Life-History Evolution on Tropidurinae Lizards: Influence of Lineage, Body Size and Climate

The study of life history variation is central to the evolutionary theory. In many ectothermic lineages, including lizards, life history traits are plastic and relate to several sources of variation including body size, which is both a factor and a life history trait likely to modulate reproductive parameters. Larger species within a lineage, for example tend to be more fecund and have larger clutch size, but clutch size may also be influenced by climate, independently of body size. Thus, the study of climatic effects on lizard fecundity is mandatory on the current scenario of global climatic change. We asked how body and clutch size have responded to climate through time in a group of tropical lizards, the Tropidurinae, and how these two variables relate to each other. We used both traditional and phylogenetic comparative methods. Body and clutch size are variable within Tropidurinae, and both traits are influenced by phylogenetic position. Across the lineage, species which evolved larger size produce more eggs and neither trait is influenced by temperature components. A climatic component of precipitation, however, relates to larger female body size, and therefore seems to exert an indirect relationship on clutch size. This effect of precipitation on body size is likely a correlate of primary production. A decrease in fecundity is expected for Tropidurinae species on continental landmasses, which are predicted to undergo a decrease in summer rainfall.

Fight versus flight: the interaction of temperature and body size determines antipredator behaviour in tegu lizards

Ectotherm antipredator behaviour might be strongly affected both by body temperature and size: when environmental temperatures do not favour maximal locomotor performance, large individuals may confront predators, whereas small animals may flee, simply because they have no other option. However, integration of body size and temperature effects is rarely approached in the study of antipredator behaviour in vertebrate ectotherms. In the present study we investigated whether temperature affects antipredator responses of tegu lizards, Tupinambis merianae, with distinct body sizes, testing the hypothesis that small tegus (juveniles) run away from predators regardless of the environmental temperature, because defensive aggression may not be an effective predator deterrent, whereas adults, which are larger, use aggressive defence at low temperatures, when running performance might be suboptimal. We recorded responses of juvenile (small) and adult (large) tegu lizards to a simulated predatory attack at five environmental temperatures in the laboratory. Most differences between the two size classes were observed at low temperatures: large tegus were more aggressive overall than were small tegus at all temperatures tested, but at lower temperatures, the small lizards often used escape responses whereas the large ones either adopted a defensive posture or remained inactive. These results provide strong evidence that body size and temperature affect the antipredator responses of vertebrate ectotherms. We discuss the complex and intricate network of evolutionary and ecological parameters that are likely to be involved in the evolution of such interactions. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Are corridors, fragment size and forest structure important for the conservation of leaf-litter lizards in a fragmented landscape?

To investigate the implications of forest fragmentation for conservation of leaf-litter lizards the importance of fragment size, corridors and forest structure was examined in 20 forest fragments and six localities within a continuous forest in the Atlantic Plateau of Sao Paulo state, Brazil. The fragments were 2-276 ha in area and had different degrees of connectivity depending on the presence or absence of corridors. Two species of lizards were dominant, Ecpleopus gaudichaudii and Enyalius perditus. Variation in forest structure among sites was important only in explaining the abundance of E. perditus. Regardless of variation in forest structure, lizard species composition, total lizard abundance, number of species and abundance of E. perditus were sensitive to fragmentation per se but not to fragment size or corridor linkage. The inhospitable matrix surrounding fragments is probably what determines the presence and abundance of E. perditus and the high er lizard richness in continuous forests. These conditions may have prevented lizard species from recolonizing the forest fragments. Our results emphasize that the conservation of this leaf-litter fauna depends on the maintenance of large tracts of continuous forests and not on the size of fragments or on the presence of forest connections. Strategies for conservation of leaf-litter lizards in such highly fragmented Atlantic Forest landscapes should consider the enlargement of landscape connectivity between fragments and continuous forest, allowing the latter areas to act as a source of individuals for fragments.

Are leaf-litter frogs and lizards affected by edge effects due to forest fragmentation in Brazilian Atlantic forest?

FAPESP Fundacao de Amparo a Pesquisa do Estado de Sao Paulo

The comparative field body temperature among Liolaemus lizards: Testing the static and the labile hypotheses

Two competing hypotheses have been suggested to explain thermal sensitivity of lizards to environmental conditions. These are the static and the labile hypotheses. The static hypothesis posits that thermal physiology is evolutionary conservative and consequently relatively insensitive to directional selection. Contrarily, the labile hypothesis states that thermal physiology does respond readily to directional selection in some lizard taxa. In this paper, we tested both hypotheses among species of Liolaemus lizards. The genus Liolaemus is diverse with about 200 species, being broadly distributed from central Peru to Tierra del Fuego at the southern end of South America. Data of field body temperature (T(b)) from Liolaemus species were collected from the literature. Based on the distributional range of the species we also collected data of mean annual ambient temperatures. We observed that both the traditional analysis and the phylogenetic approach indicate that in the genus Liolaemus T(b) of species varies in a manner that is consistent with ecological gradient of ambient temperature. The data suggest that the thermal physiology of Liolaemus lizards is evolutionarily flexible, and that this plasticity has been partially responsible for the colonization of a wide array of thermal environments. (C) 2009 Elsevier Ltd. All rights reserved.

