The systematics of the Phelsuma madagascariensis species group of day geckos (Reptilia: Gekkonidae) in the Seychelles

Conventional taxonomy of the Seychelles populations of the Phelsuma madagascariensis species group has failed to illustrate the principal patterns of phenetic variation between island populations. The taxonomic history of the populations is discussed critically, and a revised nomenclature is suggested from the results of a recent comprehensive study of the forms using multivariate morphometrics. Three phenetic groups from the granitic islands are referred to as Phelsuma sundbergi sundbergi from the north western islands; P. s. ladiguensis from the north eastern islands; and P. s. longinsulae from the southern islands. Phelsuma abbotti populations from Aldabra and Assumption are considered subspecifically distinct, and not conspecific with the granitic island forms. The other coralline island populations are closely allied to the races from the granitic islands and probably represent recent colonization or introductions by man.     

A partial systematic revision of the day geckos, Phelsuma Gray, of Madagascar (Reptilia: Squamata: Gekkonidae)

Recent survey work in Madagascar has led to significant changes in the systematics of Madagascan Phelsuma. A new species from the Masoala Peninsula in northeastern Madagascar is described, which has a nostril position typical of Phelsuma from Mauritius, Reunion and Rodriguez islands. Phelsuma breviceps was rediscovered in extreme southern Madagascar and is resurrected from the synonymy of P. mutabilis. Phelsuma breviceps has fragile skin, a probable predator escape mechanism, similar to the condition found in geckos of the genera Ailuronyx and Geckokpis. Three new synonyms are recognized. Phelsuma befotahensis and P. checki are junior synonyms of P. abbotti , and P. minuthi is a junior synonym of P. lineata. Twenty species of Phelsuma occur in Madagascar, of which 16 are endemic. A checklist and identification key are provided. Conservation problems of Phelsuma in the coastal regions of Madagascar are briefly discussed.     

Relationships and evolution of the North African geckos,Geckonia and Tarentola (Reptilia: Gekkonidae), based on mitochondrial and nuclear DNA sequences

Mitochondrial (cytochrome b and 12S rRNA) and nuclear (c-mos) genes, analyzed by a variety of methods, indicate that the distinctive northwest African gecko Geckonia chazaliae is a member of the Tarentola clade, being most closely related to the species of the western Canary and Cape Verde islands. Relationships in Tarentola as a whole are as follows: (T. americana ((T. mauritanica, T. angustimentalis) ((T. deserti, T. boehmei) ((T. b. boettgeri-South (T. b. boettgeri-North (T. b. bischoffi, T. b. hierrensis))) ((T. annularis, T. ephippiata) (Geckonia, T. delalandii, T. gomerensis, Cape Verde species)))))); nearly all nodes have high bootstrap support. Results confirm that T. americana of Cuba and the Bahamas separated at the most basal dichotomy of the phylogeny and give no positive support for the monophyly of the subgenera Tarentola s. str. and Makariogecko. The latter includes Geckonia and the subgenus Sahelogecko. Continental Tarentola appear to have invaded the Sahara desert from its northern edge. They have also colonized groups of Atlantic islands five times: a single invasion of the West Indies and three of the Canary islands, one of which then went on to invade the Cape Verde archipelago. The phylogeny corroborates anatomical evidence that the ground-dwelling Geckonia had a climbing ancestry, something that is paralleled in some southern African terrestrial gekkonids related to Pachydactylus. Distinctive derived features of Geckonia occur in other gekkonids that are ground dwelling in arid habitats and may be functionally related to this environment. The evolution of such features indicates that, although Tarentola is generally very uniform and may have been so for over 10 million years, this is not due to any overwhelming phylogenetic constraint. G. chazaliae should be included in Tarentola, as Tarentola chazaliae.     

Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences

With more than 80 species inhabiting all warm continental land masses and hundreds of intervening continental and oceanic islands, Hemidactylus geckos are one of the most species-rich and widely distributed of all reptile genera. They consequently represent an excellent model for biogeographic, ecological, and evolutionary studies. A molecular phylogeny for Hemidactylus is presented here, based on 702 bp of mtDNA (303 bp cytochrome b and 399 bp 12S rRNA) from 166 individuals of 30 species of Hemidactylus plus Briba brasiliana, Cosymbotus platyurus, and several outgroups. The phylogeny indicates that Hemidactylus may have initially undergone rapid radiation, and long-distance dispersal is more extensive than in any other reptilian genus. In the last 15 My, African lineages have naturally crossed the Atlantic Ocean at least twice. They also colonized the Gulf of Guinea, Cape Verde and Socotra islands, again sometimes on more than one occasion. Many extensive range extensions have occurred much more recently, sometimes with devastating consequences for other geckos. These colonizations are likely to be largely anthropogenic, involving the 'weedy' commensal species, H. brookii s. lat, H. mabouia, H. turcicus, H. garnotii, and H. frenatus. These species collectively have colonized the Mediterranean region, tropical Africa, much of the Americas and hundreds of islands in the Pacific, Indian, and Atlantic oceans. Five well-supported clades are discernable in Hemidactylus, with the African H. fasciatus unallocated. 1. Tropical Asian clade: (Cosymbotus platyurus (H. bowringii, H. karenorum, H. garnotii)) (H. flaviviridis (Asian H. brookii, H. frenatus)). 2. African H. angulatus and Caribbean H. haitianus. 3. Arid clade, of NE Africa, SW Asia, etc.: (H. modestus (H. citernii, H. foudai)) (H. pumilio (H. granti, H. dracaenacolus) (H. persicus, H. macropholis, H. robustus, H. turcicus (H. oxyrhinus (H. homoeolepis, H. forbesii))). 4. H. mabouia clade (H. yerburii, H. mabouia). 5. African-Atlantic clade: H. platycephalus ((H. agrius, H. palaichthus) (H. longicephalus, H. greeffi, H. bouvieri, Briba brasiliana))). Cosymbotus and Briba are synonymized with Hemidactylus, and African populations of H. brookii separated as H. angulatus, with which H. haitianus may be conspecific. Some comparatively well-sampled widespread species show high genetic variability (10-15% divergence) and need revision, including Cosymbotus platyurus, H. bowringii, Asian H. brookii, H. frenatus, H. angulatus, and H. macropholis. In contrast, most populations of H. mabouia and H. turcicus are very uniform (1-2% divergence). Plasticity of some of the morphological features of Hemidactylus is confirmed, although retention of primitive morphologies also occurs.     

The genus Paragehyra (Reptilia: Sauria: Gekkonidae) in southern Madagascar

Paragehyra Angel is a gekkonine genus endemic to Madagascar. From 1929 until the present, the genus was represented only by the holotype of the single known species. Paragehyra petiti. The type locality of P. petiti is in the driest part of Madagascar (south-west) where rainless periods may exceed 11 months and the original vegetation consists of thorn scrub or dry deciduous forest. No new individuals of P. petiti were observed. However, a second, undescribed species of the genus was discovered in October 1990 on the opposite side of the island (south-east) in primary-low montane rainforest. The new form was observed at four localities at the southernmost extremity of rainforest in Madagascar.     

Between a rock and a soft place: microtopography of the locomotor substrate and the morphology of the setal fields of Namibian day geckos (Gekkota: Gekkonidae: Rhoptropus)

Many species of gekkotans possess adhesive subdigital pads that allow them to adhere to, and move on, a wide variety of surfaces. The natural surfaces exploited by these lizards may be rough, undulant and unpredictable and therefore likely provide only limited, patchy areas for adhesive contact. Here, we examine the microtopography of rock surfaces used by seven species of Rhoptropus and compare this to several rough and smooth artificial surfaces employed in previous studies of gekkotan adhesion. These data are considered in relation to the form, configuration, compliance and functional morphology of the setal fields of these species. Our results demonstrate that natural rock surfaces are rough and unpredictable at the scale of the setal arrays, with equal amounts of variation existing within and between the various types of rock surfaces examined. Such surfaces differ from smooth and rough artificial surfaces in the proportion of surface area available for attachment and the relative predictability of surface undulance. Generally, setal field characteristics of individual species are not relatable to specific substrates, but instead are configured to allow for sufficient attachment to a wide variety of unpredictable surfaces. Our findings provide insight into the evolution and microanatomy of the adhesive system of gekkotan lizards and its adaptive relationship to topographically unpredictable surfaces.     

