Gecko Species Research Articles

A new species of naked-toed gecko Cyrtopodion sensu lato (Squamata: Sauria: Gekkonidae) from Gujarat, India

We describe a new species of naked-toed gecko of the genus Cyrtopodion Fitzinger, 1843, from the Dahod and Panchmahal districts of Gujarat state, India. The new species is diagnosable by the following suite of characters: a medium-sized Cyrtopodion (adult, snout to vent length up to 50 mm); dorsal scalation on trunk granular, intermixed with enlarged, regularly arranged transverse rows of 15 trihedral tubercles; 6 transverse rows of tubercles on the second segment of the tail; midbody scale rows across belly 20-22; midventral scales 89-97; males with a continuous series of 29-33 precloacal-femoral pores. The new species, Cyrtopodion vindhya sp. nov. is the fifth endemic species of reptile described in the last 15 years from the state of Gujarat that highlights the rich and unique diversity of this understudied region.

A new species of Phyllurus leaf-tailed gecko (Lacertilia: Carphodactylidae) from Scawfell Island, mid-east Queensland, Australia

A recent targeted reptile survey of Scawfell Island, in the South Cumberland Group, revealed a species of Phyllurus gecko that could not be morphologically assigned to any described species. Here I describe this as a new species, Phyllurus fimbriatus sp. nov., based on differences in a number of morphometric and scalation traits from congeners. Phyllurus fimbriatus sp. nov. is restricted to deeply-piled boulder habitat under rainforest canopy on Scawfell Island, approximately 50 km offshore from Mackay in mid-east Queensland. A survey in rocky, rainforest habitat on nearby Carlisle Island failed to find the species, and other nearby islands appear to lack sufficiently deep rock outcropping to support the species. Phyllurus fimbriatus sp. nov. is known from two small patches of habitat on Scawfell Island, but it is common within these, and is likely to be found in other suitable habitat patches on the island. Based on assessment of imagery, the total area of habitat occupied may be < 1 km2. The island is protected within South Cumberland National Park, but fire encroachment from adjacent dry sclerophyll habitats, climate change, competition from introduced Asian House Geckos (Hemidactylus frenatus Duméril & Bibron, 1836), and poaching are potential threats.

Fossil Squamates of Pedernales Province, Dominican Republic: Novel Record of Human-Induced Extinction and Extirpation

Hispaniola (Haiti &amp; Dominican Republic (D.R.)), is the second largest island in the Caribbean and is a hotspot of squamate diversity (~184 species), yet little is known about their fossil record and how it relates to the present. Past studies on mammalian communities suggest that many rodent species go extinct after human arrival (~7,000 y.B.P.), most significantly after European colonization (~530 y.B.P.) corresponding to a rapid loss of 70-98% of original forest. Pedernales Province in the D.R., has much of its original forest cover, yet fossil squamates from this region have not been documented. Our study of Holocene dry cave fossil sites and a new archeological site located in Pedernales allows for evaluation of extinction in squamates in an area where deforestation is not a primary concern. Using dentigerous elements, several genera of squamates (Ameiva, Anolis, Celestus, Cyclura, Leiocephalus, and Geckos) were identified. We recorded their abundance at each depth, and found Anolis, Celestus, and Geckos to be the most common fossils. From this collection we discovered two new species of Celestus, a new species of Gecko, and revised the taxonomy of Leiocephalus. One of the newly identified large Celestus species, originally only known from fossils, was later discovered in a separate cave as a living specimen. These results suggest that deforestation is not the only driver of extinction in the D.R., and that even in well preserved areas, extinction of small taxa is happening. Additionally, some cryptic taxa may be unrecognized. The arrival of domesticated animals and associated pests (mice, rats, etc.), likely had a dramatic influence on these extinctions, as well as direct human predation, first reported here. Further radiocarbon and geochemical sampling of these herpetofaunal fossils will help assess why these species went extinct, and such trends can inform modern conservation efforts as to which taxa are most at risk.

