Lizard Genus Research Articles

Testing relationships between multiple regional features and biogeographic processes of speciation, extinction, and dispersal.

The spatial and environmental features of regions where clades are evolving are expected to impact biogeographic processes such as speciation, extinction, and dispersal. Any number of regional features (such as elevation, distance, area, etc.) may be directly or indirectly related to these processes. For example, it may be that distances or differences in elevation or both may limit dispersal rates. However, it is difficult to disentangle which features are most strongly related to rates of different processes. Here, we present an extensible Multi-feature Feature-Informed GeoSSE (MultiFIG) model that allows for the simultaneous investigation of any number of regional features. MultiFIG provides a conceptual framework for incorporating large numbers of features of different types, including categorical, quantitative, within-region, and between-region features, along with a mathematical framework for translating those features into biogeographic rates for statistical hypothesis testing. Using traditional Bayesian parameter estimation and reversible-jump Markov chain Monte Carlo, MultiFIG allows for the exploration of models with different numbers and combinations of feature-effect parameters, and generates estimates for the strengths of relationships between each regional feature and core process. We validate this model with a simulation study covering a range of scenarios with different numbers of regions, tree sizes, and feature values. We also demonstrate the application of MultiFIG with an empirical case study of the South American lizard genus Liolaemus, investigating sixteen regional features related to area, distance, and elevation. Our results show two important feature-process relationships: a negative distance/dispersal relationship, and a negative area/extinction relationship. Interestingly, although speciation rates were found to be higher in Andean versus non-Andean regions, the model did not assign significance to Andean- or elevation-related parameters. These results highlight the need to consider multiple regional features in biogeographic hypothesis testing.

Do we have the necessary insights to classify species according to their conservation status? A case study in the world’s southernmost lizard genus

Endemic species generally have higher risk of extinction; therefore, it is necessary to focus conservation actions on these groups. Tools such as the Red List, through its criteria (A, B, C, D and E) allow to establish extinction risk and a conservation status for the species. Liolaemus is a highly diversified genus of lizards from southern South America. In Chile, 64% of the species within this genus are endemic, and more than 25% are threatened. However, for most of Liolaemus we have little biological information. In this study, to assess whether the current insights are sufficient to establish the conservation statuses of these species and to determine the criteria on which the current conservation statuses are based, a bibliographic review of all available information was conducted for the Chilean endemic Liolaemus species. Our results show that the most-used criteria were those of distribution (B and D). However, abundance studies were poorly represented. Species with wider distributions present the highest number of studies and more restricted species are little studied. Finally, many evaluations are based on the expert’s opinion. We conclude that further current and temporal studies of abundance and distribution, key aspects of ecology to assess species, are essential to generate an adequate conservation status of the genus.

Review of the Slender Geckos (Squamata: Gekkonidae: Hemiphyllodactylus) of the Cardamom Mountains of southwestern Cambodia, with descriptions of two new species and one country record

The gekkonid lizard genus Hemiphyllodactylus contains small, secretive species that are often overlooked in their habitats. Recent molecular and morphological analyses have resulted in the descriptions of numerous new species of Hemiphyllodactylus in mainland Southeast Asia and southern China, many of which were historically hidden within the geographically widespread taxon H. yunnanensis (Boulenger, 1903). Little is known on species diversity and distributions of Hemiphyllodactylus in Cambodia, and all previous reports from the country have been limited to provisional identifications owing to collections containing only a single female or juvenile specimen. We report on new collections of Hemiphyllodactylus specimens from opposite ends of the Cardamom Mountain range in southwestern Cambodia. Morphological and molecular data support the hypothesis that these collections contain three distinct species, two of which are described as new species and one of which is assigned to the geographically widespread H. typus Bleeker, 1860 as a new country record for Cambodia. Hemiphyllodactylus bokor sp. nov. from Preah Monivong Bokor National Park in Kampot Province and Hemiphyllodactylus samkos sp. nov. from Phnom Samkos Wildlife Sanctuary in Pursat Province are related to species from adjacent areas in southern Vietnam, southern Laos, and southeastern Thailand, as well as a clade from India, but are readily differentiated in mitochondrial DNA and in having unique combinations of mensural and meristic morphological characters. Additional species diversity of Hemiphyllodactylus likely exists in Cambodia.

