Chromosomal Evolution in the Brazilian Geckos of the Genus Gymnodactylus (Squamata, Phyllodactylidae) from the Biomes of Cerrado, Caatinga and Atlantic Rain Forest: Evidence of Robertsonian Fusion Events and Supernumerary Chromosomes

Chromosomes of the South American geckos Gymnodactylus amarali and G. geckoides from open and dry areas of the Cerrado and Caatinga biomes in Brazil, respectively, were studied for the first time, after conventional and AgNOR staining, CBG- and RBG-banding, and FISH with telomeric sequences. Comparative analyses between the karyotypes of open areas and the previously studied Atlantic forest species G. darwinii were also performed. The chromosomal polymorphisms detected in populations of G. amarali from the states of Goias and Tocantins is the result of centric fusions (2n = 38, 39 and 40), suggesting a differentiation from a 2n = 40 ancestral karyotype and the presence of supernumerary chromosomes. The CBG- and RBG-banding patterns of the Bs are described. G. geckoides has 40 chromosomes with gradually decreasing sizes, but it is distinct from the 2n = 40 karyotypes of G. amarali and G. darwinii due to occurrence of pericentric inversions or centromere repositioning. NOR location seems to be a marker for Gymnodactylus, as G. amarali and G. geckoides share a medium-sized subtelocentric NOR-bearing pair, while G. darwinii has NORs at the secondary constriction of the long arm of pair 1. The comparative analyses indicate a non-random nature of the Robertsonian rearrangements in the genus Gymnodactylus. Copyright (C) 2010 S. Karger AG, Basel

Karyotypes of a Cryptic Diploid Form of the Unisexual Leposoma percarinatum (Squamata, Gymnophthalmidae) and the Bisexual Leposoma ferreirai from the Lower Rio Negro, Amazonian Brazil

Karyotypes of Leposoma show a clear differentiation between species of the scincoides group from Brazilian Atlantic Forest (2n = 52, without distinctive size groups of chromosomes) and those of the parietale group from the Amazon (2n = 44, with 20M + 24m). In a previous study, we found that in the parietale group the parthenoform Leposoma percarinatum from the state of Mato Grosso, Brazil, exhibited a triploid karyotype (3n = 66) with 30 macrochromosomes and 36 microchromosomes. It was suggested that this karyotype arose after hybridization between a bisexual species with N = 22 (10M + 12m) and a hypothetical unisexual cryptic diploid form of the L. percarinatum complex. Herein, we describe the karyotypes for two species of the parietale group occurring sympatrically in the Arquipelago das Anavilhanas, lower Rio Negro, in Amazonian Brazil. The first represents a distinctive diploid parthenogenetic clone of the L. percarinatum complex, and the other is the recently described Leposoma ferreirai. Both species have 44 biarmed chromosomes clearly represented by 20 macrochromosomes and 24 microchromosomes and present Ag-NORs in one pair of the smallest sized microchromosomes; heteromorphism of size for these regions was detected in L. percarinatum. C-banding revealed blocks of constitutive heterochromatin on the telomeric and pericentromeric regions of macrochromosomes and some microchromosomes. The description of a diploid karyotype (2n = 44, 20M + 24m) for the L. percarinatum complex and its sympatric congener L. ferreirai provides new insight for a better understanding of the origin of parthenogenesis in the L. percarinatum complex.

Ecologia comportamental de tropidurus Hispidus e Tropidurus Semitaeniatus (Squamata, Tropiduridae) em Simpatria, em área de caatinga do nordeste do Brasil.

