A phylogeny of Chinese species in the genus Phrynocephalus (Agamidae) inferred from mitochondrial DNA sequences

We investigated the phylogenetic relationships among most Chinese species of lizards in the genus Phrynocephalus (118 individuals, collected from 56 populations of 14 well-defined species and several unidentified specimens) using four mitochondrial gene fragments (12S rRNA, 16S rRNA, cytochrome b, and ND4-tRNA(LEU)). The partition-homogeneity tests indicated that the combined dataset was homogeneous, and maximum-parsimony (MP), neighbor-joining (NJ), maximum-likelihood (ML) and Bayesian (BI) analyses were performed on this combined dataset (49 haplotypes including outgroups for 2058 bp in total). The maximum-parsimony analysis resulted in 24 equally parsimonious trees, and their strict consensus tree shows that there are two major clades representing the Chinese Phrynocephalus species: the viviparous group (Clade A) and the oviparous group (Clade B). The trees derived from Bayesian, ML. and NJ analyses were topologically identical to the MP analysis except for the position of P. mystaceus. All analyses left the nodes for the oviparous group, the most basal clade within the oviparous group, and P. mystaceus unresolved. The phylogenies further suggest that the monophyly of the viviparous species may have resulted from vicariance, while recent dispersal may have been important in generating the pattern of variation among the oviparous species. (C) 2003 Elsevier Science (USA). All rights reserved.

Effects of sublethal fenitrothion ingestion on cholinesterase inhibition, standard metabolism, thermal preference, and prey-capture ability in the Australian central bearded dragon (Pogona vitticeps, Agamidae)

The central bearded dragon (Pogona vitticeps) is a medium-sized lizard that is common in semiarid habitats in Australia and that potentially is at risk of fenitrothion exposure from use of the chemical in plague locust control. We examined the effects of single sublethal doses of this organophosphate (OP; low dose = 2.0 mg/kg; high dose = 20 mg/kg; control = vehicle alone) on lizard thermal preference, standard metabolic rate, and prey-capture ability. We also measured activities of plasma total cholinesterase (ChE) and acetylcholinesterase before and at 0, 2, 8, 24, 120, and 504 h after OP dosing. Predose plasma total ChE activity differed significantly between sexes and averaged 0.66 ± 0.06 and 0.45 ± 0.06 μmol/min/ml for males and females, respectively. Approximately 75% of total ChE activity was attributable to butyrylcholinesterase. Peak ChE inhibition reached 19% 2 h after OP ingestion in the low-dose group, and 68% 8 h after ingestion in high-dose animals. Neither OP doses significantly affected diurnal body temperature, standard metabolic rate, or feeding rate. Plasma total ChE levels remained substantially depressed up to 21 d after dosing in the high-dose group, making this species a useful long-term biomonitor of OP exposure in its habitat.

Complete mitochondrial genome of the frillneck lizard (Chlamydosaurus kingii, Reptilia; Agamidae), another squamate with two control regions.

Using PCR, the complete mitochondrial genome was sequenced in three frillneck lizards (Chlamydosaurus kingii). The mitochondria spanned over 16,761bp. As in other vertebrates, two rRNA genes, 22 tRNA genes and 13 protein coding genes were identified. However, similar to some other squamate reptiles, two control regions (CRI and CRII) were identified, spanning 801 and 812 bp, respectively. Our results were compared with another Australian member of the family Agamidae, the bearded dragon (Pogana vitticeps). The overall base composition of the light-strand sequence largely mirrored that observed in P vitticeps. Furthermore, similar to P. vitticeps, we observed an insertion 801 bp long between the ND5 and ND6 genes. However, in contrast to P vitticeps we did not observe a conserved sequence block III region. Based on a comparison among the three frillneck lizards, we also present data on the proportion of variable sites within the major mitochondrial regions.

Chorological analysis and morphological variations of Saurians of the genus uromastyx (Reptilia, Agamidae) in western Sahara: Description of two new taxa.

[ES]Se describe una nueva especie del género Uromastyx sobre la base de dos especímenes procedentes del Adrar Sutuf (Sahara Occidental). Los individuos del nuevo taxón difieren fuertemente de Uromastyx acanthinura por el gran tamaño que llegan a alcanzar los adultos, su siempre mayor número de escamas, la disposición diferente de los tubérculos en los muslos, y por su diseño y coloración. Sus características generales se asemejan a las de Uromastyx aegyptia, un hecho que sugiere la posible existencia de lagartos del complejo U. aegyptia en poblaciones relícticas a lo largo del desierto del Sahara.

