Eugongylus Group Research Articles

A comparative analysis of testicular sperm morphology in fossorial and surface‐living skinks in South Africa

AbstractWe described for the first time the spermatozoan ultrastructure of the fully pentadactyl surface‐living skink Trachylepis punctatissima, and limbless fossorial skink Acontias meleagris. The spermatozoa of both species follow the general patterns observed within the Squamata. However, several important differences were detected between the two species in the head region (shape of the anterior acrosome, size of acrosome and nucleus) and especially in the midpiece (size of the midpiece, the presence of regular rows of dense bodies, size and number of mitochondria and beginning of the fibrous sheath). Both species shared more characters with the Sphenomorphus + Egernia group than with the Eugongylus group proposed by Jamieson, Oliver, and Scheltinga (Acta Zoologica, 77, 85). Differences in the spermatozoan ultrastructure between T. punctatissima and A. meleagris could be due to distinct ecological and physiological requirements for fertilization, but additional research on these genera and within the Scincidae is required to disentangle this hypothesis, and to disentangle the phylogenetic relationships among skinks.

Taxonomic resolution to the problem of polyphyly in the New Caledonian scincid lizard genus Lioscincus (Squamata: Scincidae)

Recent genetic studies have identified the New Caledonian scincid genus Lioscincus to be polyphyletic, comprising four distinct evolutionary lineages which we recognize at the generic level. The revised concept of Lioscincus s.s. now includes only the type species Lioscincus steindachneri Bocage, 1873 and the recently described Lioscincus vivae Sadlier, Bauer, Whitaker & Smith, 2004. The three remaining lineages identified are: Leiolopisma tillieri Ineich & Sadlier, 1991 and Lioscincus maruia Sadlier, Whitaker & Bauer, 1998 for which the genus Phasmasaurus gen. nov. is proposed; Lygosoma (Mocoa) nigrofasciolatus Peters, 1869 and Leiolopisma greeri Bohme, 1979 for which the genus Epibator gen. nov. is proposed; and Lygosoma (Leiolopisma) novaecaledoniae Parker, 1926 for which the genus Caesoris gen. nov. is proposed. Each of these genera is diagnosed by a suite of morphological apomorphies which in combination is unique within the Eugongylus group of skinks of which each is a member. The revised taxonomy presented here, which recognises an additional three new genera, brings the number of endemic skink genera present in New Caledonia to 17, representing approximately 40% of the generic diversity within the Australian/Pacific region Eugongylus group of skinks as currently recognised. Although we are reluctant to erect new genera to accommodate one or two species, the data at hand suggest this as the most accurate reflection of the available genetic and morphological data.

Evaluating the performance of anchored hybrid enrichment at the tips of the tree of life: a phylogenetic analysis of Australian Eugongylus group scincid lizards.

BackgroundHigh-throughput sequencing using targeted enrichment and transcriptomic methods enables rapid construction of phylogenomic data sets incorporating hundreds to thousands of loci. These advances have enabled access to an unprecedented amount of nucleotide sequence data, but they also pose new questions. Given that the loci targeted for enrichment are often highly conserved, how informative are they at different taxonomic scales, especially at the intraspecific/phylogeographic scale? We investigate this question using Australian scincid lizards in the Eugongylus group (Squamata: Scincidae). We sequenced 415 anchored hybrid enriched (AHE) loci for 43 individuals and mined 1650 exons (1648 loci) from transcriptomes (transcriptome mining) from 11 individuals, including multiple phylogeographic lineages within several species of Carlia, Lampropholis, and Saproscincus skinks. We assessed the phylogenetic information content of these loci at the intergeneric, interspecific, and phylogeographic scales. As a further test of the utility at the phylogeographic scale, we used the anchor hybrid enriched loci to infer lineage divergence parameters using coalescent models of isolation with migration.ResultsPhylogenetic analyses of both data sets inferred very strongly supported trees at all taxonomic levels. Further, AHE loci yielded estimates of divergence times between closely related lineages that were broadly consistent with previous population-level analyses.ConclusionsAnchored-enriched loci are useful at the deep phylogeny and phylogeographic scales. Although overall phylogenetic support was high throughout the Australian Eugongylus group phylogeny, there were nonetheless some conflicting or unresolved relationships, especially regarding the placement of Pseudemoia, Cryptoblepharus, and the relationships amongst closely-related species of Tasmanian Niveoscincus skinks.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-015-0318-0) contains supplementary material, which is available to authorized users.

