Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean.

Diurnal dwarf geckos of the genus Lygodactylus are distributed in tropical and subtropical regions and live in highly diverse habitats. The genus currently comprises 79 species and several candidates for new species or subspecies. Most of these taxa occur in Sub-Saharan Africa and Madagascar, with only two described species in South America. Although the main center of diversity of Lygodactylus currently is Africa, the genus probably has a Malagasy origin, followed by two or three independent transoceanic dispersal events between Madagascar and Africa and one trans-Atlantic dispersal from Africa to South America. A few species colonised islands in the Western Indian Ocean belonging to the Zanzibar Archipelago and to the Îles Éparses. Here we examined L. grotei pakenhami from Pemba Island, L. insularis from Juan de Nova, and L. verticillatus from Europa Island to clarify their taxonomic status and their origin. Concerning L. grotei pakenhami and L. insularis, preceding studies pointed to a relation to species of the African L. capensis group. In contrast, L. verticillatus on Europa Island is considered to be conspecific with Malagasy populations. Therefore, we conducted a phylogenetic study of the African L. capensis group and the Malagasy L. verticillatus group, and examined color pattern, selected morphological characters and two mitochondrial markers (ND2 for African and 16S rRNA for Malagasy Lygodactylus). Lygodactylus grotei pakenhami from Pemba and L. grotei from mainland Africa cannot be distinguished by their scalation, but their reciprocal monophyly suggested by mitochondrial DNA, conspicuously different coloration (both in adults and hatchlings) and their high genetic distances (16.3% in ND2) support the hypothesis that these taxa represent two distinct species. Consequently, we elevate L. grotei pakenhami to species level, as Lygodactylus pakenhami Loveridge, 1941. Lygodactylus pakenhami is endemic to Pemba Island which was possibly separated from the African mainland during the late Miocene or Early Pliocene (6 million years ago). The simplest explanation for the existence of L. pakenhami on Pemba is vicariance. A recent, human-mediated transportation is excluded, as the molecular data clearly indicate a longer period of isolation. Lygodactylus insularis has been supposed to be related to the taxa 'capensis' or 'grotei'. However, it is impossible to discern the relationship of L. insularis, L. capensis and L. grotei by means of scalation or coloration alone. Our molecular phylogenetic analyses reveal that L. insularis is embedded within the L. capensis group, clearly indicating its African origin. The single gene (ND2) as well as the multigene analyses fully support a closer common origin of L. insularis and L. capensis than of L. insularis and L. grotei. However, the position of L. insularis within the clade formed by L. insularis, L. nyaneka, L. capensis sensu stricto and six L. aff. capensis groups is not clearly resolved. Lygodactylus insularis is endemic on Juan de Nova Island, an old low elevation atoll. That all L. insularis mitochondrial sequences are very similar to each other and together form a monophyletic lineage is in agreement with the hypothesis of a single dispersal event to the island. For the L. verticillatus population from Europa Island our mitochondrial data suggest close relationships to conspecific samples from the coastal regions of south-western Madagascar. As we found no relevant morphological or genetic differences between the insular and the Malagasy populations of L. verticillatus, and no remarkable genetic variation within the monophyletic lineage on Europa, we suggest a single, very recent dispersal event, perhaps human-mediated. Although the genus Lygodactylus colonised Africa, islands in the Gulf of Guinea, South America and some islands in the Western Indian Ocean, it seems-compared to other lizard genera-to be only moderately successful in transoceanic long-distance dispersal.

Multigene phylogenetic analysis of Lygodactylus dwarf geckos (Squamata: Gekkonidae).