Testing for heterogeneous rates of discrete character evolution on phylogenies

Abstract Many hypotheses in the field of phylogenetic comparative biology involve specific changes in the rate or process of trait evolution. This is particularly true of approaches designed to connect macroevolutionary pattern to microevolutionary process. We present a method to test whether the rate of evolution of a discrete character has changed in one or more clades, lineages, or time periods. This method differs from other related approaches (such as the “covarion” model) in that the “regimes” in which the rate or process is postulated to have changed are specified a priori by the user, rather than inferred from the data. Similarly, it differs from methods designed to model a correlation between two binary traits in that the regimes mapped onto the tree are fixed. We apply our method to investigate the rate of dewlap colour and/or caudal vertebra number evolution in Caribbean and mainland clades of the diverse lizard genus Anolis. We find little evidence to support any difference in the evolutionary process between mainland and island evolution for either character. We also examine the statistical properties of the method more generally and show that it has acceptable type I error, parameter estimation, and power. Finally, we discuss some general issues of frequentist hypothesis testing and model adequacy, as well as the relationship of our method to existing models of heterogeneity in the rate of discrete character evolution on phylogenies.

Molecular phylogenetic and historical biogeographical relationships of Laudakia (Squamata: Agamidae) and intraspecific differentiation of L. stoliczkana inferred from mitochondrial DNA sequences

The rock lizard genus Laudakia is representative agamid species from the arid zone, and its genus division has not been resolved yet. Laudakia stoliczkana, which occurs in both Xinjiang, China, and the Gobi Altai, Mongolia, is divided into two subspecies, Laudakia stoliczkana stoliczkana and Laudakia stoliczkana altaica, based on morphological differences, but little is known about the molecular genetic differences between the two subspecies. This study reconstructs the phylogenetic tree of Laudakia and analyses molecular differences between two subspecies of L. stoliczkana by DNA barcoding (COI and 16S). Our results show that: (1) Laudakia is monophyletic and the phylogenetic tree is broadly divided into three main branches, namely branch A (L. caucasia and L. stoliczkana), which occurs mainly in Central Asia and the Gobi Altai region to the north, branch B (L. stellio), which occurs in the Middle East, and branch C (L. tuberculata, L. papenfussi, L. himalayana, L. wui, L. stellio), which occurs mainly near the Himalayas; (2) The biogeographic analysis of Laudakia suggests that the genus probably originated at 43.72 Ma (95% confidence interval HPD: 23.53–66.12Ma) and is associated with the uplift of the Tibetan Plateau and the aridification of Central Asias subsequently; (3) Molecular genetic distances and morphological differences support the delimitation of the two subspecies of L. stoliczkana, with divergence between the two subspecies estimated to have occurred at 3.27 Ma (95% confidence interval HPD: 1.58–5.87Ma), in associated with the recent uplift of the Tian Shan Mountains. The results highlight the importance of the uplift of the Central Asian mountains and the Tibetan Plateau for the divergence of Laudakia.

Integrative taxonomy reveals two new species of karst-dwelling Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from the border region of Laos and Vietnam

An integrative taxonomic analysis of the gekkonid lizard genus Hemiphyllodactylus was conducted using molecular, morphological, color pattern, and ecological data on a clade of 19 species, including two new species from karst landscapes in northern Laos and Vietnam. Hemiphyllodactylus houaphanensis sp. nov. (Laos) and H. vanhoensis sp. nov. (Vietnam) are karst-dwelling species whose discovery further emphasizes the unrealized herpetological diversity and site-specific endemism in karst ecosystems across Southeast Asia and the need for increased field work to protect these biodiverse landscapes.

Environmental and anthropogenic factors mediating the functional connectivity of the mesquite lizard along the eastern Trans-Mexican Volcanic Belt.

Functional connectivity, the extent to which a landscape facilitates or impedes the dispersal of individuals across the landscape, is a key factor for the survival of species. Anthropogenic activities, such as urbanization, agriculture and roads, negatively impact functional connectivity of most species, particularly low-vagility species like lizards. Here, we examine how a landscape modified by anthropogenic activities affects the functional connectivity, at both broad and fine scales, of a widely distributed generalist lizard Sceloporus grammicus in the eastern Trans-Mexican Volcanic Belt, Mexico. We estimated for the first time the species' genetic structure, gene flow and functional connectivity in agricultural and forest zones using genomic data, a comprehensive landscape characterization and novel methods including gravity models. Our results showed not only marked genetic differentiation across the study region but also that functional connectivity is maintained for tens of kilometres despite S. grammicus low vagility. Specifically, we found that substrate and air temperature facilitated connectivity over broad and fine scales, respectively, while agricultural cover, relative humidity and slope were important for connectivity and gene flow. Contrastingly, forest cover and roads favoured (broad-scale) and limited (fine-scale) connectivity, likely associated with movement facilitated by small forest patches and with thermoregulation. Altogether, these results support that S. grammicus alternates its thermoregulatory behaviour depending on the distance travelled and the habitat environmental conditions, and that it can disperse through relatively modified landscapes, mainly using agricultural zones. The information obtained is crucial to understanding the response of lizards to current anthropogenic pressures and their potential to adapt.