Development of yolk sac and chorioallantoic membranes in the Lord Howe Island skink, Oligosoma lichenigerum

Development of the yolk sac of squamate reptiles (lizards and snakes) differs from other amniote lineages in the pattern of growth of extraembryonic mesoderm, which produces a cavity, the yolk cleft, within the yolk. The structure of the yolk cleft and the accompanying isolated yolk mass influence development of the allantois and chorioallantoic membrane. The yolk cleft of viviparous species of the Eugongylus group of scincid lizards is the foundation for an elaborate yolk sac placenta; development of the yolk cleft of oviparous species has not been studied. We used light microscopy to describe the yolk sac and chorioallantoic membrane in a developmental series of an oviparous member of this species group, Oligosoma lichenigerum. Topology of the extraembryonic membranes of late stage embryos differs from viviparous species as a result of differences in development of the yolk sac. The chorioallantoic membrane encircles the egg of O. lichenigerum but is confined to the embryonic hemisphere of the egg in viviparous species. Early development of the yolk cleft is similar for both modes of parity, but in contrast to viviparous species, the yolk cleft of O. lichenigerum is transformed into a tube-like structure, which fills with cells. The yolk cleft originates as extraembryonic mesoderm is diverted from the periphery of the egg into the yolk sac cavity. As a result, a bilaminar omphalopleure persists over the abembryonic surface of the yolk. The bilaminar omphalopleure is ultimately displaced by intrusion of allantoic mesoderm between ectodermal and endodermal layers. The resulting chorioallantoic membrane has a similar structure but different developmental history to the chorioallantoic membrane of the embryonic hemisphere of the egg.

Lygosomine phylogeny and the origins of Australian scincid lizards

Aim Australian scincid lizards represent three distinct groups within the cosmopolitan clade Lygosominae, the Egernia, Eugongylus and Sphenomorphus groups. This paper presents a time-calibrated phylogeny for Lygosominae that provides the necessary temporal framework for assessing the contributions of immigration from Asia and of Gondwanan inheritance in the derivation of the Australian scincid fauna. Location Australasia, Asia, Africa. Methods Phylogenetic relationships and divergence times were inferred from novel BDNF, c-mos and PTPN12 sequences (2408 aligned sites). Results Lygosomine monophyly is well supported, and there is strong support for monophyly of the Egernia, Eugongylus and Sphenomorphus groups. A sister-group relationship of Tribolonotus (distributed in Melanesia and the Papuan Region) and the Egernia group is strongly supported in both Bayesian and maximum likelihood analyses. Australian representatives of the Sphenomorphus group compose a significantly supported clade estimated to have originated c. 25 Ma. An age of c. 18 Ma is inferred for a strongly supported clade comprising Australian representatives of the Egernia group; this clade diverged from Corucia zebrata (confined to the Solomon Islands) c. 25 Ma and from Tribolonotus c. 54 Ma. A well-supported clade including all Australian Eugongylus group taxa sampled is estimated to have arisen c. 20 Ma. Main conclusions The Australian Sphenomorphus group is nested within the more inclusive Sphenomorphus group (distributed primarily in Asia and Australasia), suggesting comparatively recent descent from a colonizing Asian ancestor; the divergence times inferred here indicate that colonization occurred during the mid Cenozoic, subsequent to the rifting of Australia from Antarctica. An Oligocene origin of the extant Eugongylus group fauna of Australasia (the basal members of which are distributed in the Southwest Pacific) indicates that Eugongylus group lygosomines also dispersed to Australia relatively recently. The Egernia group diverged from Tribolonotus in the Early Eocene; however, extant Egernia group lygosomines originated only during the Late Oligocene, implying extensive pruning of stem taxa (i.e. extinction). As a result, inferences of the timing of dispersal into Australia are associated with substantial uncertainty, although independent palaeontological evidence suggests that the Egernia group entered Australia prior to the Oligocene, immediately after (or perhaps before) the separation of Australia and Antarctica.