The genus Lygodactylus is a group of small, diurnal geckos distributed mainly in Africa and Madagascar. A closely related, monotypic genus is Microscalabotes from Madagascar. We examined fragments of two moderate- to slow-evolving nuclear genes (RAG-1, RAG-2) and of two fast-evolving mitochondrial genes (16SrRNA, cytochrome b) to estimate phylogenetic relationships among 28 (of about 60) species (including Microscalabotes). Since cytochrome b provided little resolution for analysis of interspecific relationships in Lygodactylus, it was excluded from the concatenated Bayesian analysis. Our well-resolved Bayesian inference tree and its resulting clades confirm many previously suggested relationships among dwarf geckos based mainly on morphological characters, while several novel relationships lead us to regroup some species and to synonymize the genus Microscalabotes with the genus Lygodactylus. Our phylogenetic analysis well supports the monophyly of Lygodactylus (including Microscalabotes). However, it rejects monophyly of the Madagascan Lygodactylus. On the other hand, the data do not reject monophyly of the African forms. Though Africa is currently the main centre of diversity of Lygodactylus, our data suggest that the most parsimonious scenario would be a Madagascan origin, one dispersal to Africa, and one back to Madagascar.

Sexual characteristics and spermatogenesis in males of the parthenogenetic gecko Lepidodactylus lugubris (Reptilia, Gekkonidae).

Obligately parthenogenetic lizards usually are all-female populations of hybrids producing diploid oocytes by premeiotic endomitosis and quasi-normal meiosis. In an all-female strain of the gekkonid lizard Lepidodactylus lugubris several phenotypic males arose spontaneously. The sexual characteristics of these males were studied using light and electron microscopy and compared with normal males of the bisexual genus Lygodactylus. Emphasis was layed on morphology of seminiferous tubules, occurrence of spermatogenic stages and ultrastructure of spermatozoa. The phenotypic males possessed preanal pores filled with secretions and a sexual nephric segment which were exactly the same as in normal, reproductively active males. In the testes, density and morphology of non-spermatogenic cell types, the Leydig and Sertoli cells, indicate a normal production of testicular testosterone and a normal function of the blood-testis barrier, respectively. Both in the normal and the phenotypic males, all meiotic cell types of spermatogenesis can be recognised in the seminiferous tubules and are apparently identical, indicating a normal meiosis without impairment in the phenotypic males. In contrast, the differentiation process of spermatids is markedly disturbed in the phenotypic males of L. lugubris. In the normal male, spermiogenesis results in mature spermatids and spermatozoa with small elongated nuclei, an acrosomal complex, and a flagellar tail possessing one axoneme. Spermatozoa fill both the lumen of most seminiferous tubules and the lumina of ductus epididymidis and ductus deferens. In the phenotypic male, spermiogenesis results in seemingly normal spermatids and in spermatozoa with large, non-elongated, deformed nuclei and/or irregular tails possessing more than one axoneme. Both the lumen of most seminiferous tubules and the lumina of the ductus epididymidis and the ductus deferens contain relatively few spermatozoa. We suggest that the phenotypic males inherited the ability for a premeiotic endomitosis from their all-female ancestral lineage. While in females this leads to quasi-normal meiosis and diploid oocytes capable of development, the small nuclei of the spermatozoa are unable to contain a diploid set of chromosomes. Because of the high amount of deformed spermatozoa and possibly uncontrolled loss of genetic material in structurally normal, but aneuploid spermatozoa we conclude that these otherwise perfect males are infertile, thus constituting another example of gametic sterility.

The optokinetic reaction in foveate and afoveate geckos.

To investigate determinants of symmetry of the monocular horizontal optokinetic reaction (hOKR) in vertebrates, we performed behavioural studies in diurnal foveate, as well as nocturnal afoveate geckos. During binocular viewing hOKR gain was equal for movement to the left or right, during monocular stimulation, all afoveate geckos (Lepidodactylus lugubris, Gekko gecko, Eublepharis macularius) and the foveate Lygodactylus spp. exclusively reacted to temporo-nasal stimulation with stabilising head movements whereas in Phelsuma madagascariensis a naso-temporal component of hOKR could be elicited albeit much weaker. Thus, neither the presence of a fovea nor lifestyle seems to be decisive for a symmetrical monocular hOKR.