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We show that the Wotjulum frog, Litoria watjulumensis (Copland, 1957), comprises two deeply divergent mitochondrial DNA lineages that are also reciprocally monophyletic for a nuclear gene locus and have discrete distributions. The taxa are differentiated in multivariate analysis of shape but show no appreciable differences in colour and pattern. The two taxa differ substantially in the degree of female biased sexual size dimorphism, with the western taxon showing considerably more pronounced dimorphism. We subsequently resurrect Litoria (Hyla) spaldingi (Hosmer, 1964) for populations from east of the Daly River system in the Northern Territory through to western Queensland and restrict L. watjulumensis to populations from the Kimberley region of north-western Australia and the Victoria River system of the western Northern Territory. The complex advertisement call of L. spaldingi is described for the first time.

Subspecies designations for herpetofauna in Western Australia were largely coined in the 20th century where rigorous evolutionary concepts to species were not consistently applied. Rather, subspecies tended to designate geographic populations of similar-looking taxa to nominate forms, usually differing in size, pattern or colour and, at best, a few scalation differences. Here we re-evaluate two pygopodoid taxa from Western Australia using a combination of published and original genetic data coupled with a reassessment of morphology. We review these differences in light of an integrative taxonomic approach that looks to find multiple independent lines of evidence to establish the evolutionary independence of populations. For the pygopod species Pletholax gracilis, we found consistent diagnostic characters (e.g. body size, visibility of ear opening, scalation) and a deep genetic divergence between the two subspecies. We therefore raise each subspecies to full species: P. gracilis and P. edelensis. The two subspecies of the carphodactylid gecko Nephrurus wheeleri were also assessed, and we found strong genetic and morphological evidence (e.g. body size, scalation, pattern) to raise these to full species: N. wheeleri and N. cinctus. By revisiting Storr’s morphological insights and newly acquired genetic evidence, in addition to a thorough re-examination of morphological traits, our study provides a robust foundation to raise Storr’s morphological subspecies into full species based upon multiple lines of evidence. Such an approach applied to other subspecies in the Australian herpetofauna also may result in revised taxonomies.

We present information on primary type specimens for 13,282 species and subspecies of reptiles compiled in the Reptile Database, that is, holotypes, neotypes, lectotypes, and syntypes. These represent 99.4% of all 13,361 currently recognized taxa (11,050 species and 2311 subspecies). Type specimens of 653 taxa (4.9%) are either lost or not located, were never designated, or we did not find any information about them. 51 species are based on iconotypes. To map all types to physical collections we have consolidated all synonymous and ambiguous collection acronyms into an unambiguous list of 364 collections holding these primary types. The 10 largest collections possess more than 50% of all (primary) reptile types, the 36 largest collections possess more than 10,000 types and the largest 73 collections possess over 90% of all types. Of the 364 collections, 107 hold type specimens of only 1 species or subspecies. Dozens of types are still in private collections. In order to increase their utility, we recommend that the description of type specimens be supplemented with data from high-resolution images and CT-scans, and clear links to tissue samples and DNA sequence data (when available). We request members of the herpetological community provide us with any missing type information to complete the list.

For over two decades, assessments of geographic variation in mtDNA and small numbers of nuclear loci have revealed morphologically similar, but genetically divergent, intraspecific lineages in lizards from around the world. Subsequent morphological analyses often find subtle corresponding diagnostic characters to support the distinctiveness of lineages, but occasionally do not. In recent years it has become increasingly possible to survey geographic variation by sequencing thousands of loci, enabling more rigorous assessment of species boundaries across morphologically similar lineages. Here we take this approach, adding new, geographically extensive SNP data to existing mtDNA and exon capture datasets for the Gehyra australis and G. koira species complexes of gecko from northern Australia. The combination of exon-based phylogenetics with dense spatial sampling of mitochondrial DNA sequencing, SNP-based tests for introgression at lineage boundaries and newly-collected morphological evidence supports the recognition of nine species, six of which are newly described here. Detection of discrete genetic clusters using new SNP data was especially convincing where candidate taxa were continuously sampled across their distributions up to and across geographic boundaries with analyses revealing no admixture. Some species defined herein appear to be truly cryptic, showing little, if any, diagnostic morphological variation. As these SNP-based approaches are progressively applied, and with all due conservatism, we can expect to see a substantial improvement in our ability to delineate and name cryptic species, especially in taxa for which previous approaches have struggled to resolve taxonomic boundaries.