Effect of habitat structure on abundance and body conditions of two sympatric geckos, Cyrtodactylus saiyok and Cyrtodactylus tigroides, in the karst forest of western Thailand

ABSTRACT Deforestation is a major threat to biodiversity worldwide, especially in Southeast Asia. In western Thailand, reptiles are an important part of tropical rainforest ecosystems and karst forests. However, basic ecological studies of many species are largely lacking. Therefore, we studied two sympatric gecko species, Cyrtodactylus saiyok and C. tigroides, that inhabit karst forests and are restricted to western Thailand. We investigated the habitat variables that affect their abundance and body condition in Kanchanaburi Province in western Thailand. We found that the abundance of C. saiyok increased with increasing plant density and decreasing canopy cover. For both study species, body condition was positively correlated with canopy cover. The two species differed in their microhabitat use, and C. tigroides prefers more karst outcrops and stays higher above the ground. Our findings highlight the importance of an intact forest structure for the survival of geckos and thus reiterates that it is crucial to conserve karst forests in Southeast Asia.

Scratching the surface: a new species of Bent-toed gecko (Squamata, Gekkonidae, Cyrtodactylus) from Timor-Leste of the darmandvillei group marks the potential for future discoveries.

A new species of limestone-dwelling Bent-toed gecko (genus Cyrtodactylus) is described from Nino Konis Santana National Park in the far-east region of Timor-Leste. Both genetic and morphological data strongly support the evolutionary distinctness of the new species, which we describe herein as Cyrtodactylussantana sp. nov. Phylogenetic analysis based on the ND2 mitochondrial gene inferred the new species as part of the C.darmandvillei group with close genetic affinities to C.batucolus, C.seribuatensis, C.petani, C.sadleiri, and two undescribed lineages from the Moluccas in Indonesia. The new species represents the first species of Cyrtodactylus identified at the species level from Timor-Leste and fills an important gap in our understanding of the biogeography and evolutionary history of Cyrtodactylus especially in the Wallacean region. Our results strongly suggest that the diversity of Cyrtodactylus in Wallacea is still underestimated and many more unnamed species remain to be described.

Cytogenetic Analysis of Seven Species of Gekkonid and Phyllodactylid Geckos.

Geckos (Gekkota), the species-rich clade of reptiles with more than 2200 currently recognized species, demonstrate a remarkable variability in diploid chromosome numbers (2n = 16-48) and mode of sex determination. However, only a small fraction of gekkotan species have been studied with cytogenetic methods. Here, we applied both conventional (karyotype reconstruction and C-banding) and molecular (fluorescence in situ hybridization with probes for rDNA loci and telomeric repeats) cytogenetic analyses in seven species of geckos, namely Blaesodactylus&nbsp;boivini, Chondrodactylus&nbsp;laevigatus, Gekko&nbsp;badenii, Gekko cf. lionotum, Hemidactylus&nbsp;sahgali, Homopholis&nbsp;wahlbergii (Gekkonidae) and Ptyodactylus&nbsp;togoensis (Phyllodactylidae), in order to provide further insights into the evolution of karyotypes in geckos. Our analysis revealed the presence of interstitial telomeric repeats in four species, but we were not able to conclude if they are remnants of previous chromosome rearrangements or were formed by an accumulation of telomeric-like satellite motifs. Even though sex chromosomes were previously identified in several species from the genera Hemidactylus and Gekko by cytogenetic and/or genomic methods, they were not detected by us in any examined species. Our examined species either have poorly differentiated sex chromosomes or, possibly, environmental sex determination. Future studies should explore the effect of temperature and conduct genome-wide analyses in order to identify the mode of sex determination in these species.

Madagascar Leaf-Tail Geckos (Uroplatus spp.) Share Independently Evolved Differentiated ZZ/ZW Sex Chromosomes.

Geckos are an excellent group to study the evolution of sex determination, as they possess a remarkable variability ranging from a complete absence of sex chromosomes to highly differentiated sex chromosomes. We explored sex determination in the Madagascar leaf-tail geckos of the genus Uroplatus. The cytogenetic analyses revealed highly heterochromatic W chromosomes in all three examined species (Uroplatus&nbsp;henkeli, U.&nbsp;alluaudi, U. sikorae). The comparative gene coverage analysis between sexes in U.&nbsp;henkeli uncovered an extensive Z-specific region, with a gene content shared with the chicken chromosomes 8, 20, 26 and 28. The genomic region homologous to chicken chromosome 28 has been independently co-opted for the role of sex chromosomes in several vertebrate lineages, including monitors, beaded lizards and monotremes, perhaps because it contains the amh gene, whose homologs were repeatedly recruited as a sex-determining locus. We demonstrate that all tested species of leaf-tail geckos share homologous sex chromosomes despite the differences in shape and size of their W chromosomes, which are not homologous to the sex chromosomes of other closely related genera. The rather old (at least 40 million years), highly differentiated sex chromosomes of Uroplatus geckos can serve as a great system to study the convergence of sex chromosomes evolved from the same genomic region.