Isolation and characterisation of novel microsatellite and mitochondrial DNA markers for the Eastern Water Dragon (Physignathus lesueurii)

Habitat fragmentation as a result of urbanisation is a growing problem for native lizard species. The Eastern Water Dragon (Physignathus lesueurii) is a social arboreal agamid lizard, native to Australia. This species represents an ideal model species to investigate the effect of urbanisation because of their prominent abundance in the urban landscape. Here we describe the isolation and characterisation of a novel set of 74 di-, tri-, and tetramicrosatellites from which 18 were selected and optimised into two multiplexes. The 18 microsatellites generated a total 148 alleles across the two populations. The number of alleles per locus varied from 2 to 18 alleles and measures of Ho and He varied from 0.395 to 0.877 and from 0.441 to 0.880, respectively. We also present primers for four novel mitochondrial DNA (mtDNA) markers. The combined length of the four mtDNA marker pairs was 2,528 bp which included 15 nucleotides changes. In comparison to threatened species, which are generally characterised by small population sizes, the Eastern Water Dragon represents an ideal model species to investigate the effect of urbanisation on their behavioural ecology and connectivity patterns among populations.

沙蜥属部分物种分类性状评价

沙蜥属Phrynocephalus Kaup，1825隶属于爬行纲（Reptilia）有鳞目（Squamata）蜥蜴亚目（Lacertilia）鬣蜥科（Agamidae），是欧亚大陆荒漠和稀疏草原常见蜥蜴。沙蜥属的分类及系统演化关系、地理分布格局与新生代第三纪以来古地中海的变迁、青藏高原的抬升及亚洲内陆干旱荒漠化的过程有密切的关系，长期以来有关沙蜥属的研究一直受到中外学者们的关注。由于沙蜥属地理分布广、形态变异大、体色和斑纹变化复杂，虽然前人使用过许多形态性状来描述和分类沙蜥属物种，但是仍然存在许多问题。性状的分类学意义不明确是造成这些问题的主要原因之一，因此本研究针对沙蜥属常用的鉴别性状进行分类意义的分析，希望能对沙蜥属物种鉴定及分类学其它研究有所裨益。 中国沙蜥属物种主要分布于西北的干旱荒漠区域及青藏高原的大部分地区，大约为18种。 本文研究了中国境内12种沙蜥：青海沙蜥（Phrynocephalus vlangalii）、西藏沙蜥（P. theobaldi）、南疆沙蜥（P. forsythii）、变色沙蜥（P. versicolor）、旱地沙蜥（P. helioscopus）、荒漠沙蜥（P. przewalskii）、乌拉尔沙蜥（P. guttatus）、草原沙蜥（P. frontalis）、叶城沙蜥（P. axillaris）、白稍沙蜥（P. koslowi）、无斑沙蜥（P. immaculatus）和白条沙蜥（P. albolineatus），对它们的65项外部形态性状进行了观察和测量，其中数量性状29项、质量性状36项。评价了这些性状的序级性、间断性和代表性，结论如下： 1. 对于数量性状，得出了适合各级分类的数值区间； 2. 给出了在不同序级上适合分类的质量性状。 并利用各性状评价的结果，给出12种沙蜥的检索表，以及对中国沙蜥物种某些尚存在争议的问题进行了探讨。 详细记录了青海沙蜥红原亚种的骨骼系统，首次发现并命名了肘骨（elbow bone）和垫骨（stepping bone），为沙蜥属系统学研究补充了骨骼方面的证据；解剖了乌拉尔沙蜥、旱地沙蜥、荒漠沙蜥的雌体和雄体的骨骼系统，并在14项骨骼形态性状上对这3种沙蜥进行了比较。 Phrynocephalus （Squamata，Agamidae） is a familiar genus of lizards inhabited desert and sparse steppes in Eurasia. The taxonomics, phylogenetics and distribution pattern of Phrynocephalus are relative intensely to these events: the vicissitudes of the archaic Mediterranean sea since the Cainozoic, the uplift of Qingzang Plateau and the expending arid areas in the inland of Asia. Owing to the wide distribution, the large variability of the morphology and the different colors in Phrynocephalus, it is difficult to identify them. Tough many morphological characters are used to describe and discriminate them，a lot of questions still exist. One of the most important reasons is the confusion in the morphological characters. In this study, we demonstrate the validity and the invalidity of the familiar characters. There are about 18 species of the genus Phrynocephalus in China, which exist in arid desert in Northwest China and Qingzang Plateau. Twelve Chinese species was analyzed in this paper. They are P. vlangalii，P. theobaldi，P. forsythia，P. versicolor，P. helioscopus，P. przewalskii，P. guttatus，P. frontalis，P. axillaris，P. koslowi，P. immaculatus，P. albolineatus. We measure 29 quantitative characters and observe 36 qualitative characters in each individual. Through analyzing these characters, we made some conclusions as follows: 1. to every quantitative character, we get a clear numeric area to discriminate the different operational taxonomic units. 2. we chose the valid qualitative characters in these operational taxonomic units. This paper is the first to describe the “elbow bone”, which is a bone in pectoral appendage equivalent to patella，and “stepping bone”, which is a bone under carpal. A detailed description of the skeletal system of female Phrynocephalus vlangalii hongyuanensis was conducted. We also anatomise the skeletal systems of three species: P. guttatus，P. przewalskii，P. helioscopus, and compare or contrast 14 skeletal characters in them. What’s more，this paper offers some suggestions to the questions of Chinese Phrynocephalus species and keys to 12 species of Phrynocephalus basing on our conclusions on the evaluation of the morphological characters.

