE achieved better estimates of non-co-transcribed elements than other available alignment-based or mapping-based software. ExplorATE results showed high congruence with alignment-based tools with and without a reference genome, yet ExplorATE required less execution time. Likewise, ExplorATE expands and complements most previous TE analyses by incorporating the co-transcription and multi-mapping effects during quantification, and provides a seamless integration with other downstream tools within the R environment. AVAILABILITY AND IMPLEMENTATION: Source code is available at https://github.com/FemeniasM/ExplorATEproject and https://github.com/FemeniasM/ExplorATE_shell_script. Data available on request. SUPPLEMENTARY INFORMATION: Supplementary data are available at Bioinformatics online.

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Understanding the factors that cause heterogeneity among gene trees can increase the accuracy of species trees. Discordant signals across the genome are commonly produced by incomplete lineage sorting (ILS) and introgression, which in turn can result in reticulate evolution. Species tree inference using the multispecies coalescent is designed to deal with ILS and is robust to low levels of introgression, but extensive introgression violates the fundamental assumption that relationships are strictly bifurcating. In this study, we explore the phylogenomics of the iconic Liolaemus subgenus of South American lizards, a group of over 100 species mostly distributed in and around the Andes mountains. Using mitochondrial DNA (mtDNA) and genome-wide restriction site-associated DNA sequencing (RADseq; nDNA hereafter), we inferred a time-calibrated mtDNA gene tree, nDNA species trees, and phylogenetic networks. We found high levels of discordance between mtDNA and nDNA, which we attribute in part to extensive ILS resulting from rapid diversification. These data also reveal extensive and deep introgression, which combined with rapid diversification, explain the high level of phylogenetic discordance. We discuss these findings in the context of Andean orogeny and glacial cycles that fragmented, expanded, and contracted species distributions. Finally, we use the new phylogeny to resolve long-standing taxonomic issues in one of the most studied lizard groups in the New World.[Andes; ddRADSeq; introgression; lizards; mtDNA; reptiles; SNPs.].

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A new species of the Liolaemus capillitas clade is described. Liolaemus galactostictos sp. nov. differs from other members of its group by a combination of morphological and molecular traits, in particular its black dorsal coloration pattern not found in any other Liolaemus species. Liolaemus galactostictos sp. nov. is only known from its type locality. This new species is found in rocky fields surrounded by grasslands on the top of the Velasco Mountains, a ¨sky island environment¨, in northwestern Argentina. As well as other members of its clade this species seems to be strictly saxicolous, viviparous and feeds on insects.

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Liolaemus duellmani is a microendemic lizard described from southern Mendoza, Argentina, representing the southernmost distribution of the Liolaemus montanus species group. The original description was based on a small batch of preserved material and a juvenile collected alive. Along the years, this poorly known species was rarely collected or observed in the field, with no additional reports or findings published until a recent description of an adult female which was considered as a rediscovery of the species. Here, we redescribe the species and describe variation on external morphology, coloration, and some natural history observations.

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The clade Leiosaurae is composed of poorly-known species endemic to the southern region of South America. The difficulties of finding these lizards in the field, and their highly conserved morphology, have limited our taxonomic knowledge and understanding of their evolutionary histories. Here, we use data collected over 9 years to study the phylogenetic history, genetic diversity, and biogeographic history of almost all the southernmost species of Leiosaurae (except P. nigroigulus), including: Leiosaurus bellii, Diplolaemus darwinii, D. bibronii, D. sexcinctus and D. leopardinus. We use a fragment of the mitochondrial cytochrome-b gene to resolve general phylogeographic patterns, and add another mitochondrial gene and eight nuclear genes to perform species delimitation and phylogenetic analyses associated with divergence times. We found evidence for three putative new species-level taxa within L. bellii and five within Diplolaemus species, indicating high levels of geographic structure. We use a time-calibrated phylogeny to estimate ranges of ancestral distributions and to generate new hypotheses about their historical biogeography.