This study evaluated the spatial, time and alimentary niches of Tropidurus hispidus and Tropidurus semitaeniatus in sympatry in a caatinga of Rio Grande do Norte, Brazil, as well as their foraging and termoregulatory behaviors, the activity body temperature and their reproductive and fat body cycles. Monthly excursions, from October 2006 to May 2008, were conducted at the Ecological Station of the Seridó (ESEC Seridó), Serra Negra do Norte municipality, using specific methodology for investigation of the aforementioned objectives. The two species presented similarities in space niche use, mainly in rocky habitat, however they differed in vertical microhabitat use with T. hispidus using a larger vertical microhabitat range. In the dry season the time of activity of both species was bimodal. In the wet season T. semitaeniatus showed a unimodal activity period, while T. hispidus maintained an bimodal activity period. In terms of importance in the diet, to both species, Hymenoptera/Formicidae and Isoptera predominated during the dry season. In the wet season, although Hymenoptera/Formicidae had larger importance among the prey items, lizards opportunistically predated on Lepidoptera larvae, Coleoptera larvae/adults and Orthoptera nymphs/adults. The foraging intensity revealed differences between the species, mainly in the wet season, when T. semitaeniatus was more active than T. hispidus. The mean activity body temperature of T. semitaeniatus was significantly higher than that of T. hispidus. The thermoregulatory behavior showed that during the dry season T. hispidus and T. semitaeniatus spent more time in shade or under filtered sun. In the wet season, T. hispidus did not show differences in the amount of time spent among the light exposure locations, however T. semitaeniatus spent most of their time exposed to direct sun or filtered sun. The reproductive cicle of T. hispidus and T. semitaeniatus occurred from the middle of the dry season to the beginning of the wet season. In both species, female reproductive activity was influenced by precipitation, whereas males exhibited spermatozoa in their testes throughout the year, and their reproductive activity was not related with any of the climatic variables analysed. In the two species, the fat storage varied inversely with reproductive activity, and there was no difference in fat body mass between females and males. We concluded that the segregation between T. hispidus and T. semitaeniatus in this caatinga area occurs in vertical space use, in the largest vagility of T. hispidus in microhabitat use and larger range size of their alimentary xviii items. Additionally, significant seasonal differences in relation to the activity period, body temperature, and foraging and termoregulatory behaviors between these two Tropidurus species facilitate their coexistence.

Dieta e compotamento de forrageio de duas espécies de Tropidurus (Squamata,Tropiduridae) em uma área de Caatinga do Nordeste do Brasil

Tropidurus semitaeniatus and Tropidurus hispidus are lizard species that are widely distributed in Brazil. The former is endemic to the Caatingas, where they live in syntopy with the latter, which is distributed in other ecosystems and biomes. In this work, diet was evaluated including seasonal variations as well as foraging behavior, to determine the importance of these factors in the structure of these populations and the coexistence of these species in syntopy. No significant difference was found in diet composition between the two species, although T. hispidus feed on larger preys than do T. semitaeniatus; this fact, along with the different foraging behavior in the rainy season, are ecological factors that contribute to the syntopic coexistence of these species in the Caatinga area studied

Ecologia alimentar e comportamento de forrageamento de Ameivula aff. ocellifera (Squamata: Teiidae) em área de caatinga do nordeste do Brasil