Sexual selection and the evolution of complex color patterns in dragon lizards

Many species have elaborate and complex coloration and patterning, which often differ between the sexes. Sexual selection may increase the size or intensity of color patches (elaboration) in one sex or drive the evolution of novel signal elements (innovation). The latter potentially increases color pattern complexity. Color pattern complexity may also be influenced by ecological factors related to predation and environment; however, very few studies have investigated the effects of both sexual and natural selection on color pattern complexity across species. We used a phylogenetic comparative approach to examine these effects in 85 species and subspecies of Australian dragon lizards (family Agamidae). We quantified color pattern complexity by adapting the Shannon–Wiener diversity index. There were clear sex differences in color pattern complexity, which were positively correlated with both sexual dichromatism and sexual size dimorphism, consistent with the idea that sexual selection plays a significant role in the evolution of color pattern complexity. By contrast, we found little evidence of a link between environmental factors and color pattern complexity on body regions exposed to predators. Our results suggest that sexual selection rather than natural selection has led to increased color pattern complexity in males.

Morphology and histochemistry of the hyolingual apparatus in chameleons

We reexamined the morphological and functional properties of the hyoid, the tongue pad, and hyolingual musculature in chameleons. Dissections and histological sections indicated the presence of five distinctly individualized pairs of intrinsic tongue muscles. An analysis of the histochemical properties of the system revealed only two fiber types in the hyolingual muscles: fast glycolytic and fast oxidative glycolytic fibers. In accordance with this observation, motor-endplate staining showed that all endplates are of the en-plaque type. All muscles show relatively short fibers and large numbers of motor endplates, indicating a large potential for fine muscular control. The connective tissue sheet surrounding the entoglossal process contains elastin fibers at its periphery, allowing for elastic recoil of the hyolingual system after prey capture. The connective tissue sheets surrounding the m. accelerator and m. hyoglossus were examined under polarized light. The collagen fibers in the accelerator epimysium are configured in a crossed helical array that will facilitate limited muscle elongation. The microstructure of the tongue pad as revealed by SEM showed decreased adhesive properties, indicating a change in the prey prehension mechanics in chameleons compared to agamid or iguanid lizards. These findings provide the basis for further experimental analysis of the hyolingual system. © 2001 Wiley-Liss, Inc.

Experience overrides colour in lizard contests

We examined the role of conspicuous coloration in male-male contests for two species of Australian dragon lizards, Ctenophortis decresii and C. vadnappa, in which conspicuous coloration has a demonstrated predation cost. We conducted contests in which the overall conspicuousness of male coloration was manipulated using paints that matched the spectral reflectance of the lizards, as well as natural (control) contests. There was little evidence for an influence of colour on contest outcome or aggression levels for either species when all experiments were considered. However, we found a significant effect of trial order and experience on contest outcome and aggression levels (the same pair of males was used for both types of contest), despite a 2-3 week interval between contests. When we examined only the first trial between unfamiliar males, we found that male C. vadnappa that had been painted to appear more conspicuous consistently won. Comparison with the natural trials suggests that the aspect of colour manipulation that was responsible for this result was the 'hue' of the throat: males with yellower throats consistently beat males with bluer throats in both natural and painted trials. The difference in coloration of flank markings also predicted the difference in aggression scores between contestants in the natural trials. These results suggest that although colour is important in opponent assessment and in determining contest outcome in C vadnappa, previous agonistic experience can override the effects of colour and have a long-lasting influence on aggressive behaviour.