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No central online repository exists for the collection of animal images; hence it remains unclear how extensively species have been illustrated in the published literature or online. Here we compiled a list of more than 8000 reptile species (out of 11,341) that have photos in one of six popular online repositories, namely iNaturalist (6,349 species), the Reptile Database (5,144), Flickr (4,386), CalPhotos (3,071), Wikimedia (2,952), and Herpmapper (2,571). These sites have compiled over one million reptile photos, with some species represented by tens of thousands of images. Despite the number of images, many species have only one or a few images. This suggests that a considerable fraction of morphological and geographic variation is under documented or difficult to access. We highlight prominent gaps in amphisbaenians, lizards, and snakes, with geographic hotspots for species without images in Central Africa, Pacific Islands, and the Andes Mountains. We present a list of ~3,000 species without photos in any of the six databases and ask the community to fill the gaps by depositing images on one of these sites (preferably with minimal copyright restrictions).

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Coalescent-based algorithms coupled with the access to genome-wide data have become powerful tools for assessing questions on recent or rapid diversification, as well as delineating species boundaries in the absence of reciprocal monophyly. In southern South America, the diversification of Liolaemus lizards during the Pleistocene is well documented and has been attributed to the climatic changes that characterized this recent period of time. Past climatic changes had harsh effects at extreme latitudes, including Patagonia, but habitat changes at intermediate latitudes of South America have also been recorded, including expansion of sand fields over northern Patagonia and Pampas). In this work, we apply a coalescent-based approach to study the diversification of the Liolaemus wiegmannii species complex, a morphologically conservative clade that inhabits sandy soils across northwest and south-central Argentina, and the south shores of Uruguay. Using four standard sequence markers (mitochondrial DNA and three nuclear loci) along with ddRADseq data we inferred species limits and a time-calibrated species tree for the L. wiegmannii complex in order to evaluate the influence of Quaternary sand expansion/retraction cycles on diversification. We also evaluated the evolutionary independence of the recently described L. gardeli and inferred its phylogenetic position relative to L. wiegmannii. We find strong evidence for six allopatric candidate species within L. wiegmannii, which diversified during the Pleistocene. The Great Patagonian Glaciation (∼1 million years before present) likely split the species complex into two main groups: one composed of lineages associated with sub-Andean sedimentary formations, and the other mostly related to sand fields in the Pampas and northern Patagonia. We hypothesize that early speciation within L. wiegmannii was influenced by the expansion of sand dunes throughout central Argentina and Pampas. Finally, L. gardeli is supported as a distinct lineage nested within the L. wiegmannii complex.

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Hybridization is likely to occur more often between closely related taxa that have had insufficient time to diverge to the point of reproductive incompatibility; hybridization between deeply divergent lineages is rare. In squamate reptiles, hybridization has been proposed as a possible explanation for the extensive paraphyly observed in mitochondrial gene trees in several species complexes of the South American lizard genus Liolaemus. One of the best-documented cases is within the L. boulengeri and L. rothi complexes, which diverged ~5.5 million years ago. Here, we describe a comprehensive study for approaching the hybridization hypothesis between these lizard species complexes. We explored the level of gene tree discordance using the novel 'extra lineage contribution' statistics (XLC, presented in this study) that quantifies the level of gene tree discordance contribution per individual within a species. We included molecular data (12 nuclear and two mitochondrial genes) from 127 individuals, and results of a coalescent model-based analysis show that the most likely explanation for the gene tree-species tree discordance is interspecific hybridization. Our best-supported hypothesis suggests current and past hybridization between L. rothi (rothi complex) and L. tehuelche (boulengeri complex), and independently between L. rothi and L. boulengeri and L. telsen (boulengeri complex). The hybrid descendants are characterized by intermediate phenotypes between the parental species, but are more similar to L. rothi in body size. We discuss the possible role of hybridization in Liolaemus evolution.

