Partitioned Bayesian analyses, dispersal-vicariance analysis, and the biogeography of Chinese toad-headed lizards (Agamidae: Phrynocephalus): a re-evaluation

The toad-headed lizards of genus Phrynocephalus are distributed from northwestern China to Turkey and are one of the major components of the central Asian desert fauna. To date, published morphological and molecular phylogenetic hypotheses of Phrynocephalus are only partially congruent, and the relationships within the genus are still far from clear. We re-analyzed published mitochondrial gene sequence data (12S, 16S, cyt b, ND4-tRNA(Leu)) by employing partition-specific modeling in a combined DNA analysis to clarify existing gaps in the phylogeny of Chinese Phrynocephalus. Using this phylogenetic framework, we inferred the genus' historical biogeography by using weighted ancestral-area analysis and dispersal-vicariance analysis in combination with a Bayesian relaxed molecular-clock approach and paleogeographical data. The partitioned Bayesian analyses support the monophyly of Phrynocephalus and its sister-group relationship with Laudakia. An earlier finding demonstrating the monophyly of the viviparous group is corroborated. However, our hypothesis of internal relationships of the oviparous group differs from a previous hypothesis as our results do not support monophyly of the oviparous taxa. Instead, the viviparous taxa form a clade with many oviparous taxa exclusive of P. helioscopus and P. mystaceus. Our results also suggest that: (1) P. putjatia is a valid species, comprising populations from Guide, Qinghai Province and Tianzhu, Gansu Province; (2) P. hongyuanensis is not a valid species, synonymized instead with P. vlangalii; (3) P. zetangensis is not a valid species and should be included in P. theobaldi; (4) the population occurring in Kuytun, Xinjiang Uygur Autonomous Region is recognized as P. guttatus instead of P. versicolor; and (5) the Lanzhou population of P. frontalis is part of P. przewalskii. Congruent with previous hypotheses, the uplift of the Tibetan Plateau played a fundamental role in the diversification of Phrynocephalus. An evolutionary scenario combining aspects of vicariance and dispersal is necessary to explain the distribution of Phrynocephalus. Bayesian divergence-time estimation suggests that Phrynocephalus originated at the Middle-Late Miocene boundary (15.16-10.4 Ma), and diversified from Late Miocene to Pleistocene from a center of origin in Central Asia, Tarim Basin, and Junggar Basin temperate desert, followed by several rapid speciation events in a relatively short time. The proposed biogeographic scenarios also indicate that the Tarim Basin desert may be the secondary diversification center, followed by Junggar Basin temperate desert and Alashan Plateau temperate desert. In the viviparous group, the allopatric speciation of P. theobaldi and P. vlangalii may have been caused by the uplifting of Tanggula Mountain Ranges. In addition, the results of this study make an important contribution to understanding the uplift of the Tibetan Plateau and Tian Shan Mountains and the biogeography of the entire region.     

Carpus and tarsus patterns in some Arabian lizards (Reptilia: Agamidae, Lacertidae, Scincidae)

Within the families Agamidae, Lacertidae and Scincidae, the carpal and tarsal elements, the sesamoidea and the ossification pattern show considerable variability. Furthermore, carpal and tarsal structure in the lizards studied here is not unique to the genera in question and identical or very similar conditions occur in other lizards. These elements contain sufficient anatomical information to make them taxonomically valuable.     

沙蜥属Phrynocephalus生物地理学研究进展

沙蜥属Phrynocephalus作为鬣蜥科Agamidae中的古北界类群,分布区复杂的地质、地理历史对该属物种形成、分化及分布格局的形成具有重要的影响.近年来,随着生物地理学理论的不断发展,各国学者从不同角度对沙蜥属的生物地理学做了大量研究.本文主要综述了这方面的研究进展;同时基于沙蜥属的研究现状对今后的研究做出展望.     

The effects of ambient temperature and thermoregulation on locomotor performance of newborn Guide toad-headed lizards (Phrynocephalus putjatia)