This study investigated the influence of intrinsic and extrinsic factors on the feeding ecology and foraging behavior of the whiptail lizard Ameivula aff. ocellifera, a new species widely distributed in the Brazilian Caatinga, and that is in process of description. In attendance to the objectives, the Dissertation was structured in two chapters, which correspond to scientific articles, one already published and the other to be submitted for publication. In Chapter 1 were analyzed the general diet composition, the relationship between lizard size and prey size, and the occurrence of sexual and ontogenetic differences in the diet. Chapter 2 contemplates a seasonal analysis of diet composition during two rainy seasons interspersed with a dry season, and the quantitative analysis of foraging behavior during two distinct periods. The diet composition was determined through stomach analysis of lizards (N = 111) collected monthly by active search, between September 2008 and August 2010, in the Estação Ecológica do Seridó (ESEC Seridó), state of Rio Grande do Norte. Foraging behavior was investigated during a rainy and a dry month of 2012 also in ESEC Seridó, by determining percent of time moving (PTM), number of movements per minute (MPM) and prey capture rate by the lizards (N = 28) during foraging. The main prey category in the diet of Ameivula aff. ocellifera was Insect larvae, followed by Orthoptera, Coleoptera and Araneae. Termites (Isoptera) were important only in numeric terms, having negligible volumetric contribution (<2%) and low frequency of occurrence, an uncommon feature among whiptail lizards. Males and females did not differ neither in diet composition nor in foraging behavior. Adults and juveniles ingested similar prey types, but differed in prey size. Maximum and minimum prey sizes were positively correlated with lizard body size, suggesting that in this population individuals experience an ontogenetic change in diet, eating larger prey items while growing, and at the same time excluding smaller ones. The diet showed significant seasonal differences; during the two rainy seasons (2009 and 2010), the predominant prey in diet were Insect larvae, Coleoptera and Orthoptera, while in the dry season the predominant prey were Insect larvae, Hemiptera, Araneae and Orthoptera. The degree of mobility of consumed prey during the rainy seasons was lower, mainly due to a greater consumption of larvae (highly sedentary prey) during these periods. Population niche breadth was higher in the dry season, confirming the theoretical prediction that when food is scarce, the diets tend to be more generalized. Considering the entire sample, Ameivula aff. ocellifera showed 61,0 ± 15,0% PTM, 2,03 ± 0,30 MPM, and captured 0,13 ± 0,14 per minute. Foraging mode was similar to that found for other whiptail lizards regarding PTM, but MPM was relatively superior. Seasonal differences were verified for PTM, which was significantly higher in the rainy season (66,4 ± 12,1) than in the dry season (51,5 ± 15,6). It is possible that this difference represents a behavioral adjustment in response to seasonal variation in the abundance and types of prey available in the environment in each season

Frugivoria e dispersão de sementes pelo lagarto teiú Tupinambis merianae (Reptilia: Teiidae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Energetic and physiological correlates of prey handling and ingestion in lizards and snakes

In this review, we summarize the energetic and physiological correlates of prey handling and ingestion in lizards and snakes. There were marked differences in the magnitude of aerobic metabolism during prey handling and ingestion between these two groups, although they show a similar pattern of variation as a function of relative prey mass. For lizards, the magnitude of aerobic metabolism during prey handling and ingestion also varied as a function of morphological specializations for a particular habitat, prey type, and behavior. For snakes, interspecific differences in aerobic metabolism during prey handling seem to be correlated with adaptations for prey capture (venom injection vs. constriction). During ingestion by snakes, differences in aerobic metabolism might be due to differences in cranial morphology, although allometric effects might be a potentially confounded effect. Anaerobic metabolism is used for prey handling and ingestion, but its relative contribution to total ATP production seems to be more pronounced in snakes than in lizards. The energetic costs of prey handling and ingestion are trivial for both groups and cannot be used to predict patterns of prey-size selection. For lizards, it seems that morphological and ecological factors set the constraints on prey handling and ingestion. For snakes, besides these two factors, the capacity of the cardio-respiratory system may also be an important factor constraining the capacity for prey handling and ingestion. © 2001 Elsevier B.V.

Os Répteis do Município de São Paulo: Diversidade e ecologia da fauna pretérita e atual

The reptile fauna in the municipality of São Paulo is well sampled due to intensive collection in the last 100 years. In the present work we provide a checklist of reptile species in São Paulo municipality based on preserved specimens in scientific collections. The reptile fauna was also characterized by three ecological parameters: habitat use, substrate use, and diet. We recorded a total of 97 reptile species (two turtles, one crocodilian, 19 lizards, seven amphisbaenians, and 68 snakes). Approximately 70% of the lizards and 40% of the snakes are typical of forest habitats of the Serra do Mar mountain range. Other squamates are typical of open formations that occur mainly on inland Cerrado habitats. All turtles and the crocodilian are associated to riparian habitats. Approximately 63% of the lizards are predominantly terrestrial, and the remaining species are arboreal. Most species of snakes are terrestrial (38%) or subterranean/criptozoic (25%) whereas a smaller proportion are arboreal (18%) or aquatic (9%). Lizards feed upon arthropods. Almost 50% of the snake species are specialized or feed mainly upon anuran amphibians. Other important items consumed by snakes are mammals (24%), lizards (18%), subterranean vertebrates (10%), and invertebrates (earthworms, mollusks and arthropods; 15%). A total of 51 reptile species have not been recorded for the last six years. Probably many of these species are extinct in the region due the intense local urbanization and habitat loss. The survey of species collected in São Paulo municipality and received in the Instituto Butantan in recent years allowed the identification of 10 lizards at least 42 snake species already occurring in the region. The high species richness of the original fauna seems related to the geographic location of the municipality, in a contact zone between forested areas of the Atlantic Forest (ombrophilous forest) and open formations (savannas, high-altitude grassland). Thus, the original habitat composition probably allowed sympatry among different species pools typical of both open and forested formations. The extant snake fauna recorded in the last three years indicates a higher loss of the species in open formations when compared to the forested areas.