A new species of spiny-tailed gecko (Squamata: Diplodactylidae: Strophurus) from the mulga woodlands of inland Western Australia

A new species of spiny-tailed gecko (Squamata: Diplodactylidae: Strophurus) from the mulga woodlands of inland Western Australia

New Morphological and Molecular Data Reveal an Underestimation of Species Diversity of Mites of the Genus Geckobia (Acariformes: Pterygosomatidae) in India

Mites of the genus Geckobia (Acariformes: Pterygosomatidae) are permanent and highly specialised ectoparasites of geckos (Gekkota). We conducted a local study on Geckobia mites associated with the geckos of the family Gekkonidae found mainly in the territory of the Indian Institute of Science’s campus (Bangalore, India). In total, we examined 208 lizards belonging to two genera: Hemidactylus and Cnemaspis. We assessed the prevalence of the mites and identified the preferred site for their infestation. We extended the standard morphological identification of the mite species by using DNA barcode markers, partial sequences of the mitochondrial cytochrome c oxidase subunit I (COI) gene and nuclear ribosomal gene sequences: 18S rRNA and hypervariable region D2 of nuclear 28S rRNA. We checked the suitability of COI and nuclear (D2 of 28S rRNA) markers for species delimitations and identification purposes of the genus. The distance- and phylogeny-based approaches were applied: (i) to test the presence of a barcoding gap, we used the automated barcoding gap discovery tool (ABGD) and investigated intra- and interspecific genetic distances, and (ii) to reconstruct evolutionary relationships within the species, we performed maximum likelihood (ML) and Bayesian inference with Markov-Chain Monte Carlo (BI) analyses. As a result, we described five new species—Geckobia gigantea sp. n., G. treutleri sp. n., G. unica sp. n. and G. brevicephala sp. n.—from four Hemidactylus species: H. giganteus, H. treutleri, H. parvimaculatus and H. frenatus, respectively, and G. mysoriensis sp. n. from Cnemaspis mysoriensis. Additionally, we found three already described species: Geckobia indica Hirst, 1917 on H. treutleri (new host), Geckobia bataviensis Vitzhum, 1926 on H. parvimaculatus (new host) and H. frenatus (new locality) and Geckobia phillipinensis Lawrence, 1953 on H. frenatus (new locality). The diagnoses of G. indica and G. phillipinensis were improved and supplemented by descriptions of the males and juveniles. Both topologies of the BI and ML phylogenetic trees, as well as genetic distances, supported the species boundaries in the mite population shown by the morphological data. Hemidactylus frenatus was the most infected gecko species (61% prevalence), with the highest number of mite species (three spp.). The scale-mite richness was higher than expected; therefore, further research is required to evaluate the true diversity of Geckobia mites.

A new species of feather-tailed leaf-toed gecko, Kolekanos Heinicke, Daza, Greenbaum, Jackman, Bauer, 2014 (Squamata, Gekkonidae) from the poorly explored savannah of western Angola.

We here describe a new species of feather-tailed leaf-toed gecko, Kolekanos, from southern Benguela Province, Angola, based on morphological and osteological evidence, supported by phylogenetic analysis of mitochondrial data. The new species adds to the rapidly growing and newly-recognised endemic biodiversity of Angola, doubling the number of Kolekanos species, breaking the pattern observed within other closely-related African members of a clade of circum-Indian Ocean leaf-toed geckos - Ramigekko, Cryptactites and Afrogecko - all of which are presently monotypic. The new species is easily distinguished from K.plumicaudus, based on spine-like (as opposed to feather-like) scales on the margins of the original tail. Phylogenetic analyses also recovered the new taxon as monophyletic, with a well-supported sister relationship to K.plumicaudus, from which it differs by a substantial 24.1% NADH-dehydrogenase subunit 2 mitochondrial gene uncorrected p-distance.

The mutual history of Schlegel's Japanese gecko (Reptilia: Squamata: Gekkonidae) and humans inscribed in genes and ancient literature.