Flexibility in thermal requirements: a comparative analysis of the wide-spread lizard genus Sceloporus.

Adaptation or acclimation of thermal requirements to environmental conditions can reduce thermoregulation costs and increase fitness, especially in ectotherms, which rely heavily on environmental temperatures for thermoregulation. Insight into how thermal niches have shaped thermal requirements across evolutionary history may help predict the survival of species during climate change. The lizard genus Sceloporus has a widespread distribution and inhabits an ample variety of habitats. We evaluated the effects of geographical gradients (i.e. elevation and latitude) and local environmental temperatures on thermal requirements (i.e. preferred body temperature, active body temperature in the field, and critical thermal limits) of Sceloporus species using published and field-collected data and performing phylogenetic comparative analyses. To contrast macro- and micro-evolutional patterns, we also performed intra-specific analyses when sufficient reports existed for a species. We found that preferred body temperature increased with elevation, whereas body temperature in the field decreased with elevation and increased with local environmental temperatures. Critical thermal limits were not related to the geographic gradient or environmental temperatures. The apparent lack of relation of thermal requirements to geographic gradient may increase vulnerability to extinction due to climate change. However, local and temporal variations in thermal landscape determine thermoregulation opportunities and may not be well represented by geographic gradient and mean environmental temperatures. Results showed that Sceloporus lizards are excellent thermoregulators, have wide thermal tolerance ranges, and the preferred temperature was labile. Our results suggest that Sceloporus lizards can adjust to different thermal landscapes, highlighting opportunities for continuous survival in changing thermal environments.

On Segurasaurus (Squamata: Pythonomorpha), a New Genus of Lizard from the Cenomanian (Upper Cretaceous) of Portugal

Carentonosaurus soaresi was recently described in the uppermost middle Cenomanian (Upper Cretaceous) of Casais dos Carecos (Coimbra, western Portugal) based on a diverse set of new material (cervical and dorsal vertebrae) of the Pythonomorpha lizard. The main morphological characteristics observed in the vertebrae used for the diagnosis of this species are the presence of distinct lateral and subcentral foramina, highly laterally projected paradiapophyses beyond the prezygapophyses, a low subrectangular neural spine ornamented with longitudinal grooves, and dorsal vertebrae displaying a sagittal furrow along the ventral surface. Additional diagnostic details observed both in new material and the previously studied vertebrae are described herein, such as the presence of keels in the zygantrum and zygosphene. These and other important morphological characteristics present in the species soaresi are absent in the genotype Carentonosaurus mineaui and in other known Squamata, allowing for the definition of the new genus Segurasaurus.

Hematology of Liolaemus pacha (Iguania: Liolaemidae) and its relationship with mite infestation, reproductive period and body condition.

Variations in hematological profile in reptiles can be caused by multiple factors, including parasites presence. Our goals were to identify and morphologically describe blood cells of Liolaemus pacha and analyze their relationship with sex, body condition, individual reproductive/post-reproductive period and mite infestation. Blood smear analyses do not indicate the presence of hemoparasites, suggesting that the mites Neopterygosoma do not serve as vectors for these organisms, as has been proposed for other genera of ectoparasitic mites. In post-reproductive period, there was a reduction in specimens' body condition and a higher leukocyte count in uninfected lizards. This could be a consequence of the testosterone effects, in higher concentration during the reproductive season, which can increase the metabolic rate, decreasing feeding rate. Infested and non-infested lizards showed no differences in body condition, as well as in leukocyte count, hence the host's immune system could be developing infestation tolerance. Infested specimens had a higher count of monocytes, thrombocytes, heterophils and lymphocytes. Based on cells function, mites' effect could be associated with inflammatory processes, allergic reactions or infectious diseases. These results suggested a complex interaction between lizards' hematological parameters and factors associated to ectoparasites or body conditions. We consider this work as a diagnostic tool for genus Liolaemus, to evaluate health quality, with relevance to the conservation or management of this lizard's genus.