The effect of thermal environments during embryonic development as a proximate source of variation in the fitness of offspring has been examined in a wide variety of taxa, and reptiles have been proved to be excellent mode systems for research in this field. Here, we describe a study revealing the effects of ambient temperature and thermoregulation on locomotor performance of newborn ovoviviparous lizards. A 2 (background temperatures set at 18 °C or 22 °C) × 2 (allowing thermoregulation for 14 h or 10 h daily) factorial design experiment was carried out to examine the effects of ambient temperature and thermoregulation on the locomotor performance of newborn Guide toad-headed lizards (Phrynocephalus putjatia; Agamidae). Gravid females were collected in May 2010 from a population in Guide, Qinghai, northwestern China, and were transported to our laboratory in Hangzhou. Ten to fifteen females were housed together in 1200 mm × 600 mm × 700 mm (length × width × height) communal cages, which were placed in AAPS (artificial atmospheric phenomena simulator) rooms, and contained a substrate of sand (～400 mm depth), with rocks and pieces of clay tiles provided as shelter and basking sites. One light bulb (200 W) was suspended above one end of the cage to create a thermal gradient ranging from room temperature to 60 °C for 14 h or 10 h daily, and overnight temperatures followed AAPS temperatures (18 °C or 22 °C). Food (mealworms and house crickets) dusted with multivitamins and minerals and water were provided daily. Cages were checked twice daily for neonates after the first female gave birth, and neonates were immediately collected and weighed after birth.Twenty neonates from single litters of each testament were measured at birth for locomotor performance. All running trials were conducted at a body temperature of 30 °C, which was achieved by placing the newborns in an incubator at 30 °C for 30 min prior to testing. Locomotor performance was assessed by chasing the neonates along a 2-m-long racetrack, which was placed in a room at constant 30 °C, with one side of the racetrack transparent, allowing videoing with a Panasonic NV-DS77 digital video camera. The tapes were later examined with a computer using MGI VideoWave III software for PC (MGI Software Co., Toronto, Canada) for sprint speed in the fastest 250-mm interval and the maximal length. Each individual was measured five times after birth, at 15-day intervals, until 60 days.We found that locomotor performance of neonates was affected by thermoregulating opportunity, but not by background temperature. Neonates produced by females thermoregulated for 14 h daily performed better (both sprint speed and the maximal length) in the racetrack than those produced by females thermoregulated for 10 h daily. However, the interaction between background temperature and thermoregulating opportunity was not a significant source of variation in locomotor performance. Moreover, sprint speed was positively correlated to the maximal length in newborn P. putjatia. In summary, locomotor performance is a highly fitness-related trait, and this study implies that viviparity allows female P. putjatia to provide optimal temperatures for embryo development through thermoregulation, thereby producing well-performed offspring.     

沙蜥属系统学研究综述

沙蜥属隶属于鬣蜥科,是欧亚大陆荒漠和稀疏草原常见的蜥蜴。该属特征变异较大,存在许多争议。本文从其分类史、系统演化关系、生物地理学研究三个方面进行总结,并指出沙蜥属卵生类群的系统关系还比较混乱,生物地理历史过程有待进一步研究。     

荒漠沙蜥食物的两性差异

对2008年4～6月在甘肃省甘塘、阿拉善左旗、民勤、山丹采集的荒漠沙蜥样本进行了食物两性差异的研究。主要采用剖胃法对219个荒漠沙蜥胃样的6021个食物做了识别和鉴定。结果表明,荒漠沙蜥的食性为动物性,食物多为昆虫,主要取食半翅目和膜翅目昆虫,尤其是长蝽科和蚁科(合计高于70%),取食猎物的体积大小集中在0.01～20mm3。4个种群的沙蜥对长蝽科昆虫均表现为正向选择,部分种群对蚁科昆虫表现负向选择。可能是因为沙蜥有食物选择性,对长蝽科昆虫取食较多,对拟步甲科摄食较少。较多地取食蚂蚁则是因为生境中喜好食物较少,而蚁科昆虫数量多,遇见率也较高。荒漠沙蜥雌雄个体食物生态位重叠度极高,但是卡方检验的结果却表明两性在食物组成和食物大小存在显著差异。由于雌雄个体对同种食物有相同的选择性,因此导致两性在食物组成和食物大小上存在显著差异的原因可能是雄性家域范围更大,对食物有更多选择。     

The cost of living large: comparative gliding performance in flying lizards (Agamidae: Draco)

Despite exhibiting considerable interspecific variation in body mass, flying lizards of the genus Draco are isometric in their area-mass scaling relationships and exhibit no significant compensatory variation in wing aspect ratio. Thus, larger species are expected to be relatively poor gliders, in lieu of behavioral or physiological compensation, when compared with smaller congeners. Here we tested this hypothesis by conducting gliding performance trials for 11 Draco species spanning virtually the entire size range of the genus. We considered three primary performance variables: maximum velocity adjusted for wind conditions, height lost over a standard horizontal glide distance, and glide angle. Comparative analysis confirmed that larger species are relatively poor gliders and do not compensate substantially for their higher wing loadings via either behavioral or physiological mechanisms. Flying lizards were found to exhibit substantial context-dependent variation in glide performance, with smaller species often exhibiting extensive variation in height lost and glide angle between trials. Variation also was observed in empirically derived velocity profiles, with only a subset of individuals appearing to perform equilibrium glides. Such size-dependent variation in performance has important consequences for the ecology and evolution of flying lizards and other glissant taxa.     