Global biodiversity loss is a profound consequence of human activity. Disturbingly, biodiversity loss is greater than realized because of the unknown number of undocumented species. Conservation fundamentally relies on taxonomic recognition of species, but only a fraction of biodiversity is described. Here, we provide a new quantitative approach for prioritizing rigorous taxonomic research for conservation. We implement this approach in a highly diverse vertebrate group—Australian lizards and snakes. Of 870 species assessed, we identified 282 (32.4%) with taxonomic uncertainty, of which 17.6% likely comprise undescribed species of conservation concern. We identify 24 species in need of immediate taxonomic attention to facilitate conservation. Using a broadly applicable return-on-investment framework, we demonstrate the importance of prioritizing the fundamental work of identifying species before they are lost., In order to inform conservation effort, there is urgent need for rigorous taxonomic research to describe species under threat of extinction. Implementation of a new prioritization method identified 282 Australian reptile species needing taxonomic research, of which 17.6% represent undescribed species of conservation concern; this approach could be readily implemented across many faunal groups.

The methods used to detect and describe morphologically cryptic species have advanced in recent years, owing to the integrative nature of molecular and morphological techniques required to elucidate them. Here we integrate recent phylogenomic work that sequenced many genes but few individuals, with new data from mtDNA and morphology from hundreds of gecko specimens of the Gehyra variegata group from the Australian arid zone. To better understand morphological and geographical boundaries among cryptic forms, we generated new sequences from 656 Gehyra individuals, largely assigned to G. variegata group members over a wide area in Western Australia, with especially dense sampling in the Pilbara region, and combined them with 566 Gehyra sequences from GenBank, resulting in a dataset of 1,222 specimens. Results indicated the existence of several cryptic species, from new species with diagnostic morphological characters, to cases when there were no useful characters to discriminate among genetically distinctive species. In addition, the cryptic species often showed counter-intuitive distributions, including broad sympatry among some forms and short range endemism in other cases. Two new species were on long branches in the phylogram and restricted to the northern Pilbara region: most records of the moderately sized G. incognita sp. nov. are near the coast with isolated inland records, whereas the small-bodied saxicoline G. unguiculata sp. nov. is only known from a small area in the extreme north of the Pilbara. Three new species were on shorter branches in the phylogram and allied to G. montium. The moderately sized G. crypta sp. nov. occurs in the western and southern Pilbara and extends south through the Murchison region; this species was distinctive genetically, but with wide overlap of characters with its sister species, G. montium. Accordingly, we provide a table of diagnostic nucleotides for this species as well as for all other species treated here. Two small-bodied species occur in isolated coastal regions: G. capensis sp. nov. is restricted to the North West Cape and G. ocellata sp. nov. occurs on Barrow Island and other neighbouring islands. The latter species showed evidence of introgression with the mtDNA of G. crypta sp. nov., possibly due to recent connectivity with the mainland owing to fluctuating sea levels. However, G. ocellata sp. nov. was more closely related to G. capensis sp. nov. in the phylogenomic data and in morphology. Our study illustrates the benefits of combining phylogenomic data with extensive screens of mtDNA to identify large numbers of individuals to the correct cryptic species. This approach was able to provide sufficient samples with which to assess morphological variation. Furthermore, determination of geographic distributions of the new cryptic species should greatly assist with identification in the field, demonstrating the utility of sampling large numbers of specimens across wide areas.