Isolation and characterization of microsatellite loci from the invasive delicate skink (Lampropholis delicata), with cross-amplification in other Australian Eugongylus group species

The delicate skink (Lampropholis delicata), native to eastern Australia, is one of the most successful invasive reptiles in the Pacific region, with established populations in New Zealand, the Hawaiian Islands, and Lord Howe Island. Here we isolate and characterize 12 polymorphic microsatellite loci for L. delicata. Genetic variation was assessed in individuals from Sydney and also from the eastern Victoria and Tasmanian region. The number of alleles per locus ranged from 2 to 23, with high levels of heterozygosity. All microsatellite loci successfully amplified in individuals from the three invasive populations. Cross-amplification of these loci was assessed in 14 species from other Eugongylus group skink genera. These microsatellite markers will be used to investigate the invasion dynamics of L. delicata.

Origin, diversification, and systematics of the New Zealand skink fauna (Reptilia: Scincidae)

The diverse scincid lizard fauna of the largely submerged subcontinent of Zealandia (which incorporates New Zealand, New Caledonia, Lord Howe Island, Norfolk Island, and the Chatham Islands) forms a monophyletic lineage within the Eugongylus group of skinks. We use 4062 bp of mitochondrial (ND2, ND4, Cytochrome b, 12SrRNA, 16SrRNA) and nuclear (Rag-1) DNA sequence data to recover a molecular phylogeny for the New Zealand skink fauna, and investigate the origin and diversification of skinks in New Zealand. Our phylogeny includes 32 of the 33 extant described New Zealand skink species ( Cyclodina and Oligosoma), the Lord Howe Island skink ( C. lichenigera), and representatives from several New Caledonian genera. Neighbour-joining, Maximum Parsimony, Maximum Likelihood, and Bayesian phylogenetic analyses are used to demonstrate that the New Zealand skink species form a single monophyletic lineage, with C. lichenigera representing a closely related sister lineage to the New Zealand radiation. Our relaxed molecular clock analyses indicate that skinks colonised New Zealand in the early Miocene (16–22.6 mya), shortly after the ‘Oligocene drowning’ event (∼25 mya). We propose that skinks reached New Zealand from New Caledonia via long-distance overwater dispersal, with C. lichenigera persisting on volcanic islands along the Lord Howe Rise and Norfolk Ridge. Eight major genetic clades are evident within the New Zealand skink fauna, with the divergences among these clades during the early to mid-Miocene resulting in distinct open habitat, forest, and coastal radiations. Subsequent diversification in the late Miocene–Pliocene appears to coincide with tectonic activity along the Alpine Fault and the uplift of the Southern Alps. We were unable to resolve the phylogenetic affinities of O. suteri, New Zealand’s only native oviparous skink. We use the phylogeny and topology tests to resolve several taxonomic issues and assess the taxonomic status of several suspected undescribed taxa. We complete a generic revision for the New Zealand skink fauna, placing C. lichenigera and all native New Zealand species into a single genus.

Number of Maxillary Teeth in Scincid Lizards: Lineage Characteristics and Ecological Implications

Abstract The relationship between the mean number of maxillary teeth and the mean head length was investigated among 67 species of skinks, each represented by five large adults, and chosen to represent skink diversity. Only 52.4% of the potentially available morphological space defined by the four coordinates representing the ends of the range of these two variables is actually occupied by the datapoints for the species. There is no relationship between the two variables within skinks as a whole or within major lineages. The most primitive species falls close to the center of the cloud of datapoints. The ratio of the mean number of maxillary teeth to the mean head length is significantly greater in the Eugongylus group of eugongyline skinks than it is in the sphenomorphine skinks. Considering the number of teeth on other bones, these differences may be even greater than indicated by the number of teeth on only the maxilla. Because these two groups occur in broad sympatry in the Australian region, the diff...