Knowing how the present distribution of organisms was formed is an essential issue in evolutionary ecology. Recently, the distribution of organisms on Earth has been significantly changed by human-mediated dispersal due to globalization. Therefore, significant attention has been paid to such processes. However, although humankind has taken considerable time to achieve modernization, the impact of ancient human activity on ecosystems has not yet been thoroughly studied. We hypothesized that ancient urban development and transitions had a non-negligible effect on species distribution. Inferring the impact of past human activity on ecosystems from ancient literature and verifying that impact by genetic analysis and human history is an effective means of tackling this problem. As geckos, a popular neighbor of human dwellings, are good material for this model, we performed this combination approach using Schlegel's Japanese gecko, Gekko japonicus. We show that G. japonicus migrated from China to the western Japanese archipelago before Christ. The gecko species dispersed itself from western to eastern the archipelago on a time scale of thousands of years. There are many synchronizations between the dispersal history of G. japonicus and the historical development of human society. It is suggested by such synchronizations that humans have influenced the distribution of G. japonicus many times throughout its dispersal history.

Forest cover and geographical distance influence fine-scale genetic structure of leaf-toed geckos in the tropical dry forests of western Mexico

Abstract The biodiversity within tropical dry forests (TDFs) is astounding and yet poorly catalogued due to inadequate sampling and the presence of cryptic species. In the Mexican TDF, endemic species are common, and the landscape has been continually altered by geological and anthropogenic changes. To understand how landscape and environmental variables have shaped the population structure of endemic species, we studied the recently described species of leaf-toed gecko, Phyllodactylus benedettii, in coastal western Mexico. Using double-digest restriction site-associated DNA sequencing data, we first explore population structure and estimate the number of ancestral populations. The results indicate a high degree of genetic structure with little admixture, and patterns corresponding to both latitudinal and altitudinal gradients. We find that genetic structure cannot be explained purely by geographical distance, and that ecological corridors may facilitate dispersal and gene flow. We then model the spatial distribution of P. benedettii in the TDF through time and find that the coastline has been climatically suitable for the species since the Last Glacial Maximum. Landscape genetic analyses suggest that the combination of isolation by distance (IBD) and isolation by resistance (IBR; forest cover) has influenced the spatial genetic structure of the species. Overall, our genomic data demonstrate fine-scale population structure in TDF habitat, a complex colonization history, and spatial patterns consistent with both IBD and other ecological factors. These results further highlight the Mexican TDF as a diversity hotspot and suggest that continued anthropogenic changes are likely to affect native fauna.

Ecological niche overlap of two allopatric karst-adapted tiger geckos (Goniurosaurus) from northern Vietnam: microhabitat use and implications for conservation

ABSTRACT Anthropogenic pressures have caused a substantial decline of global biodiversity and have been further reported to strongly affect the ecological performance of species in their habitat, especially reptiles. Understanding the ecology of species and how species respond to habitat alterations is basic knowledge needed to develop conservation programmes and address issues of biodiversity loss. All five species of tiger geckos (Goniurosaurus) in Vietnam are known to be threatened by extinction due to anthropogenic impacts such as habitat degradation and harvesting for the international pet trade. However, conservation actions have only been initiated for the better-studied species. This study provides detailed basic data on microhabitat use of two allopatric sister species, namely Goniurosaurus huuliensis and G. luii. In total, 145 geckos (including 59 records of G. huuliensis and 86 records of G. luii) were observed during field surveys. All Goniurosaurus individuals were mostly recorded in the forest on karst formations, covered with evergreen broad-leaved woody trees, intermixed with ferns, shrubs and vines. Microhabitats of the two species were relatively similar in other traits, such as high vegetation coverage, high humidity, stable ambient temperature and dry-rock substrates. A multiple factor analysis supported that the ecological niche spaces of the two species highly overlap, even though their distribution ranges are geographically separated. We further found no intraspecific niche segregation in both species. The present data provide baseline knowledge for both in situ and ex situ conservation measures to protect species in the genus Goniurosaurus.