The scincid lizard genus Ctenotus is one of the most species-rich genera of squamate reptiles, but few molecular phylogenetic studies have been undertaken on the group. Here we assess molecular and morphological variation within C. piankai and C. duricola, an arid-adapted pair of nominate species characterized by a pattern of thin pale longitudinal lines on a dark background that occur primarily in the western deserts and Pilbara region of Australia. We sequenced mitochondrial DNA and genome-wide restriction-site associated DNA (RAD) from geographically widespread samples of these lizard taxa, with particularly dense sampling from the Pilbara region. Phylogenetic analyses of the mitochondrial cytochrome B gene and approximately 5,000 nuclear loci identified four highly divergent lineages within the two taxa. The four genetically-defined populations were concordant with geography and are distinguishable based on multiple morphological and color pattern characters, despite appearing superficially similar in appearance. Despite limited mtDNA exchange between two lineages in the Pilbara, we found no evidence for ongoing gene flow across the nuclear genome. For the western desert lineages, there was no evidence of introgression for either mtDNA or nDNA in our data. To resolve the taxonomy of the group, we redescribe C. piankai and C. duricola, and recognize the two divergent lineages as new species: C. rhabdotus sp. nov., from the south-eastern Kimberley, Ord, Victoria River and northern Tanami Desert regions of Western Australia and the Northern Territory, and C. pallasotus sp. nov., from the western Pilbara and North West Cape regions of Western Australia.

The worm-like snakes (Scolecophidia; approximately 400 nominal extant species) have a conservative morphology and are among the most poorly-known terrestrial vertebrates. Although molecular evidence has helped determine their higher-level relationships, such data have rarely been used to discriminate among species. We generated a molecular data set for the continental Australian blindsnakes (genus Ramphotyphlops) to determine the concordance of molecular and morphological information in the taxonomic recognition of species. Our dataset included 741 specimens morphologically attributed to 27 nominal Ramphotyphlops species. We proposed species hypotheses (SHs) after analysis of sequences from a variable mitochondrial gene (cytochrome b) and examined these SHs with additional evidence from a nuclear gene (prolactin receptor) and geographical data. Although the nuclear marker was not as fast-evolving and discriminating as the mitochondrial marker, there was congruence among the mitochondrial, nuclear, and geographical data, suggesting that the actual number of species is at least two times the current number of recognized, nominal species. Several biogeographical barriers and complex phytogeographical and geological patterns appeared to be involved in the division of some burrowing snake populations and, by consequence, in their diversification and speciation through isolation. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110, 427–441.

The methods used to detect and describe morphologically cryptic species have advanced in recent years, owing to the integrative nature of molecular and morphological techniques required to elucidate them. Here we integrate recent phylogenomic work that sequenced many genes but few individuals, with new data from mtDNA and morphology from hundreds of gecko specimens of theGehyra variegatagroup from the Australian arid zone. To better understand morphological and geographical boundaries among cryptic forms, we generated new sequences from 656Gehyraindividuals, largely assigned toG. variegatagroup members over a wide area in Western Australia, with especially dense sampling in the Pilbara region, and combined them with 566Gehyrasequences from GenBank, resulting in a dataset of 1,222 specimens. Results indicated the existence of several cryptic species, from new species with diagnostic morphological characters, to cases when there were no useful characters to discriminate among genetically distinctive species. In addition, the cryptic species often showed counter-intuitive distributions, including broad sympatry among some forms and short range endemism in other cases. Two new species were on long branches in the phylogram and restricted to the northern Pilbara region: most records of the moderately sizedG. incognitasp. nov. are near the coast with isolated inland records, whereas the small-bodied saxicolineG. unguiculatasp. nov. is only known from a small area in the extreme north of the Pilbara. Three new species were on shorter branches in the phylogram and allied toG. montium. The moderately sizedG. cryptasp. nov. occurs in the western and southern Pilbara and extends south through the Murchison region; this species was distinctive genetically, but with wide overlap of characters with its sister species,G. montium. Accordingly, we provide a table of diagnostic nucleotides for this species as well as for all other species treated here. Two small-bodied species occur in isolated coastal regions:G. capensissp. nov. is restricted to the North West Cape andG. ocellatasp. nov. occurs on Barrow Island and other neighbouring islands. The latter species showed evidence of introgression with the mtDNA ofG. cryptasp. nov., possibly due to recent connectivity with the mainland owing to fluctuating sea levels. However,G. ocellatasp. nov. was more closely related toG. capensissp. nov. in the phylogenomic data and in morphology. Our study illustrates the benefits of combining phylogenomic data with extensive screens of mtDNA to identify large numbers of individuals to the correct cryptic species. This approach was able to provide sufficient samples with which to assess morphological variation. Furthermore, determination of geographic distributions of the new cryptic species should greatly assist with identification in the field, demonstrating the utility of sampling large numbers of specimens across wide areas.