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.

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.

Locomotion patterns in two South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus

We quantified gait and stride characteristics (velocity, frequency, stride length, stance and swing duration, and duty factor) in the bursts of locomotion of two small, intermittently moving, closely related South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus. They occur in different environments: V rubricauda is widely distributed in open areas with various habitats and substrates, while P. tetradactylus is endemic to dunes in the semi-arid Brazilian Caatinga. Both use trot or walking trot characterised by a lateral sequence. For various substrates in a gradient of roughness (perspex, cardboard, sand, gravel), both species have low relative velocities in comparison with those reported for larger continuously moving lizards. To generate velocity, these animals increase stride frequency but decrease relative stride length. For these parameters, P. tetradactylus showed lower values than V rubricauda. In their relative range of velocities, no significant differences in stride length and frequency were recorded for gravel. However, the slopes of a correlation between velocity and its components were lower in P. tetradactylus on cardboard, whereas on sand this was only observed for velocity and stride length. The data showed that the difference in rhythmic parameters between both species increased with the smoothness of the substrates. Moreover, P. tetradactylus shows a highly specialised locomotor strategy involving lower stride length and frequency for generating lower velocities than in V. rubricauda. This suggests the evolution of a central motor pattern generator to control slower limb movements and to produce fewer and longer pauses in intermittent locomotion. (c) 2008 Elsevier GmbH. All rights reserved.

Differential staining and microchromosomal variation in karyotypes of four Brazilian species of Tupinambinae lizards (Squamata : Teiidae)

Kayotypes of four neotropical teiid lizard species (Tupinambinae) were herein studied after conventional as well as silver staining and CBG-banding: Crocodilurus amazonicus (2n = 34), Tupinambis teguixin (2n = 36), Tupinambis merianae and Tupinambis quadrilineatus (2n = 38). The karyological data for T. quadrilineatus as well as those obtained using differential staining for all species were unknown until now. The karyotypes of all species presented 12 macrochromosomes identical in morphology, but differed in the number of microchromosomes: 22 in C. amazonicus, 24 in T. teguixin and 26 in T. quadrilineatus and T. merianae. The Ag-NOR located at the secondary constriction at the distal end of pair 2 is shared by all species, contrasting with the variability observed for this character in species of the related Teiinae. CBG-banding revealed a species-specific pattern in T. quadrilineatus with conspicuous interstitial C-blocks at the proximal region of the long arm of pair 4 and the whole heterochromatic short arm of pair 6. The karyological data reported here corroborates the relationship hypothesis obtained for Tupinambis based on molecular characters. T. teguixin presents the putative ancestral karyotype for the genus with 2n = 36 whereas T. merianae and T. quadrilineatus exhibit 2n = 38, due to an additional pair of microchromosomes.

Head shape evolution in Tropidurinae lizards: does locomotion constrain diet?

Different components of complex integrated systems may be specialized for different functions, and thus the selective pressures acting on the system as a whole may be conflicting and can ultimately constrain organismal performance and evolution. The vertebrate cranial system is one of the most striking examples of a complex system with several possible functions, being associated to activities as different as locomotion, prey capture, display and defensive behaviours. Therefore, selective pressures on the cranial system as a whole are possibly complex and may be conflicting. The present study focuses on the influence of potentially conflicting selective pressures (diet vs. locomotion) on the evolution of head shape in Tropidurinae lizards. For example, the expected adaptations leading to flat heads and bodies in species living on vertical structures may conflict with the need for improved bite performance associated with the inclusion of hard or tough prey into the diet, a common phenomenon in Tropidurinae lizards. Body size and six variables describing head shape were quantified in preserved specimens of 23 species, and information on diet and substrate usage was obtained from the literature. No phylogenetic signal was observed in the morphological data at any branch length tested, suggesting adaptive evolution of head shape in Tropidurinae. This pattern was confirmed by both factor analysis and independent contrast analysis, which suggested adaptive co-variation between the head shape and the inclusion of hard prey into the diet. In contrast to our expectations, habitat use did not constrain or drive head shape evolution in the group.

