Integrative analyses of speciation and divergence in Psammodromus hispanicus (Squamata: Lacertidae).

ABSTRACT: BACKGROUND: Genetic, phenotypic and ecological divergence within a lineage is the result of past and ongoing evolutionary processes, which lead ultimately to diversification and speciation. Integrative analyses allow linking diversification to geological, climatic, and ecological events, and thus disentangling the relative importance of different evolutionary drivers in generating and maintaining current species richness. RESULTS: Here, we use phylogenetic, phenotypic, geographic, and environmental data to investigate diversification in the Spanish sand racer (Psammodromus hispanicus). Phylogenetic, molecular clock dating, and phenotypic analyses show that P. hispanicus consists of three lineages. One lineage from Western Spain diverged 8.3 (2.9-14.7) Mya from the ancestor of Psammodromus hispanicus edwardsianus and P. hispanicus hispanicus Central lineage. The latter diverged 4.8 (1.5-8.7) Mya. Molecular clock dating, together with population genetic analyses, indicate that the three lineages experienced northward range expansions from southern Iberian refugia during Pleistocene glacial periods. Ecological niche modelling shows that suitable habitat of the Western lineage and P. h. edwardsianus overlap over vast areas, but that a barrier may hinder dispersal and genetic mixing of populations of both lineages. P. h. hispanicus Central lineage inhabits an ecological niche that overlaps marginally with the other two lineages. CONCLUSIONS: Our results provide evidence for divergence in allopatry and niche conservatism between the Western lineage and the ancestor of P. h. edwardsianus and P. h. hispanicus Central lineage, whereas they suggest that niche divergence is involved in the origin of the latter two lineages. Both processes were temporally separated and may be responsible for the here documented genetic and phenotypic diversity of P. hispanicus. The temporal pattern is in line with those proposed for other animal lineages. It suggests that geographic isolation and vicariance played an important role in the early diversification of the group, and that lineage diversification was further amplified through ecological divergence.

Integrative analyses of speciation and divergence in Psammodromus hispanicus (Squamata: Lacertidae).

BackgroundGenetic, phenotypic and ecological divergence within a lineage is the result of past and ongoing evolutionary processes, which lead ultimately to diversification and speciation. Integrative analyses allow linking diversification to geological, climatic, and ecological events, and thus disentangling the relative importance of different evolutionary drivers in generating and maintaining current species richness.ResultsHere, we use phylogenetic, phenotypic, geographic, and environmental data to investigate diversification in the Spanish sand racer (Psammodromus hispanicus). Phylogenetic, molecular clock dating, and phenotypic analyses show that P. hispanicus consists of three lineages. One lineage from Western Spain diverged 8.3 (2.9-14.7) Mya from the ancestor of Psammodromus hispanicus edwardsianus and P. hispanicus hispanicus Central lineage. The latter diverged 4.8 (1.5-8.7) Mya. Molecular clock dating, together with population genetic analyses, indicate that the three lineages experienced northward range expansions from southern Iberian refugia during Pleistocene glacial periods. Ecological niche modelling shows that suitable habitat of the Western lineage and P. h. edwardsianus overlap over vast areas, but that a barrier may hinder dispersal and genetic mixing of populations of both lineages. P. h. hispanicus Central lineage inhabits an ecological niche that overlaps marginally with the other two lineages.ConclusionsOur results provide evidence for divergence in allopatry and niche conservatism between the Western lineage and the ancestor of P. h. edwardsianus and P. h. hispanicus Central lineage, whereas they suggest that niche divergence is involved in the origin of the latter two lineages. Both processes were temporally separated and may be responsible for the here documented genetic and phenotypic diversity of P. hispanicus. The temporal pattern is in line with those proposed for other animal lineages. It suggests that geographic isolation and vicariance played an important role in the early diversification of the group, and that lineage diversification was further amplified through ecological divergence.

A New Species of Sand Racer, Psammodromus (Squamata: Lacertidae), from the Western Iberian Peninsula

A new species of lacertid lizard of the genus Psammodromus is described from the Iberian Peninsula. Genetic and recently published phenotypic data support the differentiation of Psammodromus hispanicus into three, and not as previously suggested two, distinct lineages. Age estimates, lineage allopatry, the lack of mitochondrial and nuclear haplotype sharing between lineages, ecological niche divergence, and the current biogeographic distribution, indicated that the three lineages correspond to three independent species. Here, we describe a new species, Psammodromus occidentalis sp. n., which is genetically different from the other sand racers and differentiated by the number of femoral pores, number of throat scales, snout shape, head ratio, green nuptial coloration, and number of supralabial scales below the subocular scale. We also propose to upgrade the two previously recognized subspecies, Psammodromus hispanicus hispanicus Fitzinger, 1826 from central Spain and Psammodromus hispanicus edwardsianus (Dugès, 1829) from eastern Spain, to the species level: Psammodromus hispanicus stat. nov. and Psammodromus edwardsianus stat. nov. Given that the holotype of Psammodromus hispanicus was lost, we designate a neotype. We also analysed museum specimens of P. blanci, P. microdactylus and P. algirus to describe differentiation of the Psammodromus hispanicus lineages/species from their closest relatives.