The optimal division of resources into offspring size vs. number is one of the classic problems in life-history evolution. Importantly, models that take into account the discrete nature of resource division at low clutch sizes suggest that the variance in offspring size should decline with increasing clutch size according to an invariant relationship. We tested this prediction in 12 species of lizard with small clutch sizes. Contrary to expectations, not all species showed a negative relationship between variance in offspring size and clutch size, and the pattern significantly deviated from quantitative predictions in five of the 12 species. We suggest that the main limitation of current size-number models for small clutch sizes is that they rely on assumptions of hierarchical allocation strategies with independence between allocation decisions. Indeed, selection may favour alternative mechanisms of reproductive allocation that avoid suboptimal allocation imposed by the indivisible fraction at low clutch sizes.

A wide range of organisms use chemical and visual cues in mate attraction and courtship; however, chemical discrimination relevant to reproduction and the interplay between these two types of communication are poorly understood in reptiles. We experimentally tested the ability of male Eulamprus heatwolei, a scincoid lizard, to discriminate between sexually receptive and non-receptive females in two ways. First, we conducted 155 staged encounters between males and females over 29 days to determine the start and the duration of the female receptive period based on the date of copulations. These data suggest that the receptive period lasted for approximately 7 days in late October under controlled laboratory conditions. We also recorded 6,330 individual male and female behaviours during these trials to evaluate the frequency of female courtship and rejection behaviours and the intensity of male courtship behaviour. Female courtship increased sharply during the receptive period and then diminished. The disparity between female courtship behaviours and female rejection behaviours was greatest during the receptive period. Male courtship intensity increased sharply prior to the receptive period, peaked during the receptive period and thereafter declined rapidly. To determine if males were using visual cues, chemical cues or both from females, we conducted an experiment during and after the receptive period wherein male lizards were presented with a choice of two retreat sites treated either with the odour of large sexually receptive females, odour of small sexually non-receptive females or no odour (control). Males preferred the scent of females over the odourless control, and analysis using a special form of a generalized linear model, the Bradley–Terry model, showed a clear order of retreat site preferences, with large sexually receptive females favoured over small non-receptive females over the odourless control. We speculate that males use vision to find females and then use their chemosensory ability to chemically evaluate female sexual receptivity once the pair are in close proximity.

Developmental plasticity and large egg size are traits that can benefit offspring early in life, but their coevolution has rarely been examined. I experimentally manipulated water depth and food availability of larvae of the frog Crinia georgiana, a species with a large egg size that breeds in shallow seeps that often dry before tadpoles have metamorphosed. Tadpoles that were never fed completed metamorphosis at about the same time as tadpoles under constant conditions, although at much smaller body sizes and with higher mortality. Except for the earliest stage tested (Gosner stage 28), tadpoles that experienced a decrease in water depth accelerated development and completed metamorphosis earlier than tadpoles kept in constant conditions. Acceleration was not possible when food was withheld, except when water levels were also lowered and only during later developmental stages. Because acceleration of development was initiated at earlier larval stages than in small-egged species that breed in ephe...