Formal taxa, species groups, and perception of the genus Diplodactylus (Reptilia: Gekkonidae)

Genera with large numbers of species present particular difficulties; the analysis of relationships, of included taxa may be roblematic. One attempt to aproach this problem involves the reco of clusters of species tIat may be informally assembgd into species groups. The problems tE:g the recognition of such assemblages may induce are exlored. It is not that the species groups, as originail formulated, are problematic as they are initiafy erected to demarcate clusters withm an imperfectly known phylogeny of a supposedly monophyletic group. These species groups, however, tend to become recognized as “taxa” rather than operationaf clusters and as such tend to influence the approach to the inclusive taxon taken by subsequent workers. Rather than testing the concept of the species grous, there is a tendency to retain them and to insert other groups between them that do not exactly fit the original scheme. The establishment of species groups, first used to clarify a complex situation, has teen a source of problems for subseuent workers. The history of this aproach is traced for the gekkonid genus Diplodactylus and the problems that have arisen are outlined.     

Phylogenetic systematics of the genus Echinococcus (Cestoda: Taeniidae)

Echinococcosis is a serious helminthic zoonosis in humans, livestock and wildlife. The pathogenic organisms are members of the genus Echinococcus (Cestoda: Taeniidae). Life cycles of Echinococcus spp. are consistently dependent on predator–prey association between two obligate mammalian hosts. Carnivores (canids and felids) serve as definitive hosts for adult tapeworms and their herbivore prey (ungulates, rodents and lagomorphs) as intermediate hosts for metacestode larvae. Humans are involved as an accidental host for metacestode infections. The metacestodes develop in various internal organs, particularly in liver and lungs. Each metacestode of Echinococcus spp. has an organotropism and a characteristic form known as an unilocular (cystic), alveolar or polycystic hydatid. Recent molecular phylogenetic studies have demonstrated that the type species, Echinococcus granulosus, causing cystic echinococcosis is a cryptic species complex. Therefore, the orthodox taxonomy of Echinococcus established from morphological criteria has been revised from the standpoint of phylogenetic systematics. Nine valid species including newly resurrected taxa are recognised as a result of the revision. This review summarises the recent advances in the phylogenetic systematics of Echinococcus, together with the historical backgrounds and molecular epidemiological aspects of each species. A new phylogenetic tree inferred from the mitochondrial genomes of all valid Echinococcus spp. is also presented. The taxonomic nomenclature for Echinococcus oligarthrus is shown to be incorrect and this name should be replaced with Echinococcus oligarthra.     

Complete mitochondrial genome of the red-spotted tokay gecko (Gekko gecko, Reptilia: Gekkonidae): comparison of red- and black-spotted tokay geckos

Here, we sequenced the complete mitochondrial genome of the red-spotted tokay gecko (Squamata: Gekkonidae). The genome is 16,590 bp in size. Its gene arrangement pattern was identical with that of black-spotted tokay gecko. We compared the mitochondrial genome of red-spotted tokay gecko with that of the black-spotted tokay gecko. Nucleotide sequence of the two whole mitochondrial genomes was 97.99% similar, and the relatively high similarity seems to indicate that they may be separated at the subspecies level. The information of mitochondrial genome comparison of the two morphological types of tokay gecko is discussed in detail.     

Molecular phylogeny,biogeography and chromosome evolution of Malagasy dwarf geckos of the genus Lygodactylus (Squamata,Gekkonidae)