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Two new species of the Liolaemus donosobarrosi clade are described. Liolaemus tirantii sp. nov. and Liolaemus calliston sp. nov. differ from other members of their clade by a combination of coloration characters, morphometric and molecular traits. Liolaemus tirantii sp. nov. is known from three localities separated only a few kilometers from each other and Liolaemus calliston sp. nov. is known only from the type locality. Both species inhabit a region strongly impacted by oil and gas extraction but their conservation status is unknown.

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Homonota is a Neotropical genus of nocturnal lizards characterized by the following combination of characters: absence of femoral pores, infradigital lamellae not dilated, claws without sheath, inferior lamellae laterally not denticulate, and presence of a ceratobranchial groove. Currently the genus is composed of 10 species assembled in three groups: two groups with four species, and the fasciata group with only two species. Here, we analyzed genetic and morphologic data of samples of Homonota fasciata from Paraguay; according to Maximum Likelihood and Bayesian inference analyses, the Paraguay population represents an undescribed species. Additionally, morphological analysis of the holotype of H. fasciata (MNHN 6756) shows that it is morphologically different from the banded, large-scaled Homonota commonly referred to as "H. fasciata". Given the inconsistency between morphological characters of the name-bearing type of H. fasciata and the species commonly referred to as H. fasciata, we consider them as different taxa. Thus, H. fasciata is a species inquirenda which needs further studies, and we resurrect the name H. horrida for the banded, large-scaled Homonota. The undescribed species from Paraguay is similar to H. horrida, but can be differentiated by the high position of the auditory meatus relative to the mouth commissure (vs. low position in H. horrida); and less developed tubercles on the sides of the head, including a narrow area between the orbit and the auditory meatus covered with small granular scales with or without few tubercles (vs. several big tubercles on the sides of the head even in the area between the orbit and the auditory meatus). The new species is distributed in the Dry Chaco in South America. With the formal description of this species, the actual diversity of the genus Homonota is increased to 12 species. Furthermore, we infer phylogenetic relationships for 11 of the 12 described species of the genus, based on 11 molecular markers (two mitochondrial and nine nuclear genes), with concatenated and species tree approaches.

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The consequences of global climate change can already be seen in many physical and biological systems and these effects could change the distribution of suitable areas for a wide variety of organisms to the middle of this century. We analyzed the current habitat use and we projected the suitable area of present conditions into the geographical space of future scenarios (2050), to assess and quantify whether future climate change would affect the distribution and size of suitable environments in two Pristidactylus lizard species. Comparing the habitat use and future forecasts of the two studied species, P. achalensis showed a more restricted use of available resource units (RUs) and a moderate reduction of the potential future area. On the contrary, P. nigroiugulus uses more available RUs and has a considerable area decrease for both future scenarios. These results suggest that both species have a moderately different trend towards reducing available area of suitable habitats, the persistent localities for both 2050 CO2 concentration models, and in the available RUs used. We discussed the relation between size and use of the current habitat, changes in future projections along with the protected areas from present-future and the usefulness of these results in conservation plans. This work illustrates how ectothermic organisms might have to face major changes in their availability suitable areas as a consequence of the effect of future climate change.

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The integrative taxonomy framework allows developing robust hypotheses of species limits based on the integration of results from different data sets and analytical methods. In this work, we test a candidate species hypothesis previously suggested based on molecular data, with geometric and traditional morphometrics analyses (multivariate and univariate). This new lizard species is part of the Phymaturus patagonicus group (payuniae clade) that is distributed in Neuquén and Mendoza provinces (Argentina). Our results showed that Phymaturus rahuensis sp. nov. differs from the other species of the payuniae clade by a higher number of midbody scales, and fewer supralabials scales, finger lamellae and toe lamellae. Also, its multidimensional spaces, both based on continuous lineal variables and geometric morphometrics (shape) characters, do not overlap with those of the other species in this clade. The results of the morphometric and geometric morphometric analyses presented here, coupled with previously published molecular data, represent three independent lines of evidence that support the diagnosis of this new taxon.