The blindsnake genus Anilios (formerly Ramphotyphlops ) is the largest and most diverse genus of snakes in Australia with 45 currently recognized species. Recent molecular genetic studies of the genus have identified high levels of cryptic diversity within many taxa, suggesting true species diversity is greatly underestimated. Anilios leptosoma is a slender blindsnake endemic to the mid-west of Western Australia. Although morphological variation has been identified within the species in the past, the systematics and true diversity remained unstudied. Here we use recent molecular data to guide a reappraisal of morphology in order to provide a taxonomic revision of the A. leptosoma species complex. We redescribe A. leptosoma and describe two new species that occur to the south of most of true A. leptosoma ’s distribution: A. systenos sp. nov. and A. obtusifrons sp. nov. Anilios systenos sp. nov. is known from the Geraldton region with the furthest record only 100 km to the north-east, a very small range for a species of snake. Anilios obtusifrons sp. nov. has an even smaller distribution, as it is only known from a small coastal area south of Kalbarri and may represent a range-restricted taxa. All species are genetically divergent from each other and can be distinguished by consistent morphological characteristics, including the shape of the snout, the termination point of the rostral cleft and number of mid-body scale rows and ventral scales.

Multiple paternity has rarely been reported in anuran amphibians, with only three previous documented examples. For the Australian frog Crinia georgiana, we observed synchronous polyandry in an average of 44% of matings observed at four field sites. This suggests matings involving more than one male are common in this species. One to eight males were observed in amplectant groups with second males amplexed ventrally. Genetic analyses, using allozyme electrophoresis, of offspring from two matings indicated that at least two of three possible males fathered offspring. Third males were unlikely to have shared paternity, explained by their inappropriate position during amplexus. Multiple paternity may be more common in frogs than has been reported. Copyright 1999 The Association for the Study of Animal Behaviour.

Life-history models suggest that ‘costs of reproduction’ can influence the evolution of sexual dimorphism, but empirical data on this effect are scarce. We tested the idea using ‘flying lizards’ (Asian agamids of the genus Draco), because the evolution of ‘flight’ (gliding) is likely to have increased the degree to which pregnancy affects maternal locomotor ability (and hence, we infer, has increased the ‘costs of reproduction’). As predicted, Draco display patterns of sexual dimorphism that are different from those seen in most other lizards. The ‘wings’ (gliding membranes) of female D. melanopogon are significantly larger than those of males of the same body size, a dimorphism that we attribute to natural selection on the ability to glide while encumbered with eggs. Comparisons with non-flying lizards suggest that the same selective pressure has also reversed pre-existing patterns of sexual dimorphism in body size and body proportions (relative head size, relative tail length). Measurements of wing-loading, centre of balance and gliding distances of preserved specimens support the hypothesis that a larger body, relatively larger head, and longer tail may improve a female's aerial mobility when she is gravid. Thus, selection for locomotor ability in egg-burdened females (reduction in ‘costs of reproduction’) may have been responsible for a reversal of sexually-selected traits in this lineage.

The isolated uplands of the Australian arid zone are known to provide mesic refuges in an otherwise xeric landscape, and divergent lineages of largely arid zone taxa have persisted in these regions following the onset of Miocene aridification. Geckos of the genus Heteronotia are one such group, and have been the subject of many genetic studies, including H. spelea, a strongly banded form that occurs in the uplands of the Pilbara and Central Ranges regions of the Australian arid zone. Here we assess the systematics of these geckos based on detailed examination of morphological and genetic variation. The H. spelea species complex is a monophyletic lineage to the exclusion of the H. binoei and H. planiceps species complexes. Within the H. spelea complex, our previous studies based on mtDNA and nine nDNA loci found populations from the Central Ranges to be genetically divergent from Pilbara populations. Here we supplement our published molecular data with additional data gathered from central Australian samples. In the spirit of integrative species delimitation, we combine multi-locus, coalescent-based lineage delimitation with extensive morphological analyses to test species boundaries, and we describe the central populations as a new species, H. fasciolatus sp. nov. In addition, within the Pilbara there is strong genetic evidence for three lineages corresponding to northeastern (type), southern, and a large-bodied melanic population isolated in the northwest. Due to its genetic distinctiveness and extreme morphological divergence from all other Heteronotia, we describe the melanic form as a new species, H. atra sp. nov. The northeastern and southern Pilbara populations are morphologically indistinguishable with the exception of a morpho-type in the southeast that has a banding pattern resembling H. planiceps from the northern monsoonal tropics. Pending more extensive analyses, we therefore treat Pilbara H. spelea as a single species with phylogenetic structure and morphological heterogeneity.