Response to the future climate change effect of two leopard geckos’ distribution (Sauria: Eublepharidae) in Iran

This study investigates the climate change effects on the distribution range of two species of nocturnal lizards in Iran. Using climate change scenarios recently (2021) published by IPCC, two species of leopard geckos that are distributed in the northeast ( Eublepharis turcmenicus ) and southwest ( Eublepharis angramainyu ) of Iran were selected and analyzed because they are only members of the family Eublepharidae in Iran and under high risk of extinction in the wild. We found that the most important bioclimatic variables that could affect the presence of these species are the minimum temperature in the coldest month of the year and the mean temperature in the wettest season; the Mean Temperature of the Driest Quarter and the Mean Temperature of the Coldest Quarter are the most important variables in future (2070) for E. angramainyu and E. turcmenicus , respectively. These variables are important directly at the beginning and end of the hibernation period of ectotherms such as lizards and can disrupt their life cycle by changing this period. Temperature changes over the next 50 years may not be a significant threat to these two species, as their range will be much wider than the current time, indicating that these species will be able to adapt to new conditions.

Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascars rainforest.

The Lygodactylus madagascariensis species group, constituting the subgenus Domerguella, currently contains five valid species of inconspicuous dwarf geckos from Madagascars humid forests, but at least 18 deep genetic lineages have been revealed by recent molecular studies. Given the high morphological similarity of these lineages, taxonomic resolution of this astonishing diversity requires efforts to correctly delimit species, as well as assigning the available nomina to the species-level lineages identified. We here combine DNA sequences of one mitochondrial and two nuclear-encoded gene fragments with morphometric measurements and scale counts, and report evidence for a species status of most of the previously identified lineages. In particular, we rely on sympatric and often even syntopic occurrence of several of these lineages without evidence for genetic admixture, and consistent with subtle morphological differences. Furthermore, the very high divergences of 7.423.8% pairwise distances in the relatively conserved mitochondrial 16S rRNA gene, combined with a lack of haplotype sharing in the nuclear-encoded genes and differences in scale counts convinced us that most of the other, allopatrically distributed lineages also represent distinct species. We elevate L. madagascariensis petteri to species level and formally name eight new species: L. salvi sp. nov., a species from the Sambirano region in northern Madagascar, previously called L. sp. 8; L. tantsaha sp. nov. (L. sp. 10), a species occurring sympatrically with L. madagascariensis and L. petteri on Montagne dAmbre in far northern Madagascar; L. roellae sp. nov. (L. sp. 17), a species characterized by a striped coloration in all known specimens, from northern Madagascar; L. winki sp. nov. (L. sp. 18), an unstriped species from northern Madagascar but belonging to a subclade mostly distributed in the eastern rainforests of the island; L. ulli sp. nov. (L. sp. 21), a species from the same subclade as L. winki but known only from the Marojejy Massif in the North East; L. fritzi sp. nov. (L. sp. 11), a further species of this subclade from coastal lowlands in the Northern Central East; L. hodikazo sp. nov. (L. sp. 23) known from a single specimen collected at the Tsingy de Bemaraha and therefore the only Domerguella species known from the West region of Madagascar; and L. hapei sp. nov. (L. sp. 26), an enigmatic species from the Sambirano region characterized by a striped pattern on the throat that is otherwise unknown in the subgenus. Three additional deep mitochondrial lineages of Domerguella were identified in our analysis, but could not be further analyzed due to the lack or scarcity of voucher specimens. More field work and collection of voucher specimens is needed to understand their status. Furthermore, the taxonomy of the Domerguella subclade occurring in eastern Madagascar, with three described species (L. guibei, L. miops, L. fritzi), two synonyms (L. septemtuberculatus, Microscalabotes spinulifer) and at least two further deep genetic lineages co-occurring in a relatively small area, requires further revisionary work, possibly aided by target-enrichment sequencing of the respective name-bearing types.

Rediscovery of Luperosaurus sorok Das, Lakim, and Kandaung, 2008 (Squamata, Gekkonidae) With Notes on Its Taxonomy and Natural History

Gekko (Rhacogekko) sorok is a small gecko species that was described in 2008 based on a single specimen from Borneo. We rediscovered multiple specimens of this species in the Lambir Hills National Park, Malaysian Borneo. The specimens constitute the second record of the species and the first record from Sarawak State. We describe the specimens and validate the taxonomic status of the species, which was formerly treated as Luperosaurus until recently and is now placed within the genus Gekko (Rhacogekko). Molecular phylogenetic analysis revealed that G. sorok is nested within a clade that includes Luperosaurus sensu stricto and is strongly supported as the sister lineage to L. joloensis. We tentatively propose that this species not be treated as Gekko (Rhacogekko) sorok, but rather as Luperosaurus sorok. We also comment on the natural history of this rare species and suggest that it is a high canopy dweller.