Répteis da Estação Ecológica Serra Geral do Tocantins, Brasil Central

Os Cerrados sul-americanos abrigam alta diversidade de répteis, incluindo elevado número de endemismos. No entanto, o conhecimento desta diversidade é ainda incompleto frente à acelerada transformação das paisagens naturais no Brasil central. Constituem, portanto, uma das regiões prioritárias para estudo e conservação da biodiversidade mundial. Estudos intensivos sobre a fauna de répteis do Cerrado são necessários e urgentes para melhor compreensão dos processos que levaram à sua origem e distribuição e para subsidiar ações de conservação. Por meio de métodos padronizados, amostramos duas regiões ainda inexploradas da Estação Ecológica Serra Geral do Tocantins, situada na região do Jalapão. Registramos 45 espécies de répteis para a EESGT e entorno, o que representa uma riqueza alta e comparável à de outras regiões bem amostradas do Cerrado. Curvas de acumulação e estimadores indicam que a riqueza local de lagartos e anfisbenídeos aproxima-se da riqueza real enquanto a de serpentes é subestimada. A distribuição não-aleatória das espécies na paisagem concorda com evidências anteriores sugerindo utilização diferencial dos hábitats pelos répteis. Reunindo os resultados do presente estudo com os de levantamentos prévios realizados na região, registramos 88 espécies de répteis para o Jalapão sendo oito registros novos que incluem Bachia oxyrhina uma espécie recém descrita da região. As espécies da área apresentam três padrões gerais de distribuição: (1) espécies endêmicas do Cerrado, (2) espécies compartilhadas com domínios da diagonal de formações abertas sul-americanas, e (3) espécies de ampla ocorrência, compartilhadas também com ecossistemas florestais. Prevalecem espécies de ampla distribuição, porém é grande o número de espécies típicas do Cerrado, incluindo cinco possivelmente endêmicas do Jalapão, e há contribuição importante da fauna da Caatinga. A distribuição dos répteis em escala local e regional demonstra a necessidade de considerar a heterogeneidade paisagística para o planejamento de diretrizes visando à conservação em regiões do Cerrado. Por sua grande extensão, posição biogeográfica e complexidade de relevo e tipos de hábitat, a EESGT tem papel fundamental para a preservação e conhecimento da diversidade de répteis do Cerrado.

Herpetofauna do Parque Estadual Carlos Botelho, São Paulo, Brasil

O presente trabalho consiste em um inventário da herpetofauna do Parque Estadual Carlos Botelho (PECB), localizado na região da Serra de Paranapiacaba, Estado de São Paulo. Os dados foram obtidos por meio de coletas em seis áreas dentro do PECB durante um período de 76 dias distribuídos em um ano, e também por meio de consulta a coleções científicas para obtenção de dados secundários. São apresentados resultados sobre a biologia e ocorrência das espécies no PECB e no Brasil, além de fotografias das diferentes espécies que compõem a herpetofauna do PECB. A herpetofauna do PECB pode ser considerada uma das mais diversificadas de São Paulo, com 65 espécies de anfíbios confirmadas e 59 espécies de répteis registrados neste trabalho. Das 65 espécies de anfíbios, 84% (55 spp.) são endêmicas das formações florestais da Mata Atlântica. Devido às características do relevo do PECB, foram encontrados diferentes padrões altitudinais para os anfíbios: 46% das espécies foram registradas apenas em altitudes acima de 500 m, enquanto que 9% são exclusivas das regiões abaixo de 400 m e 45% ocorrem em todas as áreas amostradas do Parque. Das 59 espécies de répteis do PECB, foram registradas 10 espécies de lagartos, 48 de serpentes e um quelônio. Dentre as serpentes coletadas no PECB, a jararaca Bothrops jararaca foi a mais frequente, com 26,9% (N = 14) do total registrado. Espécies de difícil amostragem, como Echinanthera cephalostriata (13,5%; N = 7) e Taeniophallus affinis (7,7%; N = 4), também foram numerosas no PECB. Dentre os lagartos, Enyalius iheringii foi a espécie mais abundante, com 50% (N = 16) de registros. Uma análise de agrupamento entre 25 taxocenoses de anfíbios brasileiras, incluindo o PECB, resultou na formação de quatro agrupamentos principais. A anurofauna do PECB é mais relacionada com as taxocenoses do Parque Estadual Jacupiranga (0,68) e do Parque Estadual Intervales (0,66). Estes Parques se encontram geograficamente próximos e constituem um dos maiores fragmentos preservados de Mata Atlântica no Brasil. Este trabalho é o primeiro a apresentar a lista de répteis do PECB, alem de complementar o conhecimento em relação a fauna de anfíbios do PECB.