Molecular phylogeny of the scincid lizards of New Caledonia and adjacent areas: Evidence for a single origin of the endemic skinks of Tasmantis

We use ∼1900 bp of mitochondrial (ND2) and nuclear (c-mos and Rag-1) DNA sequence data to recover phylogenetic relationships among 58 species and 26 genera of Eugongylus group scincid lizards from New Caledonia, Lord Howe Island, New Zealand, Australia and New Guinea. Taxon sampling for New Caledonian forms was nearly complete. We find that the endemic skink genera occurring on New Caledonia, New Zealand and Lord Howe Island, which make up the Gondwanan continental block Tasmantis, form a monophyletic group. Within this group New Zealand and New Zealand + Lord Howe Island form monophyletic clades. These clades are nested within the radiation of skinks in New Caledonia. All of the New Caledonian genera are monophyletic, except Lioscincus. The Australian and New Guinean species form a largely unresolved polytomy with the Tasmantis clade. New Caledonian representatives of the more widespread genera Emoia and Cryptoblepharus are more closely related to the non-Tasmantis taxa than to the endemic New Caledonian genera. Using ND2 sequences and the calibration estimated for the agamid Laudakia, we estimate that the diversification of the Tasmantis lineage began at least 12.7 million years ago. However, using combined ND2 and c-mos data and the calibration estimated for pygopod lizards suggests the lineage is 35.4–40.74 million years old. Our results support the hypothesis that skinks colonized Tasmantis by over-water dispersal initially to New Caledonia, then to Lord Howe Island, and finally to New Zealand.

Using ancient and recent DNA to explore relationships of extinct and endangered Leiolopisma skinks (Reptilia: Scincidae) in the Mascarene islands

Phylogenetic analysis, using 1455 bp of recent mtDNA (cytochrome b 714 bp, 12S rRNA 376 bp) and nuclear (c- mos 365 bp) sequence from 42 species and 33 genera of Scincidae, confirms Leiolopisma telfairii, now confined to Round island off Mauritius, is a member of the mainly Australasian Eugongylus group of the Lygosominae. Ancient mtDNA (cytochrome b 307 bp, 12S rRNA 376 bp) was also extracted from subfossils of two other Mascarene taxa that are now extinct: the giant L. mauritiana from Mauritius and Leiolopisma sp., known only from fragmentary remains from Réunion. Sequence divergences of 4.2–5.7% show that all three forms were distinct and form a clade. There is restricted evidence that L. mauritiana and L. sp. from Réunion were sister species. Monophyly and relationships suggest Leiolopisma arose from a single transmarine invasion of the oceanic Mascarene islands from Australasia, 5600–7000 km away. This origin is similar to that of Cryptoblepharus skinks and Nactus geckos in the archipelago but contrasts with Phelsuma day geckos, which appear to have arrived from Madagascar where Mascarene Cylindraspis tortoises may also have originated. Diversification of the known species of Leiolopisma occurred from about 2.3–3.4 Mya, probably beginning on Mauritius with later invasion of Réunion. The initial coloniser may have had a relatively large body-size, but L. mauritiana is likely to have become gigantic within the Mascarenes. Other relationships supported by this investigation include the following. Scincines: Pamelaescincus + Janetaescincus, and Androngo ( Amphiglossus, Paracontias). Lygosomines: Sphenomorphus group—( Sphenomorphus, Lipinia ( Ctenotus, Anomalopus ( Eulamprus and Gnypetoscincus))): Egernia group— Egernia ( Cyclodomorphus, Tiliqua); Eugongylus group—( Oligosoma, Bassiana. ( Lampropholis ( Niveoscincus, Carlia))).