Osteologia do neurocrânio de Iguana iguana iguana (Squamata: Iguanidae)

Resumo: O crânio representa o segmento com conspícuas adaptações que, nos lagartos, podem ser conservativas ou impulsionadas por pressões seletivas. Objetivando subsidiar o conhecimento morfológico dos répteis, fornecemos uma descrição detalhada dos ossos que formam o neurocrânio de Iguana iguana iguana com base na análise de três esqueletos secos de espécimes adultos. O crânio da referia espécie possui características basais entre os lagartos sem o fechamento das aberturas cranianas e formato geral triangular. As estruturas ósseas que formam a base craniana apresentam muitas fusões, principalmente no assoalho. Na face caudal o exoccipital e o opistótico estão fundidos e formam o otoccipital, que contribui para a formação dos terços laterais do côndilo occipital. A parte central do côndilo é formada pelo supraoccipital. Fusões e estruturas esqueléticas presentes em Iguana são similares aos demais lagartos. Não foram descritas autapomorfias no neurocrânio para esta espécie.

Activation of a P2Y4-like purinoceptor triggers an increase in cytosolic [Ca2+] in the red blood cells of the lizard Ameiva ameiva (Squamata, Teiidae)

An increasing number of pathophysiological roles for purinoceptors are emerging, some of which have therapeutic potential. Erythrocytes are an important source of purines, which can be released under physiological and physiopathological conditions, acting on purinergic receptors associated with the same cell or with neighboring cells. Few studies have been conducted on lizards, and have been limited to ATP agonist itself. We have previously shown that the red blood cells (RBCs) of the lizard Ameiva ameiva store Ca2+ in the endoplasmic reticulum (ER) and that the purinergic agonist ATP triggers a rapid and transient increase of [Ca2+]c by mobilization of the cation from internal stores. We also reported the ability of the second messenger IP3 to discharge the ER calcium pool of the ER. Here we characterize the purinoceptor present in the cytoplasmic membrane of the RBCs of the lizard Ameiva ameiva by the selective use of ATP analogues and pyrimidine nucleotides. The nucleotides UTP, UDP, GTP, and ATPgammaS triggered a dose-dependent response, while interestingly 2MeSATP, 2ClATP, alpha, ß-ATP, and ADP failed to do so in a 1- to 200-µm con- centration. The EC50 obtained for the compounds tested was 41.77 µM for UTP, 48.11 µM for GTP, 53.11 µM for UDP, and 30.78 µM for ATPgammaS. The present data indicate that the receptor within the RBCs of Ameiva ameiva is a P2Y4-like receptor due to its pharmacological similarity to the mammalian P2Y4 receptor.

Phylogeny, species limits, and biogeography of the Brazilian lizards of the genus Eurolophosaurus (Squamata : Tropiduridae) as inferred from mitochondrial DNA sequences

Phylogenetic relationships and divergence times for 10 populations of the three recognized ""species"" of Brazilian lizards of genus Eurolophosaurus were estimated from 1229 bp of cyt b, COI, 12S, and 16S rRNA mitochondrial gene segments. Eurolophosaurus is monophyletic and the basal split within the genus separates E divaricatus from a clade comprising E amathites and E nanuzae. Three populations of E divaricatus, which occurs along the western bank of Rio S (a) over tildeo Francisco, were consistently grouped together. Oil the east bank of the river, E amathites and E nanuzae from state of Bahia were recovered as the sister group of E nanuzae populations from state of Minas Gerais. The paraphyly of E nanuzae and the high divergence levels among populations of E divaricatus strongly suggest that species limits in Eurolophosaurus should be revised. Even considering an extreme evolutionary rate of 2.8% sequence divergence per million years for the four gene segments analyzed together, E. divaricatus would have separated from the two other species by at least 5.5 my ago, and E. amathites from E nanuzae populations from Bahia and Minas Gerais, respectively, by 1.5 and 3.5 my. The paleolacustrine hypothesis and changes in the course of the river potentially explain faunal divergence in the area, but divergences are much older than previously admitted. (C) 2007 Elsevier Inc. All rights reserved.