Three more novel species of South Asian Cnemaspis Strauch, 1887 (Squamata, Gekkonidae) from Kalakad Mundanthurai Tiger Reserve, Tamil Nadu, India

We describe three distinct, small-bodied, scansorial species of south Asian Cnemaspis from Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu, India—Cnemaspis azhagusp. nov. from Thirukurungudi forest range, Cnemaspis mundanthuraiensissp. nov. from Mundanthurai forest range and Cnemaspis kalakadensissp. nov. from Kalakad forest range. Phylogenetic analyses using a partial sequence of the mitochondrial ND2 gene and general morphology places each of the three new species in the beddomei, gracilis and littoralis clades, respectively. The three new species are diagnosed from all other described members of their respective clades by a suite of differing morphological characters including snout vent length, number of dorsal tubercle rows at mid-body, number of paravertebral tubercles, presence or absence of spine-like scales on flanks, number of ventral scales across belly at mid-body, number of ventral scales from mental to anterior border of cloaca, number of lamellae under digit IV of pes, number of femoral and/or precloacal pores and poreless scales separating these series, as well as subtle colouration differences. We also provide some novel characters of tail tuberculation of the three new species described herein. With the discovery of these three new species, eight species of geckos including five Cnemaspis are now known to be endemic to KMTR.

Spatial patterns and habitat use of penned and hard-released arboreal geckos translocated to an offshore island free of introduced mammals

Temporary penning prior to release is a strategy increasingly being used in lizard translocations to improve site fidelity and increase chances of translocation success. However, it is yet to be tested on a range of lizard taxa. Between 2015 and 2018, 49 individuals of a New Zealand endemic arboreal gecko species (ngahere gecko, Mokopirirakau “southern North Island”) were translocated to mammal-free Mana Island near Wellington as mitigation for a development project.

Description of a new species of leopard geckos, Eublepharis Gray, 1827 from Eastern Ghats, India with notes on Eublepharis hardwickii Gray, 1827

A recent molecular phylogenetic study of the genusEublepharisGray, 1827 revealed the presence of a genetically divergent lineage sister toEublepharis hardwickiiGray, 1827. We re-assessed the taxonomy ofEublepharis hardwickiibased on museum material in light of the molecular phylogenetic study and the results support the recognization of two species distributed on either sides of the Brahmani River. A redescription ofE. hardwickiiis presented based on the holotype and additional museum material along-with the description of a new species,Eublepharis pictussp. nov.from Andhra Pradesh and Odisha. The presence of yet another distinct lineage in such close geographical proximity in the northern Eastern Ghats highlights the significance of these relic forests and advocates the need for conservation prioritization.

The Global and Western Indian Ocean Dispersal of House Geckos From Asia Using Historical and Mitochondrial DNA Perspectives

Seven species of house geckos occur across the scattered islands of the Indian Ocean. Two of these, Hemidactylus frenatus and H. parvimaculatus are both widespread and possess distribution profiles that suggest pre-European, or perhaps natural dispersal to some islands. Of these, only H. frenatus currently has sufficient molecular data to begin exploring dispersal patterns. This species is one of the most successful reptile colonists, as demonstrated by its global, pantropical distribution. While in some areas, such as Australia and continental South America, its dispersal patterns are both recent and well-known, early historical records of Hemidactylus in the Indian Ocean islands suggest earlier and/or potentially non-human-mediated dispersals. Here, we reviewed the historical literature and combined those reports with an assessment of mitochondrial DNA diversity of a global sampling of H. frenatus samples that included modern and museum specimens. Our results corroborate previous studies and demonstrate the relatively high diversity within this species’ native range in Southeast Asia. In addition, the phylogenetic analysis suggests both a potential cryptic species complex, as well as global geographic structuring of different H. frenatus mitochondrial lineages. This has important implications for many comparative studies of this complex. Frequent and ongoing dispersals and colonizations complicate the identification of potentially older migration patterns. Further assessments including additional samples and analyses of additional genetic markers are necessary to disentangle older from more recent dispersals within this intriguing species.