CALCIUM TRANSPORT ACROSS THE UTERINE EPITHELIUM OF PREGNANT LIZARDS

Calcium is a major requirement for developing embryos, but an insufficient amount is present in egg-yolk to sustain embryonic development in most reptiles. Consequently, the eggshell acts as an extra-lectithal source of calcium in oviparous amniote vertebrates, including squamates. A number of anatomical and physiological changes must accompany the evolutionary transitions from oviparity to viviparity, including the reduction or elimination of the eggshell. In our quest to understand the evolution of viviparity, we have used indirect immunofluorescence to characterize Ca2 ATPase pumps in the uterus of a targeted range of species of Eugongylus group skinks and bird species at various stages of the reproductive cycle, including pregnancy. During egg-shelling, apical localisation of Ca2 ATPase pumps occurs in the glandular epithelium of the uterus in the oviparous lizard Lampropholis guichenoti and the King quail Coturnix chinensis, presumably to allow movement of calcium against its electrochemical gradient into the lumen of the glandular duct and ultimately to the uterine lumen and developing embryo. Interestingly, structurally similar shell glands containing Ca2 ATPase pumps occur in the uterine lamina propria (stroma) of viviparous species, although we have so far seen them only in early vitellogenic N. ocellatus and non-pregnant female Pseudemoia spenceri. Our findings suggest that the down regulation of Ca2 ATPase in the shell glands may be an important contributor to the evolution of viviparity through the loss of an eggshell.

A Phylogenetically Important Lygosomine Skink Resurrected from Taxonomic Obscurity: Lygosoma unilineatum de Rooij, 1915

Lygosoma unlineatum de Rooij 1915 is a large New Guinea skink known from five specimens. It was previously associated with taxa now considered to be in the Sphenomorphus group of lygosomine skinks. However, it is a member of the Eugongylus group of lygosomines, best placed in the relatively primitive genus Eugongylus. Within the Eugongylus group, the species has two unique primitive characters: nine premaxillary teeth (versus 211) and pterygoid teeth. The species also has one unique derived character: fusion of the first supralabial and the nasal scale. The two primitive characters make the species especially important in reconstructing the phylogeny of the Eugongylus group.

Placental ontogeny of the Australian scincid lizardsNiveoscincus coventryiandPseudemoia spenceri

Models for the evolution of placentation among squamate reptiles have been strongly influenced by early work on one lineage of Australian scincid lizards: one of three lineages that include species with specialized allantoplacentation. The discovery of two types of complex allantoplacentae among species currently assigned to the Eugongylus species group led Weekes ([1935] Proc. Zool. Soc. Lond., 2:625-645) to propose that a graded evolutionary sequence was exemplified by the morphological series of allantoplacental types that existed among Australian skinks. One of the model's strengths is the incorporation of testable inferences of the relationship between placental structure and function. However, descriptions of placental structure for some species are incomplete and subsequent taxonomic revisions have resulted in confusion concerning the species studied. One of the ambiguities is the identity of Lygosoma weekesae (Kinghorn [1929] Proc. Linn. Soc. N.S.W., 54:32-33), one of two species for which the most specialized type of allantoplacenta was defined. The distinguishing characteristic of this placental type is the presence of a structure known as a placentome. Subsequent to the original species description, Lygosoma weekesae was named a synonym for Pseudemoia spenceri (Rawlinson [1974] Mem. Natn. Mus. Vict., 35:87-96), yet the placental description (Weekes [1929] Proc. Linn. Soc. N.S.W., 54:34-60) is thought to be of Niveoscincus coventryi (Rawlinson [1975] Mem. Natn. Mus. Vic., 36:1-16). We describe placental ontogeny of N. coventryi and P. spenceri as a contribution to study of comparative placentation among lygosomatine skinks. We conclude that the placental description for L. weekesae could not have been N. coventryi, because a placentome is not a component of the allantoplacenta of N. coventryi. Further, the allantoplacenta of this species does not conform to previously defined categories for Eugongylus group skinks. The allantoplacenta of P. spenceri contains a placentome that is structurally congruous with the original description of placentation for L. weekesae (Weekes [1929] Proc. Linn. Soc. N.S.W., 54:34-60). Comparison of extraembryonic membrane and placental structure among the four viviparous and one oviparous species of Eugongylus group skinks for which data are available shows that each species exhibits characters that distinguish it from other species, while additional characters are shared among all, or some, species. Based on a recent phylogenetic hypothesis, the distribution of allantoplacental types among these species does not support Weekes' ([1935] Proc. Zool. Soc. Lond., 2:625-645) model of a graded sequence for the evolution of squamate placentation.