Ocorrência de Giardia, Cryptosporidium e microsporídios em animais silvestres em área de desmatamento no Estado de São Paulo, Brasil

A ocorrência de Giardia, Cryptosporidium e microsporídios foi investigada por meio da análise de 98 amostras fecais de animais silvestres capturados em uma área de desmatamento para a construção das barragens de Paraitinga e Biritiba, localizadas nos Municípios de Mogi das Cruzes, Salesópolis e Biritiba-Mirim, no Estado de São Paulo. As amostras foram obtidas de 46 roedores, 21 marsupiais, 16 sapos, nove morcegos, três primatas e três lagartos. As técnicas de centrífugo-flutuação com sulfato de zinco, de Kinyoun e a coloração de Gram-Chromotrope foram utilizadas, respectivamente, para a pesquisa de Giardia, de Cryptosporidium e de microsporídios. O total de animais parasitados por um dos protozoários investigados foi de 17,35% (17/98). Cistos de Giardia foram encontrados em amostras fecais de dois pequenos roedores da espécie Coendou villosus (ouriço-cacheiro). Os três animais positivos para Cryptosporidium foram roedores das espécies Akodon montensis, Thaptomys nigrita (ambos conhecidos como ratos do mato) e Sciurus aestuans (serelepe ou caxinguelê). Esporos de microsporídios foram encontrados nas fezes de 12 animais, sendo seis roedores das espécies Oligoryzomys sp.(um), Akodon montensis (três) e Coendou villosus (dois), três marsupiais pertencentes às espécies Didelphis aurita (dois) e Marmosops incanus (um) e três morcegos da espécie Diphylla ecaudata. Este é o primeiro relato de microsporidiose em animais silvestres no Brasil. A presente investigação enfatiza a importância de animais silvestres, particularmente pequenos mamíferos, como potenciais fontes de infecção desses protozoários para outras populações animais, incluindo o homem, em áreas de desmatamento.

Status of Early 19th-Century Names Authored in Parallel by Wied and Schinz for South American Reptiles and Amphibians, with Designations of Three Nomina Protecta

Prince Maximilian zu Wied's great exploration of coastal Brazil in 1815-1817 resulted in important collections of reptiles, amphibians, birds, and mammals, many of which were new species later described by Wied himself The bulk of his collection was purchased for the American Museum of Natural History in 1869, although many ""type specimens"" had disappeared earlier. Wied carefully identified his localities but did not designate type specimens or type localities, which are taxonomic concepts that were not yet established. Information and manuscript names on a fraction (17 species) of his Brazilian reptiles and amphibians were transmitted by Wied to Prof. Heinrich Rudolf Schinz at the University of Zurich. Schinz included these species (credited to their discoverer ""Princ. Max."") in the second volume of Das Thierreich ... (1822). Most are junior objective synonyms of names published by Wied. However, six of the 17 names used by Schinz predate Wied's own publications. Three were manuscript names never published by Wied because he determined the species to be previously known. (1) Lacerta vittata Schinz, 1822 (a nomen oblitum) = Lacerta striata sensu Wied (a misidentification, non Linnaeus nec sensu Merrem) = Kentropyx calcarata Spix, 1825, herein qualified as a nomen protectum. (2) Polychrus virescens Schinz, 1822 = Lacerta marmorata Linnaeus, 1758 (now Polychrus marmoratus). (3) Scincus cyanurus Schinz, 1822 (a nomen oblitum) = Gymnophthalmus quadrilineatus sensu Wied (a misidentification, non Linnaeus nec sensu Merrem) = Micrablepharus maximiliani (Reinhardt and Lutken, ""1861"" [1862]), herein qualified as a nomen protectum. Qualifying Scincus cyanurus Schinz, 1822, as a nomen oblitum also removes the problem of homonymy with the later-named Pacific skink Scincus cyanurus Lesson (= Emoia cyanura). The remaining three names used by Schinz are senior objective synonyms that take priority over Wied's names. (4) Bufo cinctus Schinz, 1822, is senior to Bufo cinctus Wied, 1823; both, however, are junior synonyms of Bufo crucifer Wied, 1821 = Chaunus crucifer (Wied). (5) Agama picta Schinz, 1822, is senior to Agama picta Wied, 1823, requiring a change of authorship for this poorly known species, to be known as Enyalius pictus (Schinz). (6) Lacerta cyanomelas Schinz, 1822, predates Teius cyanomelas Wied, 1824 (1822-1831) both nomina oblita. Wied's illustration and description shows cyanomelas as apparently conspecific with the recently described but already well-known Cnemidophorus nativo Rocha et al., 1997, which is the valid name because of its qualification herein as a nomen protectum. The preceding specific name cyanomelas (as corrected in an errata section) is misspelled several ways in different copies of Schinz's original description (""cyanom las,"" ""cyanomlas,"" and cyanom""). Loosening, separation, and final loss of the last three letters of movable type in the printing chase probably accounts for the variant misspellings.