Phylogeny of North African Agama lizards (Reptilia: Agamidae) and the role of the Sahara desert in vertebrate speciation

The origin of Saharan biodiversity is poorly understood, in part because the geological and paleoclimatic events that presumably shaped species diversity are still controversial, but also because few studies have explored causal explanations for the origin of Saharan diversity using a phylogenetic framework. Here, we use mtDNA (16S and ND4 genes) and nDNA (MC1R and CMOS genes) to infer the relationships and biogeographic history of North African agamas (genus Agama). Agamas are conspicuous, diverse and abundant African lizards that also occur in the Saharan xeric and mesic environments. Our results revealed the presence of three Agama lineages in North Africa: one Afrotropical, one Sahelo-Saharan, and one broadly distributed in North Africa and mainly Saharan. Southern Mauritania contains the highest known diversity, with all three lineages present. Results suggest that agamas colonized the Sahara twice, but only one lineage was able to radiate and diversify there. Species in the Saharan lineage are mostly allopatric, and their splitting, genetic diversity and distribution are greatly explained by mountain ranges. One species in this lineage has colonized the Mediterranean climatic zone (A. impalearis), and another one the Sahel savannah (A. boueti). The other lineage to colonize the Sahara corresponds to A. boulengeri, an eminently Sahelian species that also inhabits Saharan mountain ranges in Mauritania and Mali. Phylogenetic analyses indicate that allopatric montane populations within some currently recognized species are also genetically divergent. Our study therefore concludes that vicariant speciation is a leading motor of species diversification in the area: Inside the Sahara, associated to mountain-ranges isolated by dune seas and bare plains; outside, associated to less harsh climates to the North and South. Paleoclimatic oscillations are suggested as causal explanations of the vicariant distribution and origin of species. Agamas are thought to have colonized northern Africa during wet periods, with subsequent dry periods fragmenting species distribution and leading to allopatric populations associated to milder and wetter climates in the Mediterranean, Sahel, and in Saharan mountains, in an island-model fashion. Finally, our results support the synonymization of A. castroviejoi with A. boueti, the reciprocal monophyly of all other North African agamas, and suggest one candidate species within A. boulengeri.     

奇台沙蜥生长过程中的两性异形

研究奇台沙蜥Phrynocephalus grumgrzimailoi头、尾、腋胯距大小在个体发育过程中的变化.成体体长(SVL)无显著的两性差异,两性异形主要表现为雄性个体有较大的头部(头长和头宽)和尾部,雌性具有较大的腋胯距.头、尾、腋胯距大小的两性异形在幼体就已存在,并随个体发育的进行变得更加显著.不同年龄组两性个体头部、尾部、腋胯距随SVL呈异速增长,表现为两性头部的增长速率在个体发育过程中逐渐增大,尾部的增长速率逐渐减慢,腋胯距在雌性蜥蜴中增长速率逐渐增大,在雄性中逐渐变小.     

The role of prostaglandins and the hypothalamus in thermoregulation in the lizard, Phrynocephalus przewalskii (Agamidae)

Typically, small lizards rely heavily on behavioral thermoregulation rather than physiological mechanisms to control their rates of warming and cooling. We tested the hypothesis that prostaglandins participate in mediating the cardiovascular response to heating and cooling and temperature regulating neurons in the hypothalamus of the small lizard Phrynocephalus przewalskii. In vivo and in vitro treatments, heart rates (HRs) were all found to be higher during heating than during cooling, hysteresis was distinct below 30 and 26°C, respectively. In vivo, as administration of COX inhibitor, there were no differences in HR between heating and cooling at any body temperature and administration of agonist prostaglandins only produced a significant effect on HR below 25°C. Single-unit activity was recorded extracellularly in vitro with microelectrodes, found the firing rate of the continuous unit increased 23% when the temperature of the artificial cerebrospinal fluid dropped from 30–20°C. We conclude that prostaglandins appear to play only a limited role in modulating heart activity in Phrynocephalus przewalskii and suggest that cold-sensitive neurons in the preoptic and anterior hypothalamus (PO/AH) are involved in thermoregulatory control during heating or cooling.     