Recent work on species complexes of the pebble-mimic dragons of the Australian genus Tympanocryptis has greatly clarified evolutionary relationships among taxa and also indicated that species diversity has been severely underestimated. Here we provide a morphological and molecular appraisal of variation in the T. cephalus species-group and find evidence for recognizing five species-level lineages from Western Australia. Four species-level lineages are strongly supported with a combined mitochondrial and nuclear DNA Bayesian analysis (a fifth population from the Gascoyne region lacked tissue samples). Morphologically, we found subtle, yet consistent, differences among the populations in scalation, color and pattern. True T. cephalus Gunther is restricted to the coastal Pilbara region and characterized by five dark blotches on the dorsum, keeled ventrals, and other characters. Two other lineages within the Pilbara, from the Hamersley range and Fortescue/northern Pilbara region, differed from T. cephalus senso stricto by possessing a more elongate body and a plain dorsum. Furthermore, the Hamersley lineage differed from the Fortescue lineage by possessing slightly more reddish coloration and feeble keeling on the snout. Although there are few specimens and no tissue samples available for the Gascoyne population, these individuals are larger, have rugose scales on the snout, and possess scattered enlarged tubercles with three large blotches on the dorsum. The name T. cephalus gigas Mitchell is available for this population. The most widespread lineage, and the one best represented in collections and in field guides, occurs throughout central Western Australia. These Goldfield populations are characterized by a protruding snout, narrow rostral, and uniform reddish-brown coloration, often with a dark wash. Based on the genetic and morphological differences, we redescribe T. cephalus, resurrect and elevate T. gigas to a full species and designate a neotype for this taxon, and describe three lineages as new species (T. diabolicus sp. nov., T. fortescuensis sp. nov., T. pseudopsephos sp. nov.).

We present an assessment of the systematic status of populations currently assigned to Eremiascincus fasciolatus across Australia using morphological data. Our analyses show that these populations actually comprise four species: a large and robust, narrow-banded taxon from eastern Queensland and three slender, desert-dwelling taxa from the western, northern, and south-eastern Australian arid zone respectively. These data necessitate: 1) the redescription of E . fasciolatus , a species endemic to the dry forests of eastern Queensland, 2) the resurrection of E . pallidus for the western taxon, 3) the resurrection of E . intermedius for the northern taxon, and 4) the description of the south-eastern desert-dwelling taxon, which largely occurs in the Lake Eyre Basin, as a new species. The new species, Eremiascincus phantasmus sp. nov., can be distinguished from congeners by its large size, pale color, depressed snout, and small circular ear opening, as well as by several scale characteristics. Body proportions, a depressed snout, small ear opening, and pale color suggest that this species is fossorial and predominantly occurs in habitats with loose substrate, such as dune systems and sandy plains. A synopsis of each species and a key to the narrow-banded species of Eremiascincus are provided.

Egernia depressa is an extremely spiny species of scincid lizard that occurs in several populations with highly variable morphology in western Australia. Using a combination of fi xed morphological character differences and mitochondrial DNA sequence data, we found evidence for four species level groups within the complex. We restrict E. depressa to the log-inhabiting population from south-western Australia and resdescribe the species, and describe three new species from the arid zone: two from the Pilbara and one from the central ranges. In addition to the genetic differences, the species differ in head size, limb length, tail shape, colouration and scalation. Many of the morphological characters appear to be adaptations to log or rock-dwelling, with the log-dwelling E. depressa having brown colouration, large head, limbs and tail and long thin spines on the body and tail. The two Pilbara species are not each other's closest relatives, yet they resemble each other the closest, probably owing to a suite of characters adapted for living in rock crevices such as yellow to reddish colouration, smaller head and limbs, narrower tail and short strong spines on the body and tail. The central ranges species appears to have a combination of characters from log and rock-dwelling forms and is the most isolated of the four species.