Variação sazonal da temperatura corpórea no lagarto Teiú, Tupinambis merianae (Squamata, Lacertilia, Teiidae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Taxocenose de serpentes (Squamata: Serpentes) em uma área de transição cerrado-caatinga no município de Castelo do Piauí, Piauí, Brasil

Foi realizado um inventário da fauna de serpentes da área de transição entre Cerrado e Caatinga no Município de Castelo do Piauí. Foram realizadas seis expedições entre Outubro de 2005 e Julho de 2006, em fitofisionomias de Cerrado Rupestre, Cerrado Aberto e Mata Secundária de Cerrado Típico, que totalizaram 120 dias de trabalho de campo. Foram utilizados três métodos: Armadilhas de Interceptação e Queda, Procura Limitada por Tempo e Encontros Ocasionais. Foram registradas 18 espécies. A família Colubridae apresentou a maior riqueza de espécies, assim como nas diferentes formações da América do Sul. A comunidade é composta por Xenodontinae (11 espécies), seguido por Colubrinae (Mastigodryas bifossatus e Spilotes pullatus) e apenas uma espécie de Dipsadinae (Leptodeira annulata), refletindo o padrão evolutivo das linhagens filogenéticas dos colubrídeos e a estrutura da comunidade. Thamnodynastes sp. (n= 7), Philodryas nattereri (n= 5) e Phimophis iglesiasi (n= 5) foram as espécies mais abundantes na área estudada, diferente da dominância de viperídeos observada em outras taxocenoses de serpentes no Brasil. Os estimadores de riqueza Chao 2 e Jack1 indicam que a comunidade é composta por aproximadamente 24 espécies. Procura Limitada por Tempo apresentou o melhor desempenho dentre os métodos utilizados, embora a utilização dos três métodos seja recomendada para um inventário mais completo de serpentes. Foi demonstrada a predominância de serpentes terrestres e criptozóicas, diurnas, ovíparas e que possivelmente alimentam-se na sua maioria de anfíbios e lagartos. A Análise de Coordenadas Principais, análise de agrupamento, similaridade e padrão de distribuição das espécies dentre os biomas, mostram que a área estudada é mais similar a áreas abertas (Cerrado, Caatinga e Pantanal). A comunidade de serpentes de Castelo do Piauí sofre influência direta da fauna dos biomas Cerrado e Caatinga, de acordo com os estudos florísticos da área, apontando associação preferencial a taxocenoses de caatinga. As análises do PCO e de Agrupamento sugerem que a hipótese sobre composição mista das faunas de cerrado e caatinga, como apontada em outros trabalhos, possa ser conseqüência de análises inadequadas.

Gastrointestinal nematodes of the lizard Tropidurus hispidus (Squamata: Tropiduridae) from a semi-arid region of north-eastern Brazil

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

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

Skull represents the segment with conspicuous adaptations that, in lizards, may be conservative or promoted by selective pressures. The aim of assisting the morphological knowledge of reptiles, we provide a detailed description of the neurocranium of Iguana iguana iguana based on analysis of three dried adult skeletons. The skull of this species has basal characteristics in lizards without closure of cranial openings and general triangular shape. Bony structures that form the caudal base have many fusions, especially on the floor. In the caudal face the exoccipital and the opisthotic are fused and form the otooccipital, which contributes to the formation of the lateral part of the condyle. The central part is formed by the condyle supraocciopital. Fusions and skeletal structures in Iguana are similar to other lizards. There are no autopomorphies in the neurocranium for this species.