Cladogenesis and phylogeography of the lizard Phrynocephalus vlangalii (Agamidae) on the Tibetan plateau

Phrynocephalus vlangalii is restricted to dry sand or Gobi desert highlands between major mountain ranges in the Qinghai (Tibetan) Plateau. Mitochondrial DNA (mtDNA) sequence (partial ND2, tRNA(Trp) and partial tRNA(Ala)) was obtained from 293 Phrynocephalus sampled from 34 sites across the plateau. Partitioned Bayesian and maximum parsimony phylogenetic analyses revealed that P. vlangalii and two other proposed species (P. erythrus and P. putjatia) together form a monophyletic mtDNA clade which, in contrast with previous studies, does not include P. theobaldi and P. zetangensis. The main P. vlangalli clade comprises seven well-supported lineages that correspond to distinct geographical areas with little or no overlap, and share a most recent common ancestor at 5.06 +/- 0.68 million years ago (mya). This is much older than intraspecific lineages in other Tibetan animal groups. Analyses of molecular variance indicated that most of the observed genetic variation occurred among populations/regions implying long-term interruption of maternal gene flow. A combined approach based on tests of population expansion, estimation of node dates, and significance tests on clade areas indicated that phylogeographical structuring has been primarily shaped by three main periods of plateau uplift during the Pliocene and Pleistocene, specifically 3.4 mya, 2.5 mya and 1.7 mya. These periods corresponded to the appearance of several mountain ranges that formed physical barriers between lineages. Populations from the Qaidam Basin are shown to have undergone major demographic and range expansions in the early Pleistocene, consistent with colonization of areas previously covered by the huge Qaidam palaeolake, which desiccated at this time. The study represents one of the most detailed phylogeographical analyses of the Qinghai Plateau to date and shows how geological events have shaped current patterns of diversity.     

The origin of parachuting locomotion in gekkonid lizards (Reptilia: Gekkonidae)

Several recent studies have focused on the ability of the gekkonid Ptychozoon lionatum to parachute, utilizing its enlarged lateral body folds. Little attention has been paid, however, to the evolution of such behavious and here several aspects pertaining to the origin of parachuting are integrated. Behavioural shifts and morphological modifications are considered together and evaluated in the light of ecological information. It is apparent that certain behavioural shifts were the precursors of subsequent morphological changes. The structural design of the body folds is also considered in relation to parachuting behaviour and both the gross morphology and histology are shown to be significantly modified to permit effective parachuting. The absene of musculature in the lateral body flaps of geckos is confirmed and a model for the integration of the various factors involved in effective parachuting is presented. Here the importance of the structure-function and habitat-behaviour sets are emphasized together with their new interrelationships when a protoadaptation becomes functional.     

Global diversity of lizards in freshwater (Reptilia: Lacertilia)

No lizards are strictly aquatic, but at least 73 species in 11 different families can be considered to regularly utilize freshwater habitats. There are no aquatic lizards in the Nearctic or Palearctic regions, whereas the Neotropics, Southeast Asia, and the Indo-Australian Archipelago support the greatest diversity of freshwater forms, particularly in the families Gymnophthalmidae, Scincidae and Varanidae. A number of larger aquatic lizards are harvested for food and for the reptile skin trade and several are CITES listed. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Diet,Food Intake of Phrynocephalus frontalis （Agamidae） and Its Potential Role in Desert Habitat

We examined the dietary diversity and food intake of Phrynocephalus frontalis, compared the difference of insect diversity in the natural habitats with different lizard densities, and discussed the potential role of this lizard in the desert ecosystem. The results show that： （1） arthropodans of the orders Coleoptera, Hymenoptera and Hemiptera were major dietary components of P. frontalis; （2） coleoptera larvae always formed the predominant component of lizard diets; （3） dietary diversities of P. frontalis were not significantly different between summer and autumn or between the two sexes; （4） the similarity in trophic niches between seasons was 0.756, whereas the similarity in trophic niches between sexes was 0.994; （5） stomach content weight of lizards varied significantly among different seasons, but there was no significant difference in stomach content weight between sexes; （6） insect diversity differed significantly among the groups of the habitat with different degrees of lizard density, and the habitat with moderate lizards density had the highest insect diversity. We infer that P. frontalis prey mainly on insects and change their diet and food intake with season; males and females consumed similar preys in types and weights. As an important predator, P. frontalis could affect the insect community in the arid ecosystem of Hunshandak Desert on the Mongolian Plateau.