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The lizard genus Teius is composed by three species: Teius teyou, T. oculatus, and T. suquiensis and is distributed in South America, east of Andes. Teius teyou and T. oculatus have wide parapatric distributions with contact zones. Teius suquiensis is present in a small range along a sympatric area of the former species. In this work we analyze the distribution of the three species of Teius across its whole geographic range, examining its relationships with climatic parameters. We based our analysis on specimens in collections and literature records. Our analysis shows that the genus Teius is distributed from central Bolivia southwards to north of Río Negro Province in northern Patagonia, Argentina. Teius teyou reaches the northernmost range limit of the genus whereas T. oculatus occupies the southernmost limit. Teius oculatus is related to open and moist environments whereas T. teyou is more adapted to xeric and forested areas. Teius suquiensis is present in xerophytic areas of Dry Chaco and Espinal. Climatic factors in the distribution of the distribution of the two widespread species show marked differences and seasonality.

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An accurate estimation of species and population geographic ranges is essential for species-focused studies and conservation and management plans. Knowledge of the geographic distributions of reptiles from Patagonian Argentina is in general limited and dispersed over manuscripts from a wide variety of topics. We completed an extensive review of reptile species of central Patagonia (Argentina) based on information from a wide variety of sources. We compiled and checked geographic distribution records from published literature and museum records, including extensive new data from the LJAMM-CNP (CENPAT-CONICET) herpetological collection. Our results show that there are 52 taxa recorded for this region and the highest species richness was seen in the families Liolaemidae and Dipsadidae with 31 and 10 species, respectively. The Patagónica was the phytogeographic province most diverse in species and Phymaturus was the genus of conservation concern most strongly associated with it. We present a detailed species list with geographical information, richness species, diversity analyses with comparisons across phytogeographical provinces, conservation status, taxonomic comments and distribution maps for all of these taxa.

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A new species of the Andean-Patagonian Liolaemus elongatus clade is described. Liolaemus crandalli sp. nov. differs from other members of its clade by a combination of coloration characters, scale counts and genetic traits. Liolaemus crandalli sp. nov. is known only from an isolated volcanic mountain in northwestern Patagonia above 1500 m.a.s.l. unconnected with other habitat suitable for species of the Liolaemus elongatus clade.

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A new species of lizard of the saxicolous and viviparous genus Phymaturus from Patagonia is described. The new species is a member of the Phymaturus patagonicus species group distributed on volcanic plateaus, mountains or rocky outcrops of southern Argentina. Phymaturus yachanana sp. nov. differs from the other members of the patagonicus group by a dorsal pattern of small and medium-sized white spots on a brown background, the number of scales around the midbody, genetic differences in mitochondrial and nuclear markers, and allopatric geographic distribution in an isolated group of hills. As other species of the group, the new species is viviparous and herbivorous. 

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Until recently, most phylogeographic approaches have been unable to distinguish between demographic and range expansion processes, making it difficult to test for the possibility of range expansion without population growth and vice versa. In this study, we applied a Bayesian phylogeographic approach to reconstruct both demographic and range expansion in the lizard Liolaemus darwinii of the Monte Desert in Central Argentina, during the Late Quaternary. Based on analysis of 14 anonymous nuclear loci and the cytochrome b mitochondrial DNA gene, we detected signals of demographic expansion starting at ~55 ka based on Bayesian Skyline and Skyride Plots. In contrast, Bayesian relaxed models of spatial diffusion suggested that range expansion occurred only between ~95 and 55 ka, and more recently, diffusion rates were very low during demographic expansion. The possibility of population growth without substantial range expansion could account for the shared patterns of demographic expansion during the Last Glacial Maxima (OIS 2 and 4) in fish, small mammals and other lizards of the Monte Desert. We found substantial variation in diffusion rates over time, and very high rates during the range expansion phase, consistent with a rapidly advancing expansion front towards the southeast shown by palaeo-distribution models. Furthermore, the estimated diffusion rates are congruent with observed dispersal rates of lizards in field conditions and therefore provide additional confidence to the temporal scale of inferred phylogeographic patterns. Our study highlights how the integration of phylogeography with palaeo-distribution models can shed light on both demographic and range expansion processes and their potential causes.

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