Techniques of offspring size manipulation, "allometric engineering," were used in combination with studies of natural selection to elucidate the causal relation between egg size and offspring survival of lizards. The results experimentally validate premises underlying theories of optimal egg size: fecundity selection favoring the production of large clutches of small eggs was balanced by survival selection favoring large offspring. However, large hatchlings did not always have the highest survival, contrary to most theoretical expectations. Optimizing selection on offspring size per se was the most common pattern. Moreover, matches between average and optimal egg size were qualitative, not quantitative, perhaps reflecting known functional constraints on the production of large eggs.

Although the south-western Australian region is recognised as a global biodiversity hotspot, there are still significant gaps in our understanding of the biodiversity of this region. We present a phylogenetic study of the heath dragons (Rankinia adelaidensis and R. parviceps) from this region, incorporating a 1612-bp section of mtDNA and two nuclear introns, Gapdh (~244 bp) and Enol (~330 bp). In addition, we present a generic-level analysis of three gene regions (mtDNA, Gapdh, BDNF), which provides clear evidence that Rankinia adelaidensis and R. parviceps are not closely related to Rankinia diemensis from eastern Australia. Instead, the heath dragons are strongly supported as forming a clade with the genus Ctenophorus. In addition, we find that there are significant levels of haplotype divergence between currently recognised subspecies of the heath dragons (R. a. adelaidensis, R. a. chapmani, R. p. parviceps, R. p. butleri). We suggest that the genetic divergences between subspecies result from geographic isolation in allopatry owing to habitat preferences, followed by drift and/or selection. On the basis of these deep divergences and consistent morphological differences between subspecies, we recommend elevating all taxa to full species, and provide a taxonomic revision of the genera Rankinia and Ctenophorus.

Adaptive phenotypic plasticity in life-history traits is predicted to evolve when populations occur in heterogeneous environments. Anuran larvae of many species cannot escape their aquatic environment until metamorphosis and therefore should show plasticity in response to conditions experienced as tadpoles. In this study, we manipulated the aquatic environments of Crinia georgiana tadpoles in the laboratory to mimic variation among ponds in resources and drying conditions in nature. This species breeds in very shallow water in winter and ponds frequently dry between bouts of rain, especially towards spring. Tadpoles kept in constant conditions at different levels of food metamorphosed at different body sizes but showed no plasticity in metamorphic timing. Tadpoles fed only lettuce metamorphosed at sizes similar to those of field-collected tadpoles, whereas tadpoles fed a more protein-rich food metamorphosed at unusually large sizes, indicating that the seeps where C. georgiana tadpoles occur are poor in nutrients. When we decreased food and water levels, tadpoles at later developmental stages were able to accelerate development and metamorphose earlier than tadpoles kept under constant conditions. Furthermore, tadpoles in very shallow water with no access to food metamorphosed earlier and at smaller body sizes than tadpoles with a more moderate decrease in depth that were able to continue feeding. Rapid development and the ability to accelerate metamorphosis in C. georgiana tadpoles are consistent with adaptation in a heterogeneous environment where larvae are under strong time constraints.