Genetic structure, phylogeny, and biogeography of Brazilian eyelid-less lizards of genera Calyptommatus and Nothobachia (Squamata, Gymnophthalmidae) as inferred from mitochondrial DNA sequences

Calyptommatus and Nothobachia genera of gymnophthalmid lizards are restricted to sandy open habitats on Sao Francisco River margins, northeastern Brazil. Phylogenetic relationships and geographic distribution of the four recognized species of Calyptommatus were analyzed from partial mitochondrial cyt b, 12S, and 16S rRNA genes sequencing, taking allopatric populations of the monotypic Nothobachia ablephara as the outgroup. In Calyptommatus a basal split separated C. sinebrachiatus, a species restricted to the eastern bank of the river, from the three other species. In this clade, C. confusionibus, found on western margin, was recovered as the sister group of the two other species, C. leiolepis and C. nicterus, from opposite margins. According to approximate date estimations, C. sinebrachiatus would have separated from the other congeneric species by 4.4-6.5 my, and C. nicterus, also from eastern bank, would be diverging by 1.8-2.6 my from C. leiolepis, the sister species on the opposite margin. C. confusionibus and C. leiolepis, both from western sandy areas, would be differentiating by 2.8-5.0 my. Divergence times of about 3.0-4.0 my were estimated for allopatric populations of Nothobachia restricted to western margin. Significant differences in 16S rRNA secondary structure relatively to other vertebrates are reported. Distinct evolutionary patterns are proposed for different taxa in those sandy areas, probably related to historical changes in the course of Sao Francisco River. (C) 2010 Elsevier Inc. All rights reserved.

Two new species of Cnemidophorus (Squamata: Teiidae) of the C. ocellifer group, from Bahia, Brazil

Two new species of Cnemidophorus are described from the right bank of the Sao Francisco river, in the northwestern part of state of Bahia, Brazil. Both species are assigned to the Cnemidophorus ocellifer group and are distinguished from all other congeners on the basis of lepidosis and color pattern. One of them, Cnemidophorus cyanurus, shares with the species of the subgroup of C. littoralis (C. abaetensis, C. littoralis and C. venetacaudus), a bluish green tail, spurs on the heels of males, 6-7 supraciliaries, a high number of femoral pores (27-45), a row of enlarged scales in the dorsal part of the humerus, and 8 to 10 rows of ventral scales. The second species, Cnemidophorus nigrigula, shares with the C. ocellifer subgroup (composed of C. ocellifer, C. mumbuca, C. jalapensis and C. confusionibus) a low number of femoral pores (1421), enlarged scales in the temporal region posterior to the third subocular, 5 supraciliaries, 6 to 8 rows of ventral scales, and a brown tail color. It is also characterized by males being conspicuously larger than females and by females retaining the juvenile color pattern, which is lost in adult males. The latter characteristic has not been reported in any species of the C. ocellifer group before now. The two new species occur sympatrically at Santo Inacio.

Comparative Cranial Osteology of Fossorial Lizards From the Tribe Gymnophthalmini (Squamata, Gymnophthalmidae)

Squamates (lizards, snakes and amphisbaenians) are represented by a large number of species distributed among a wide variety of habitats. Changes in body plan related to a fossorial habit are a frequent trend within the group and many morphological adaptations to this particular lifestyle evolved convergently in nonrelated species, reflecting adaptations to a similar habitat. The fossorial lifestyle requires an optimal morphological organization for an effective use of the available resources. Skeleton arrangement in fossorial squamates reflects adaptations to the burrowing activity, and different degrees of fossoriality can be inferred through an analysis of skull morphology. Here, we provide a detailed description of the skull morphology of three fossorial gymnophthalmid species: Calyptommatus nicterus, Scriptosaura catimbau, and Nothobachia ablephara. J. Morphol. 271: 1352-1365, 2010. (C) 2010 Wiley-Liss, Inc.

New Species of Earless Lizard Genus Heterodactylus (Squamata: Gymnophthalmidae) from the Highlands of Chapada Diamantina, State of Bahia, Brazil

A new species of Heterodactylus is described based on two specimens obtained in the highlands of Chapada Diamantina, state of Bahia, Brazil. The new lizard is characterized by very elongate body and tail, absence of external ear opening, presence of moveable eyelids, absence of prefrontals and frontoparietals, a vestigial interparietal, 37-39 dorsal and 27-29 ventral transverse scale rows, 23-25 scales around midbody, six gular scale rows, and 10-11 and 14-15 fourth finger and fourth toe infradigital lamellae, respectively. The new species is most similar to Heterodactylus lundii from which it differs by the absence of contact between frontal and interparietal, by having wider than long parietals, smooth posterior dorsal scales, posterior ventral scales almost twice longer than wide, a lower number of scales around midbody, last supralabial in contact with the granules of the ear depression, and a more elongate body. The new species occurs about 1,100 km north of the northernmost known record of H. lundii. Species of Heterodactylus seem to be restricted to areas of cold climates associated with high latitudes and mountainous areas of eastern Brasil.