Evolution of reptilian placentation: Development of extraembryonic membranes of the Australian scincid lizards,Bassiana duperreyi (Oviparous) andPseudemoia entrecasteauxii (Viviparous)

A prominent model for the evolution of placentation among Reptilia is based on placental structure among species in the Eugongylus group of Australian lygosomatine skinks. We studied the development of the extraembryonic membranes of an oviparous species, Bassiana duperreyi, and a viviparous species, Pseudemoia entrecasteauxii, within this taxonomic group. We observed differences in the timing of development of shared features and in the structure of extraembryonic membrane epithelia in the two species. In the viviparous species, there is earlier vascularization of the yolk sâc and increased vascular support for the abembryonic yolk sac splanchnopleure. Structural differences between species result in partitioning of the egg into two distinct hemispheres and produce epithelia which appear functionally histotrophic in both the chorioallantoic membrane and the bilaminar omphalopleure of the viviparous species. We propose that the evolution of placentation in P. entrecasteauxii involved a combination of heterochrony and structural innovation. Further, because our interpretation of placental structure of this species provides new information relevant to placental function, we propose a revision of a classic model for the evolution of placentation among Reptilia. This model predicts specific relationships among reproductive characteristics and thus is testable by comparative analysis among other species within the Eugongylus group of Australian skinks. © 1996 Wiley-Liss, Inc.

The Ultrastructure of the Spermatozoa of Squamata—I. Scincidae, Gekkonidae and Pygopodidae (Reptilia)

Abstract Squamate autapomorphies seen in sperm of the Scincidae (e.g. Ctenotus robustus, Carlia pectoralis, Cryptoblepharus virgatus, and Lampropholis delicata) are penetration of the fibrous sheath of the axoneme into the midpiece, and the paracrystalline subacrosomal cone. Sphenomorphus group spermatozoa (e.g. Ctenotus) and the Egernia group (Tiliqua) differ from the more derived Eugongylus group (C. virgatus, L. delicata and C. pectoralis) in that the acrosome is elongate and apically depressed; the perforatorium is strongly oblique; the midpiece is relatively short, with four dense ring structures in longitudinal succession; mitochondria are columnar; and enlarged peripheral fibres 3 and 8 do not show the gross anterior enlargement seen in Carlia and Lampropholis. Heteronotia binoei (Gekkonidae) sperm have no epinuclear electron‐lucent region; nuclear shoulders are smooth, as in sphenomorph but not Eugongylus group skinks; mitochondria are columnar; unlike skinks, the median surfaces of the mitochondria are indented by triangular, sometimes longitudinally, interconnected dense bodies. In Lialis burtonis (Pygopodidae) sperm, the perforatorium extends virtually to the tip of the fore‐shortened apically domed acrosome; nuclear shoulders are absent; the mitochondria alternate singly or in groups with one or more dense bodies which also form an interrupted collar around the distal centriole. Spermatozoal ultrastructure suggests that a common ancestry of snakes and pygopods deserves consideration.

Karyotypes of Two Lygosomine Lizards of the Genus Emoia (Squamata: Scincidae) from Malaysia and Micronesia

Mitotic chromosomes of Emoia atrocostata from Sarawak, Malaysia, and E. caeruleocauda from Guam and Belau, Micronesia, are described for the first time for the genus. Both of the species have 2 n = 30 chromosomes, forming two distinct size-groups. The larger size-group accommodates four metacentric pairs. Of the smaller size-group, pairs 5, 8, and 10 are metacentric, pair 6 are submetacentric, and pairs 9 and 11 – 15 appear to be telocentric. In E. atrocostata pair 7 is submetacentric, whereas it is subtelocentric in E. caeruleocauda . The results seem to predict low chromosomal variability in the genus Emoia , and support Greer’s (1979) assignment of the genus to the Eugongylus Group.