Comparative approaches to understanding speciation: a case study in Xantusia lizards

Abstract The diversity of life forms that surround us formed through a variety of processes. Speciation researchers use a suite of comparative approaches to understand both the generalities and the variance underpinning this diversification. Here, we summarize these approaches and what they have taught us, and we then apply them to understand speciation in the night lizards Xantusia, a genus of small lizards found in the arid regions of western North America. Although Xantusia species span both a range of phenotypic and genetic divergence, we find that speciation in Xantusia appears to be a predictable consequence of divergence in extended periods of allopatry. We conclude by identifying possible areas of growth for comparative studies of speciation.

Islands, saurians and parasites

A review and discussion of the results obtained after more than 40 years of research on parasites of insular saurian reptiles in some areas of the Mediterranean and the Atlantic has been carried out. Twenty-seven species and subspecies belonging to seven different genera of lizards were analysed, revealing the existence of diverse helminth communities, related to different biotic and abiotic characteristics of the hosts and habitats. The biogeographical and evolutionary particularities of the endemic parasites found are detailed and commented, as well as the peculiarities of the helminth fauna of the hosts in relation to their type of feeding or their age and size. The capacity and opportunity of saurian hosts in the dispersal of parasite species and the importance of parasites as bioindicators are also analysed.

Single hybrid population but multiple parental individuals at the origin of parthenogenetic rock lizards Darevskia sapphirina and D. bendimahiensis Schmidtler, & Eiselt Darevsky (1994) endemic to the area of Lake Van in East Turkey

Among vertebrates, obligate parthenogenesis is only found in Squamata, where it always has a hybrid origin and a few lizard genera contain most of the known hybridogenous parthenogenetic taxa. Parthenogenesis thus seems to be pre-conditioned at the genus level, but it is not clear how often the encounter between two parental sexually reproducing species can result in the parthenogenetic offspring, nor whether the success of such hybridization event requires certain conditions or the specific time frame. To address this question, we studied the rock lizards of genus Darevskia, where a pair of parental species, D. valentini and D. raddei, as well as their parthenogenetic daughter species D. bendimahiensis and D. sapphirina, are found in close proximity NE of the Lake Van in East Anatolia. Using ddRAD-seq genotyping on 19 parental and 18 hybrid individuals, we found that (i) all parthenogenetic individuals from both D. bendimahiensis and D. sapphirina have a monophyletic origin tracing back to a single initial hybrid population, but their current genetic variation is geographically structured; (ii) unlike the most probable paternal ancestor, the genetically closest extant population of the maternal ancestor is not the geographically nearest one; and (iii) in the parthenogens, about 1% of loci carry multiple haplotypes, frequently differentiated by multiple substitutions. This pattern, in addition to biases in the relative frequency of haplotypes of maternal and paternal origin, does not appear compatible with a scenario of the entire parthenogenic clonal population having descended from a single pair of parental individuals. Instead, the data suggest that multiple parental individual ancestries still persist in the parthenogenetic gene pool. This supports the notion that although hybridization leading to parthenogenesis is generally rare at the level of species, it may be more common at the individual/population level once the right conditions are met.

First fossil record of Varanus (Reptilia, Squamata) from Switzerland and the earliest occurrences of the genus in Europe

We here describe two isolated fossil teeth from the Early Miocene (MN 4) of Hüenerbach, Switzerland. The material is tentatively referred to Varanus, marking the first occurrence of this iconic lizard genus in the Swiss fossil record, while it also represents one of its earliest occurrences from Europe, being coeval with other records from Spain, France, and the Czech Republic. Important diagnostic features in the teeth of extant Varanus are discussed, coupled with comparisons with related taxa. The prominent serration in one of the two Swiss teeth is an unusual feature for European varanids. In addition, the sole other lizard specimen from Hüenerbach is also described, being referred to an indeterminate lacertid, with potential affinities to the genus Janosikia.