We performed a phylogenetic analysis using nuclear (RAG‐1, RAG‐2) and mitochondrial (16S) markers, a statistical Bayesian reconstruction of ancestral distribution areas and a karyological analysis on most Malagasy species of the gekkonid genus Lygodactylus. The phylogenetic analysis largely confirms major basal branching pattern of previous molecular studies, but highlights significant differences concerning both the relationships between different species groups as well as those within groups. The biogeographic analysis supports a Malagasy origin of Lygodactylus, an oversea dispersal to continental Africa and a return to Madagascar. The L. madagascariensis group (also including a new candidate species identified herein) is the most basal clade in Lygodactylus, and the sister group of a clade with all the remaining species. The second most basal clade is the L. verticillatus group, placed as the sister group of a clade comprising African and Malagasy species. The sister lineage of the L. verticillatus group originated the African radiation through an oversea dispersal out of Madagascar. Eventually, the sister lineage of the L. capensis group originated secondary dispersals from Africa to Madagascar. In Madagascar, lineage diversification in different species groups mainly occurred from southern to northern and eastern regions. Dispersal, vicariance and paleoclimatic refugia probably played a relevant role in the evolutionary history of closely related taxa and in speciation mechanisms. The cytogenetic analysis evidenced a high karyotypic variability in Lygodactylus (from 2n = 34 to 2n = 40), which is at least partly consistent with the phylogenetic relationships and the composition of the various species group. Chromosome evolution occurred independently in different lineages, mainly through a reduction in the chromosome number and starting from a putative primitive karyotype of 2n = 40 with all telocentric elements.     

Molecular systematics of Stenodactylus (Gekkonidae), an Afro-Arabian gecko species complex

We conducted a phylogenetic analysis of Stenodactylus geckos using mitochondrial and three nuclear genes in order to understand the divergence within this genus. Stenodactylus is a complex with deep divergences that date to at least the Miocene; these patterns are seen in several other complexes in this region, indicating important and shared biogeographic processes affecting several taxonomic groups. Divergence between disjunct populations from three species in the Arabian Peninsula may have arisen because of Pliocene and Pleistocene restructuring of sand dunes. As currently recognized, Stenodactylus is not a monophyletic genus with respect to Tropiocolotes. We resurrect the monotypic genus Pseudoceramodactylus to address this problem of monophyly.     

Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa

Abstract.  Within a framework of historical analysis of Eneopterinae crickets, the genus Pseudolebinthus Robillard gen.n. and two new species P. africanus Robillard, sp.n. and P. whellani Robillard, sp.n. , endemic from south-east Africa, are described. A cladistic analysis using 198 morphological characters and 47 terminals assessed the phylogenetic position of the new taxa within the subfamily. The resultant topologies support the previously proposed phylogeny for the subfamily and contained tribes. The monophyly of Pseudolebinthus is supported strongly as well as its sister relationship with Xenogryllus within the tribe Xenogryllini. A key to Eneopterinae tribes, Xenogryllini genera and Pseudolebinthus species is given. Taxonomic, evolutionary and acoustic issues raised by the recognition of Pseudolebinthus are discussed.     

Long-distance colonization and radiation in gekkonid lizards, Tarentola (Reptilia: Gekkonidae), revealed by mitochondrial DNA sequences

Morphological systematics makes it clear that many non-volant animal groups have undergone extensive transmarine dispersal with subsequent radiation in new, often island, areas. However, details of such events are often lacking. Here we use partial DNA sequences derived from the mitochondrial cytochrome b and 12S rRNA genes (up to 684 and 320 bp, respectively) to trace migration and speciation in Tarentola geckos, a primarily North African clade which has invaded many of the warmer islands in the North Atlantic Ocean. There were four main invasions of archipelagos presumably by rafting. (i) The subgenus Neotarentola reached Cuba up to 23 million years (Myr) ago, apparently via the North Equatorial current, a journey of at least 6000 km. (ii) The subgenus Tarentola invaded the eastern Canary Islands relatively recently covering a minimum of 120 km. (iii) The subgenus Makariogecko got to Gran Canaria and the western Canary Islands 7-17.5 Myr ago, either directly from the mainland or via the Selvages or the archipelago of Madeira, an excursion of 200-1200 km. (iv) A single species of Makariogecko from Gomera or Tenerife in the western Canaries made the 1400 km journey to the Cape Verde Islands tip to 7 Myr ago by way of the south-running Canary current. Many journeys have also occurred within archipelagos, a minimum of five taking place in the Canaries and perhaps 16 in the Cape Verde Islands. Occupation of the Cape Verde archipelago first involved an island in the northern group, perhaps São Nicolau, with subsequent spread to its close neighbours. The eastern and southern islands were colonized from these northern islands, at least two invasions widely separated in time being involved. While there are just three allopatric species of Makariogecko in the Canaries, the single invader of the Cape Verde Islands radiated into five, most of the islands being inhabited by two of these which differ in size. While size difference may possibly be a product of character displacement in the northern islands, taxa of different sizes reached the southern islands independently.     