Two New Species of Menetia from Northeastern Queensland, with Comments on the Generic Diagnoses of Lygisaurus and Menetia

The Australian skink genera Lygisaurus and Menetia are diagnosed on the basis of derived morphological characters. Menetia consists of the six species traditionally included therein (alanae, amaura, concinna, greyii, maini, and surda), plus three additional species (timlowi, koshlandae, and sadlieri). Menetia koshlandae and M. sadlieri are described as new and along with M. timlowi, most recently assigned to Lygisaurus, are shown to constitute a distinct subgroup within Menetia. A key is provided to all nine species of Menetia, and some notes are presented on reproduction, habitat and color for three species. Menetia is a group of small, ground-dwelling skinks occurring in arid, semi-arid, and seasonally dry habitats in Australia. Six species are currently recognized in the genus. Recently, a species originally described as M. timlowi by Ingram (1977), was reallocated to the genus Lygisaurus, a group of small, ground-dwelling skinks from mesic habitats in New Guinea and eastern Australia (Ingram and Covacevich, 1988). The reason for the reallocation was the realization that the species lacked the one uniquelyderived character of Menetia (supraoculars two instead of three or four) and its similarity to Lygisaurus burnettii. On re-examining a longknown but undescribed species of small skink from northeastern Queensland which closely resembles M. timlowi, I came to the conclusion that these two species are probably more closely related to Menetia than to Lygisaurus. Quite recently a single specimen of a second undescribed species also closely resembling M. timlowi has been discovered in the collections of the Queensland Museum. The purpose of this paper is to update the generic diagnoses of Lygisaurus and Menetia and to describe the two new species of Menetia. I also take the opportunity to provide some general notes on reproduction and habitat in three species of Menetia. MATERIALS AND METHODS The paravertebral scales were counted from the first scale falling wholly outside of an imagetia is a group of small, ground-dwelling i ks o curring in arid, semi-ari , and seasonl dry habi ats in Australia. Six species are rently recognized in the genus. Recently, a ecies originally described as M. timlowi by ram (1977), was realloca ed to the genus Lyisaurus, a group of small, ground-dwelling inary line drawn along the posterior edge of the thighs held perpendicular to the long axis of the body forward to and including the anteriormost nuchal. The supraand subdigital scales were counted on the fourth toe of the pes beginning in the former case with the first scale wholly on the toe, and in the latter case with the first transversely enlarged scale wholly on the toe. The posteriormost supraciliary was taken as the scale just lateral to the suture between the posteriormost supraocular and the anteriormost pretemporal. Nuchals were all the transversely enlarged paravertebral scales just posterior to the parietals. The presacral vertebrae were counted on X-rays. Limb length was measured along the posterior edge of the outstretched limb and including the claw. The polarity of character states was inferred by reference to the successively more distant outgroups: Eugongylus, the most generally primitive taxon in the Eugongylus group; southeast Asian Mabuya, the most generally primitive lygosomines, and Eumeces, the most generally primitive scincid (Greer, 1979). DIAGNOSES OF LYGISAURUS AND MENETIA B th Lygisaurus and Menetia are members of the Eugongylus group of lygosomine skinks (Greer, 1979), and both show the following derived character states vis a vis this group: supranasals absent; frontoparietals fused; postsupralabial single; external ear opening small; 268 This content downloaded from 157.55.39.243 on Thu, 06 Oct 2016 04:55:39 UTC All use subject to http://about.jstor.org/terms

Lankascincus, a New Genus of Scincid Lizards from Sri Lanka, with Descriptions of Three New Species