A new Amphisbaena with chevron-shaped anterior body annuli from state of Pernambuco: Brazil (Squamata: Amphisbaenidae)

A new species of Amphisbaena is described from Fazenda Porto Seguro, municipality of Buique, state of Pernambuco, in the Caatingas of northeastern Brazil based on four specimens. The new species is a small and slender amphisbaenian with four precloacal pores, 333-337 body annuli, 22-23 tail annuli with discrete evidence of an autotomic site on the 10-12 tail annuli, and 14 dorsal and 17-18 ventral segments per annuli at midbody. The high number of body annuli, the presence of chevron-shaped anterior body annuli, and the fusion of frontal scales distinguish Amphisbaena supernumeraria sp. nov. from its congeners.

A new species of lizard genus Gymnodactylus Spix, 1825 (Squamata: Gekkota: Phyllodactylidae) from Serra do Sincora, northeastern Brazil, and the status of G. carvalhoi Vanzolini, 2005

A new species of gekkonid lizard genus Gymnodactylus Spix, 1825 is described on the basis of material collected in the ""campos rupestres"" (open rocky communities) of Mucuge municipality, northeastern Brazil. G. vanzolinii sp. nov. differs from its congeners in the number of transverse and longitudinal rows of dorsal tubercles and color pattern. It is thought to be closer to G. guttulatus which also occurs and is restricted to the rocky communities from highland open areas in the Espinhaco mountain range. The discovery leads to a reevaluation of the taxonomic status of G. carvalhoi Vanzolini, 2005, which is here considered a synonym of G. amarali Barbour, 1925.

New species of Amphisbaena with a nonautotomic and dorsally tuberculate blunt tail from state of Bahia, Brazil (Squamata, Amphisbaenidae)

A new species of Amphisbaena is described from Fazenda Caraibas, municipality of Mucuge, state of Bahia, Brazil, in the northern portion of the Serra do Espinhaco. The new species is a small amphisbaenian without precloacal pores, 210-213 body annuli, 12-13 tail annuli without evident autotomic site, and 14 dorsal and 14-15 ventral segments per annuli at midbody. The striking difference of this form is the presence of small tubercles on the dorsal region of its tail. This feature is unique among its congeners, although Rhineura floridana, a North American amphisbaenian, has tubercles on its tail. We suggest that the presence of tubercles on the tail of Amphisbaena sp. nov. and Rhineura floridana has arisen independently.

Two new species of Cnemidophorus (Squamata: Teiidae) from the Caatinga, Northwest Brazil

Two syntopic species of Cnemidophorus are described from the Caatingas of the Parque Nacional da Serra das Confusoes (PNSC), located in the Southwestern region of the state of Piaui in Brazil. Both species are assigned to the ocellifer group, differing from all other members of the group by their distinct color pattern and lepidosis. Besides these differences, both new species share a number of particular features with other members of the group. One of them, C. venetacaudus, shares with C. abaetensis and C. littoralis the presence of spurs in the heels of males, six supraciliar scales, a high number of femoral pores (from 21-45), a row of enlarged scales in the dorsal region of the arm, 8-10 rows of ventral scales, and a bright bluish-green tail, while the other species, C. confusionibus, shares with C. ocellifer, C. mumbuca, and C. jalapensis a low number of femoral pores, enlarged scales in the temporal region (posterior to third subocular), 5 supraciliar scales, and 6-8 rows of ventral scales. Based on these comparisons, we suggest that the ocellifer group is more complex than previously admitted, being composed by at least two morphologically recognizable species subgroups.

New Species of Anops Bell, 1833 (Squamata, Amphisbaenia) from Jalapao Region in the Brazilian Cerrado

A new species of keel-headed amphisbaenian of the genus Anops is described from the Cerrado of the Jalapao region, Tocantins state, Brazil. This new species of Anops is described from a single specimen, which may be easily distinguished from the other species of the genus, Anops bilabialatus and Anops kingii, by showing an extremely narrow head (37.2% head length); a row of eight occipitals anterior to the first body annulus; temporal present, mental and postmental fused; four postgenial rows located between the malars; and two malars posterior to the second infralabial. The new species is the first of the genus found in the Cerrado core area, and, based on the available records, the single species in the genus may be restricted to this region.