A new large species of gecko of the genus Hemidactylus Oken, 1817 (Reptilia: Sauria: Gekkonidae) from the Eastern Ghats,India

A new large species of gecko superficially resembling Hemidactylus maculatus Duméril & Bibron and its allied large Hemidactylus is described from Eastern Ghats. The new species, Hemidactylus kangerensis sp. nov. can be easily distinguished from members of the H. maculatus complex based on the number of femoral pores, i.e. 18–21 on each side, separated medially by 4 non-pored scales. A phylogenetic analysis based on partial sequence of mitochondrial gene cytochrome b shows that the gecko is a member of the “H. prashadi” clade and is sister to H. maculatus, from which it differs in an uncorrected pairwise sequence divergence of 10%. The discovery of a new large gecko from Eastern Ghats advocates the need for biodiversity assessment across the neglected Eastern Ghats.     

Temperature effects on oxygen consumption of two nocturnal geckos,Ptyodactylus hasselquistii (Donndorff) andBunopus tuberculatus (Blanford) (Reptilia: Gekkonidae) in Saudi Arabia

Summary Oxygen consumption rates of the lizardsPtyodactylus hasselquistii (adult and subadults) andBunopus tuberculatus (adult) were determined in relation to ambient temperatures ranging from 10 to 35°C using a double-chamber, volumetric closed system. The metabolic rate-temperature curves for both species were triphasic and similar in shape, but O₂ consumption differed between species.The low thermal dependence at temperatures between 15 and 25°C, common to both species, was correlated with the lizards' dual mode of thermoregulatory activity and ecology.     

Temperature-Dependent Sex Determination in Gekko japonicus (Gekkonidae, Reptilia)

The effects of temperature on sexual differentiation in early development of the gekkonid lizard, Gekko japonicus were studied. The eggs were collected within 24 hr after the oviposition and were incubated at 20, 24, 28 and 32°C. The number of eggs hatched was 14 at 24°C, 20 at 28°C and 21 at 32°C. Hatching never occurred at 20°C. The hatched lizards without Müllerian ducts were judged as males. The sex of all lizards with Müllerian ducts were identified histologically. The sex ratios, male/(male+female), were 0.07 at 24°C, 0.75 at 28°C and 0.24 at 32°C. The disparities of the sex ratio from 1/2 were statistically significant and differences in the sex ratio with various incubation temperatures were also significant. These sex ratios can probably be best interpreted by a temperature-dependent sex determination. The different sex ratios do not seem to be related to a predetermination of sex with a differential mortality.     

Cryptic diversity within the Moroccan endemic day geckos Quedenfeldtia (Squamata: Gekkonidae): a multidisciplinary approach using genetic,morphological and ecological data

Quedenfeldtia (Boettger, 1883) is a genus of diurnal geckos, endemic to the Atlas Mountains in Morocco, with two species being recognized: Quedenfeldtia moerens and Quedenfeldtia trachyblepharus. Quedenfeldtia moerens is found across a wide variety of habitats, from sea level to 3000 m a.s.l., whereas Q. trachyblepharus occupies exclusively high mountain regions reaching up to 4000 m a.s.l. This differentiation, offers an interesting model for study biogeographical patterns and evolutionary scenarios in a North African endemic. Analysis of two mitochondrial (12S rRNA and ND4) and four nuclear (ACM4, MC1R, PDC, and Rag1) DNA markers revealed high genetic variation, consistent with other recent phylogeographical studies, and with the two currently described species. However, within each species, a subdivision into two groups with geographical consistence was found. Multivariate morphological analyses confirmed the existence of two main phenotypes, whereas ecological niche modelling identified various environmental variables associated with the distribution of each species, and helped to predict occurrences outside the confirmed ranges. The results obtained in the present study indicate the possible existence of additional ‘cryptic’ species within this genus, a condition found in many North African reptiles, and particularly common in geckos. In general, North African montane fauna appears to reflect the occurrence of diverse palaeoendemics, as seen in Central Africa Mountain systems, rather than the pattern of recent postglacial recolonization observed in Europe. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.