Life history influences how fire affects genetic diversity in two lizard species

'Fire mosaics' are often maintained in landscapes to promote successional diversity in vegetation with little understanding of how this will affect ecological processes in animal populations such as dispersal, social organization and re-establishment. To investigate these processes, we conducted a replicated, spatiotemporal landscape genetics study of two Australian woodland lizard species [Amphibolurus norrisi (Agamidae) and Ctenotus atlas (Scincidae)]. Agamids have a more complex social and territory structure than skinks, so fire might have a greater impact on their population structure and thus genetic diversity. Genetic diversity increased with time since fire in C. atlas and decreased with time since fire in A. norrisi. For C. atlas, this might reflect its increasing population size after fire, but we could not detect increased gene flow that would reduce the loss of genetic diversity through genetic drift. Using landscape resistance analyses, we found no evidence that postfire habitat succession or topography affected gene flow in either species and we were unable to distinguish between survival and immigration as modes of postfire re-establishment. In A. norrisi, we detected female-biased dispersal, likely reflecting its territorial social structure and polygynous mating system. The increased genetic diversity in A. norrisi in recently burnt habitat might reflect a temporary disruption of its territoriality and increased male dispersal, a hypothesis that was supported with a simulation experiment. Our results suggest that the effects of disturbance on genetic diversity will be stronger for species with territorial social organization.

Spectral sensitivity of cone photoreceptors and opsin expression in two colour-divergent lineages of the lizard Ctenophorus decresii

Intraspecific differences in sensory perception are rarely reported but may occur when a species range extends across varying sensory environments, or there is coevolution between the sensory system and a varying signal. Examples in colour vision and colour signals are rare in terrestrial systems. The tawny dragon lizard Ctenophorus decresii is a promising candidate for such intraspecific variation, because the species comprises two geographically and genetically distinct lineages in which throat colour (a social signal used in intra- and inter-specific interactions) is locally adapted to the habitat and differs between lineages. Male lizards from the southern lineage have UV-blue throats, whereas males from the northern lineage are polymorphic with four discrete throat colours that all show minimal UV reflectance. Here, we determine the cone photoreceptor spectral sensitivities and opsin expression of the two lineages, to test whether they differ, particularly in the UV wavelengths. Using microspectrophotometry on retinal cone photoreceptors, we identified a long-wavelength-sensitive (LWS) visual pigment, a 'short' and 'long' medium-wavelength-sensitive (MWS) pigment and a short-wavelength-sensitive (SWS) pigment, all of which did not differ in λmax between lineages. Through transcriptome analysis of opsin genes we found that both lineages express four cone opsin genes, including the SWS1 opsin with peak sensitivity in the UV range, and that amino acid sequences did not differ between lineages with the exception of a single leucine to valine substitution in the RH2 opsin. Counts of yellow and transparent oil droplets associated with LWS+MWS and SWS+UVS cones, respectively, showed no difference in relative cone proportions between lineages. Therefore, contrary to predictions, we find no evidence of differences between lineages in single cone photoreceptor spectral sensitivity or opsin expression. However, we confirm the presence of four single cone classes, suggesting tetrachromacy in C. decresii, and we also provide the first evidence of UV sensitivity in agamid lizards.

Concordance in phylogeography and ecological niche modelling identify dispersal corridors for reptiles in arid Australia

Aim: Using the rock-specialist agamid Ctenophorus caudicinctus as a model, we test hypothesized biogeographical dispersal corridors for lizards in the Australian arid zone (across the western sand deserts), and assess how these dispersal routes have shaped phylogeographical structuring. Location: Arid and semi-arid Australia. Methods: We sequenced a c. 1400 bp fragment of mtDNA (ND2) for 134 individuals of C. caudicinctus as well as a subset of each of the mtDNA clades for five nuclear loci (BDNF, BACH1, GAPD, NTF3, and PRLR). We used phylogenetic methods to assess biogeographical patterns within C. caudicinctus, including relaxed molecular clock analyses to estimate divergence times. Ecological niche modelling (Maxent) was employed to estimate the current distribution of suitable climatic envelopes for each lineage. Results: Phylogenetic analyses identified two deeply divergent mtDNA clades within C. caudicinctus - an eastern and western clade - separated by the Western Australian sand deserts. However, divergences pre-date the Pleistocene sand deserts. Phylogenetic analyses of the nuclear DNA data sets generally support major mtDNA clades, suggesting past connections between the western C. c. caudicinctus populations in far eastern Pilbara (EP) and the lineages to the east of the sand deserts. Ecological niche modelling supports the continued suitability of climatic conditions between the Central Ranges and the far EP for C. c. graafi. Main conclusions: Estimates of lineage ages provide evidence of divergence between eastern and western clades during the Miocene with subsequent secondary contact during the Pliocene. Our results suggest that this secondary contact occurred via dispersal between the Central Ranges and the far EP, rather than the more southerly Giles Corridor. These events precede the origins of the western sand deserts and divergence patterns instead appear associated with Miocene and Pliocene climate change.