We demonstrate that egg size in side-blotched lizards is heritable (parent-offspring regressions) and thus will respond to natural selection. Because our estimate of heritability is derived from free-ranging lizards, it is useful for predicting evolutionary response to selection in wild populations. Moreover, our estimate for the heritability of egg size is not likely to be confounded by nongenetic maternal effects that might arise from egg size per se because we estimate a significant parent-offspring correlation for egg size in the face of dramatic experimental manipulation of yolk volume of the egg. Furthermore, we also demonstrate a significant correlation between egg size of the female parent and clutch size of her offspring. Because this correlation is not related to experimentally induced maternal effects, we suggest that it is indicative of a genetic correlation between egg size and clutch size. We synthesize our results from genetic analyses of the trade-off between egg size and clutch size with previously published experiments that document the mechanistic basis of this trade-off. Experimental manipulation of yolk volume has no effect on offspring reproductive traits such as egg size, clutch size, size at maturity, or oviposition date. However, egg size was related to offspring survival during adult phases of the life history. We partitioned survival of offspring during the adult phase of the life history into (1) survival of offspring from winter emergence to the production of the first clutch (i.e., the vitellogenic phase of the first clutch), and (2) survival of the offspring from the production of the first clutch to the end of the reproductive season. Offspring from the first clutch of the reproductive season in the previous year had higher survival during vitellogenesis of their first clutch if these offspring came from small eggs. We did not observe selection during these prelaying phases of adulthood for offspring from later clutches. However, we did find that later clutch offspring from large eggs had the highest survival over the first season of reproduction. The differences in selection on adult survival arising from maternal effects would reinforce previously documented selection that favors the production of small offspring early in the season and large offspring later in the season-a seasonal shift in maternal provisioning. We also report on a significant parent-offspring correlation in lay date and thus significant heritable variation in lay date. We can rule out the possibility of yolk volume as a confounding maternal effect-experimental manipulation of yolk volume has no effect on lay date of offspring. However, we cannot distinguish between genetic effects (i.e., heritable) and nongenetic maternal effects acting on lay date that arise from the maternal trait lay date per se (or other unidentified maternal traits). Nevertheless, we demonstrate how the timing of female reproduction (e.g., date of oviposition and date of hatching) affect reproductive attributes of offspring. Notably, we find that date of hatching has effects on body size at maturity and fecundity of offspring from later clutches. We did not detect comparable effects of lay date on offspring from the first clutch.

Background Molecular studies have revealed that many putative 'species' are actually complexes of multiple morphologically conservative, but genetically divergent 'cryptic species'. In extreme cases processes such as non-adaptive diversification (speciation without divergent selection) could mask the existence of ancient lineages as divergent as ecologically and morphologically diverse radiations recognised as genera or even families in related groups. The identification of such ancient, but cryptic, lineages has important ramifications for conservation, biogeography and evolutionary biology. Herein, we use an integrated multilocus genetic dataset (allozymes, mtDNA and nuclear DNA) to test whether disjunct populations of the widespread nominal Australian gecko species Crenadactylus ocellatus include distinct evolutionary lineages (species), and to examine the timing of diversification among these populations. Results We identify at least 10 deeply divergent lineages within the single recognised species Crenadactylus ocellatus, including a radiation of five endemic to the Kimberley region of north-west Australia, and at least four known from areas of less than 100 km2. Lineages restricted to geographically isolated ranges and semi-arid areas across central and western Australia are estimated to have began to diversify in the late Oligocene/early Miocence (~20-30 mya), concurrent with, or even pre-dating, radiations of many iconic, broadly sympatric and much more species-rich Australian vertebrate families (e.g. venomous snakes, dragon lizards and kangaroos). Conclusions Instead of a single species, Crenadactylus is a surprisingly speciose and ancient vertebrate radiation. Based on their deep divergence and no evidence of recent gene flow, we recognise each of the 10 main lineages as candidate species. Molecular dating indicates that the genus includes some of the oldest vertebrate lineages confounded within a single species yet identified by molecular assessments of diversity. Highly divergent allopatric lineages are restricted to putative refugia across arid and semi-arid Australia, and provide important evidence towards understanding the history and spread of the Australian arid zone, suggesting at a minimum that semi-arid conditions were present by the early Miocene, and that severe aridity was widespread by the mid to late Miocene. In addition to documenting a remarkable instance of underestimation of vertebrate species diversity in a developed country, these results suggest that increasing integration of molecular dating techniques into cryptic species delimitation will reveal further instances where taxonomic conservatism has led to profound underestimation of not only species numbers, but also highly significant phylogenetic diversity and evolutionary history.