Evolution of form and function: morphophysiological relationships and locomotor performance in tropidurine lizards

Locomotor capacity is often considered an excellent measure of whole animal performance because it requires the integrated functioning of many morphological, physiological (and biochemical) traits. However, because studies tend to focus on either structural or functional suits of traits, we know little on whether and how morphological and physiological traits coevolve to produce adequate locomotor capacities. Hence, we investigate the evolutionary relationships between morphological and physiological parameters related to exercise physiology, using tropidurine lizards as a model. We employ a phylogenetic principal component analysis (PCA) to identify variable clusters (factors) related to morphology, energetic metabolism and muscle metabolism, and then analyze the relationships between these clusters and measures of locomotor performance, using two models (star and hierarchical phylogenies). Our data indicate that sprint performance is enhanced by simultaneous evolutionary tendencies affecting relative limb and tail size and physiological traits. Specifically, the high absolute sprint speeds exhibited by tropidurines from the sand dunes are explained by longer limbs, feet and tails and an increased proportion of glycolytic fibers in the leg muscle, contrasting with their lower capacity for overall oxidative metabolism [principal component (PC1)]. However, when sprint speeds are corrected for body size, performance correlates with a cluster (PC3) composed by moderate loads for activity metabolic rate and body size. The simultaneous measurement of morphological and physiological parameters is a powerful tool for exploring patterns of coadaptation and proposing morphophysiological associations that are not directly predictable from theory. This approach may trigger novel directions for investigating the evolution of form and function, particularly in the context of organismal performance.

Six species of Amazonian Woodcreepers (Aves: Dendrocolaptidae) preying upon lizards and frogs

Dietary data from a large sample of woodcreepers (16 spp., n?=?139), revealed that six species of dendrocolaptids occasionally feed upon lizards and frogs. These birds, which are mainly insectivorous, encounter and feed on lizards while perching on tree trunks, probably in association with army-ant swarm feeding behaviour. Frog intake may be related to declines in the abundance of invertebrate prey. The bones recovered were identified as one small species of gecko, Gonatodes humeralis, and at least one anuran. We estimate that in the entire sample, about eight lizards and two frogs were ingested. The partially digested gecko material allows determination of which bones are more resistant to digestion, although it is possible that these elements were differentially retained in the stomach. These elements correspond to the more frequently preserved bones in the fossil record of geckos, indicating that the same portions of the skeleton persist under the processes of both digestion and fossilization.

Divergent calcium signaling in RBCs from Tropidurus torquatus (Squamata – Tropiduridae) strengthen classification in lizard evolution

Abstract Background We have previously reported that a Teiid lizard red blood cells (RBCs) such as Ameiva ameiva and Tupinambis merianae controls intracellular calcium levels by displaying multiple mechanisms. In these cells, calcium stores could be discharged not only by: thapsigargin, but also by the Na+/H+ ionophore monensin, K+/H+ ionophore nigericin and the H+ pump inhibitor bafilomycin as well as ionomycin. Moreover, these lizards possess a P2Y-type purinoceptors that mobilize Ca2+ from intracellular stores upon ATP addition. Results Here we report, that RBCs from the tropidurid lizard Tropidurus torquatus store Ca2+ in endoplasmic reticulum (ER) pool but unlike in the referred Teiidae, these cells do not store calcium in monensin-nigericin sensitive pools. Moreover, mitochondria from T. torquatus RBCs accumulate Ca2+. Addition of ATP to a calcium-free medium does not increase the [Ca2+]c levels, however in a calcium medium we observe an increase in cytosolic calcium. This is an indication that purinergic receptors in these cells are P2X-like. Conclusion T. torquatus RBCs present different mechanisms from Teiid lizard red blood cells (RBCs), for controlling its intracellular calcium levels. At T. torquatus the ion is only stored at endoplasmic reticulum and mitochondria. Moreover activation of purinergic receptor, P2X type, was able to induce an influx of calcium from extracelullar medium. These studies contribute to the understanding of the evolution of calcium homeostasis and signaling in nucleated RBCs.