Intricate covariation between exploration and avoidance learning in a generalist predator

Abstract Many predators avoid unprofitable prey by learning to use visual features of the prey as reliable indicators of quality. However, individual variation in avoidance learning is rarely examined in detail. It has been hypothesized that better avoidance learning ability might correlate with faster exploration tendency, but available data are limited in both quantity and scope. In this study, we examined the covariation between exploration, foraging decisions, and avoidance learning in a generalist lizard Eutropis multifasciata to test the prediction that faster explorers are also better avoidance learners. We also examined how sex, population, and color of unpalatable prey might mediate the exploration-avoidance learning covariation. We collected data on exploration and foraging behavior in individuals from two allopatric populations and quantified changes in foraging decisions over five daily learning trials, in which individuals were presented with normal- and bitter-tasting prey that differed consistently in color. Even though bitter prey elicited strong negative responses, lizards overall did not avoid consuming fewer such prey with learning. Instead, they learned to prioritize on palatable prey as the experiment progressed. In concordance with our prediction, we found that faster explorers were generally better avoidance learners, even though sex, population, and prey color were also important. Our study represents a rare experimental test of the exploration-avoidance learning covariation, especially in non-avian systems. Our results suggest that unpalatability might be an ineffective defense against generalist predators such as E. multifasciata and that faster explorers might impose stronger selection for the evolution of warning signals in unprofitable prey.

Protection from overheating of simulated Sceloporus horridus lizards in a biosphere reserve of seasonally dry tropical forest in central Mexico

In seasonally dry ecosystems, loss of vegetation cover leads to warmer microclimates that can increase lizards’ body temperatures to the point of threatening their performance. Preserving vegetation by establishing protected areas may mitigate these effects. We used remote sensing to test these ideas in the Sierra de Huautla Biosphere Reserve (REBIOSH) and surrounding areas. First, we determined whether vegetation cover was higher in the REBIOSH compared to adjacent unprotected areas to the north (NAA) and south (SAA). Then, we used a mechanistic niche model to test whether simulated Sceloporus horridus lizards in the REBIOSH experienced a cooler microclimate, higher thermal safety margin, longer foraging duration, and lower basal metabolic rate compared to adjacent unprotected areas. We compared these variables between 1999, when the reserve was declared, and 2020. We found that vegetation cover increased from 1999 to 2020 in all three areas; it was higher in the REBIOSH than in the more anthropized NAA, and was intermediate in the less anthropized SAA in both years. The microclimate temperature decreased from 1999 to 2020 and was lower in the REBIOSH and SAA than in the NAA. Thermal safety margin increased from 1999 to 2020; it was higher in the REBIOSH than in the NAA and intermediate in the SAA. Foraging duration increased from 1999 to 2020 and was similar among the three polygons. Basal metabolic rate decreased from 1999 to 2020 and was higher in the NAA than in the REBIOSH and SAA. Our results suggest that the REBIOSH provides cooler microclimates that increase the thermal safety margin and lower the metabolic rate of this generalist lizard compared to the NAA, and that the REBIOSH could contribute to increased vegetation cover in its surroundings. Besides, protecting original vegetation cover is an important part of climate change mitigation strategies more generally.

Hiding on jagged karst pinnacles: A new microendemic genus and species of a limestone-dwelling agamid lizard (Squamata: Agamidae: Draconinae) from Khammouan Province, Laos.

We describe a unique new species and genus of agamid lizard from the karstic massifs of Khammouan Province, central Laos. Laodracon carsticola Gen. et sp. nov. is an elusive medium-sized lizard (maximum snout-vent length 101 mm) specifically adapted to life on limestone rocks and pinnacles. To assess the phylogenetic position of the new genus amongst other agamids, we generated DNA sequences from two mitochondrial gene fragments (16S rRNA and ND2) and three nuclear loci ( BDNF, RAG1 and c-mos), with a final alignment comprising 7 418 base pairs for 64 agamid species. Phylogenetic analyses unambiguously place the new genus in the mainland Asia subfamily Draconinae, where it forms a clade sister to the genus Diploderma from East Asia and the northern part of Southeast Asia. Morphologically, the new genus is distinguished from all other genera in Draconinae by possessing a notably swollen tail base with enlarged scales on its dorsal and ventral surfaces. Our work provides further evidence that limestone regions of Indochina represent unique "arks of biodiversity" and harbor numerous relict lineages. To date, Laodracon carsticola Gen. et sp. nov. is known from only two adult male specimens and its distribution seems to be restricted to a narrow limestone massif on the border of Khammouan and Bolikhamxai provinces of Laos. Additional studies are required to understand its life history, distribution, and conservation status.

How agricultural land use affects the abundance and prevalence of monoxenous and heteroxenous helminths in the generalist lizard Tropidurus hispidus.