-A lineage of lygosomine skinks endemic to Sri Lanka is diagnosed and given generic rank with the name of Lankascincus. The group consists of six species, three of which are described as new. Brief accounts are provided for each species, and there is a key to aid their identification. As part of an ongoing review of the lygosomine scincid lizard assemblage it has become clear that amongst the Sphenomorphus of Sri Lanka there is an easily diagnosable, monophyletic group of six species, among which are three undescribed species. In this paper I diagnose this endemic Sri Lankan supraspecific group, give it generic rank, discuss its relationships, and describe the new species in the context of a brief review of all the species in the taxon. Collection abbreviations throughout follow Leviton et al. (1985), with the exception of the Australian Museum (AM, not AMS) and the addition of the E. H. TaylorH. M. Smith collection (EHT-HMS), now dispersed. Lankascincus new genus Type Species.-Lygosoma fallax Peters, 1860:184; designated herein. Diagnosis.-Compared to other lygosomine skinks, the skinks of this genus share the following combination of derived character states: size small (maximum SVL, 58 mm); nuchals undifferentiated; subocular scale row complete and non-interdigitating with supralabials; external ear opening reduced, approximately three times size of narial opening; postmental contacts first infralabial only; third pair of chin scales separated from infralabial row; outer preanals overlap inner; scales perfectly smooth; subdigital lamellae with postaxial groove basally; premaxillary teeth >11; palatal rami of pterygoids slightly expanded posteromedially; fat bodies absent; clutch size constant, two or one, depending on species. For a general description of the species in the taxon see Taylor (1950) and Deraniyagala (1953), and for some general morphometrics see Table 1. Referred Species (6).-Sphenomorphus deignani Taylor, 1950; Lankascincus deraniyagalae, new species; Lygosoma fallax Peters, 1861; Lankascincus gansi, new species; Eumeces taprobanense Kelaart, 1854; Lankascincus taylori, new species. Distribution and Life History.--The genus is endemic to Sri Lanka, occurring in both the lowlands and the highlands. It is diurnal and terrestrial. It is oviparous, and the clutch size is a constant one or two, depending on species. Relationships. -Although the skinks discussed here have previously been placed in the genus Sphenomorphus, it is uncertain whether they belong to this major group of lygosomines or to the Eugongylus group (Greer, 1979). Of the group characters that can be assessed in Lankascincus, it possesses three of the Eugongylus group characters (premaxillary teeth 11; inner preanals overlapped by outer-Greer, 1990; scales on dorsal surface of digits mostly in single row), and none of the group characters. However, the hemipenis character that is the single most important feature of the group (deeply bifurcate) has not yet been examined in Lankascincus due to lack of suitable material. Needless to say it would be highly desirable to know the morphology of the hemipenis of Lankascincus. Regardless of its major group relationships, Lankascincus may be most closely related to the southern Indian genus Ristella, a group whose own major group relationships are also unclear for the same reasons as apply to Lankascincus. Both taxa share a larger number of derived character states with each other than with any other lygosomine, including small size, undifferentiated nuchals, a complete subocular scale row, reduced external ear opening, postmental in contact with only one infralabial on each side, and slightly posteromedially expanded palatal rami of the pterygoids. Lankascincus differs from Ristella in having the derived character states: scales smooth, lower secondary temporal bordered posteriorly by two tertiary temporals instead of one, third pair of chin scales separated from infralabials, and clutch size a constant two or one. Ristella differs from Lankascincus in having the derived character states: supraoculars contacted by frontal three instead of two, digits on the manus four, and claws retractile. The This content downloaded from 157.55.39.45 on Thu, 01 Sep 2016 05:28:41 UTC All use subject to http://about.jstor.org/terms

Immunological Relationships and Generic Revision of the Australian Lizards Assigned to the Genus Leiolopisma (Scincidae, Lygosominae)

The phylogenetic relationships of the Australian scincid lizards currently assigned to the genus Leiolopisma have been examined by quantitative micro-complement fixation (MC'F) comparisons of serum albumin. The results of these comparisons do not support the monophyly implicit in these species' current congeneric status, but suggest instead that the Australian species of Leiolopisma belong to several distinct phyletic lineages within the Eugongylus group. These findings are supported by several sets of non-biochemical characters, including features of scalation, osteology and karyotype. None of the Australian species shares a close relationship with the type-species of Leiolopisrna (L. telfairii), and so a new taxonomic arrangement is proposed which distributes them among the following genera: Bartleia, gen. nov. (jigurru); Bassiana, gen. nov. (duperreyi, platynotum and trilineata); Cautula, gen. nov. (zia); Niveoscincus, gen. nov. (coventryi, greeni, metallicus, microlepidotus, ocellatus, orocryptus, palfreymani and pretiosus); and Pseudemoia Fuhn, 1967 (baudini, entrecasteauxii Group 1; entrecasteauxii Group 2, rawlinsoni and spenceri). Preliminary comparisons suggest that other Leiolopisma species, from New Caledonia, Lord Howe I. and New Zealand, belong to phyletic lineages which are distinct from any of the Australian 'Leiolopisma' and from the type-species.