Among the forms of anthropogenic disturbance, agricultural land use is one of the main threats to biodiversity. Understanding how interactions between parasites and hosts are affected by agricultural land use allows predictions of how these anthropogenic impacts affect parasites. Although parasitism patterns are affected by agricultural land use, it is noteworthy that different groups of parasites can respond differently to these environmental alterations. While heteroxenous species need more than one host to complete their life cycle and tend to be more harmed by anthropization, monoxenous species, which need only one host to complete their life cycle, tend to be less harmed. In this work, we evaluate how agricultural land use affects the abundance and prevalence of parasitism for monoxenous and heteroxenous helminths in the generalist lizard Tropidurus hispidus in Caatinga Domain, Brazil. We recorded differences in abundance and prevalence of heteroxeneous (higher in conserved areas) and monoxenous helminths (higher in agricultural areas). Heteroxenous helminths that have lizards as definitive hosts are mainly obtained through diet. Tropidurus hispidus predominantly consumes insects, so it is possible that the lower abundance and prevalence of heteroxenous parasites in agricultural areas, beyond habitat simplification, is related to the decrease in the insect population. As monoxenous species do not need an intermediate host, it is possible that this aspect has influenced their greater success in anthropogenic environments than heteroxenous species. This contrasting result reinforces the need for a separate assessment between these groups when evaluating effects of land use.

A Phylogenetic Assessment of Nematodes (Oxyuroidea: Pharyngodonidae) Infecting Moroccan Lizards.

Molecular tools can be used to estimate the phylogeny of species and to identify cryptic diversity, but their use for parasites has lagged behind that of free-ranging organisms. As an example, in North Africa there is minimal molecular data available for helminth parasites of lizards. In this work we used two molecular markers (the nuclear 18S rRNA and the mitochondrial Cytochrome c Oxidase subunit 1) to investigate the diversity of nematodes of the family Pharyngodonidae parasitizing three genera of lizards from Morocco (Chalcides, Quedenfeldtia and Tarentola) and to explore their co-evolutionary history. Morphological assessments indicated that members of three genera were present: Spauligodon, Thelandros, and Parapharyngodon. Phylogenetic analysis of 18S rRNA sequences indicated the monophyly of the genus Spauligodon, and that some lineages could be distinguished, including Spauligodon auziensis from the host species Tarentola mauritanica, and another unnamed lineage from hosts of the genus Chalcides. However, with this slow-evolving marker some species could not be distinguished. The genus Thelandros was not monophyletic, although relationships were not strongly supported. Analysis of the faster evolving mitochondrial marker clearly separated various species of Spauligodon, as well as distinct unnamed lineages identified in the host genus Chalcides and the host Quedenfeldtia moerens.

Diversification in the mountains: Evolutionary history and molecular phylogeny of Anatolian rock lizards

Mountains play a key role in forming biodiversity by acting both as barriers to gene flow among populations and as corridors for the migration of populations adapted to the conditions prevailing at high elevations. The Anatolian and the Zagros Mountains are located in the Alpine-Himalayan belt. The formation of these mountains has influenced the distribution and isolation of the animal population since the late Cenozoic. Apathya is a genus of lacertid lizards distributed along these mountains with two species, i.e., Apathya cappadocica and Apathya yassujica. The taxonomy status of lineages within the genus is complicated. In this study, we tried to collect extensive samples from throughout the distribution range, especially within the Zagros Mountains. Also, we used five genetic markers, two mitochondrial (COI and Cyt b) and three nuclear (C-mos, NKTR, and MCIR), to resolve the phylogenetic relationships within the genus and explain several possible scenarios that shaped multiple genetic structures. The combination of results in the current study indicated eight well-support monophyletic lineages that separated to two main groups; group 1 including A. c. cappadocica, A. c. muhtari and A. c. wolteri, group 2 contains four regional clades Turkey, Urmia, Baneh and Ilam, and finally a single clade belonging to the species A. yassujica. In contrast to previous studies, Apathya cappadocica urmiana was divided into four clades and three clades were recognized within Iranian boundaries. The clades have dispersed from Anatolia to adjacent regions in the south of Anatolia and the western Zagros Mountains. According to the evidence generated in this study this clade is paraphyletic. Based on our assumption, orogeny activities and also climate fluctuations in Middle Miocene and Pleistocene have influenced to formation of lineages. In this study we revisit the taxonomy of the genus and demonstrate that the species diversity was substantially underestimated. Our findings suggest that each of the eight clades corresponding to subspecies and distinct geographic regions deserve to be promoted to species level.