Patterns of infestation by the trombiculid mite Eutrombicula alfreddugesi in four sympatric lizard species (genus Tropidurus) in northeastern Brazil

We studied the parasitism by the chigger mite Eutrombicula alfreddugesi on four sympatric lizard species of the genus Tropidurus in Morro do Chapéu, Bahia state, Brazil: T. hispidus, T. cocorobensis, T. semitaeniatus and T. erythrocephalus. For each species, we investigated the patterns of infestation and analyzed to which extent they varied among the hosts. We calculated the spatial niche breadth of the chigger mite on the body of each host species and the distribution of mites along the hosts' bodies for each Tropidurus species. All four species of Tropidurus at Morro do Chapéu were parasited by the chigger mite, with high (97-100%) prevalences. Host body size significantly explained the intensity of mite infestation for all species, except T. erythrocephalus. The body regions with highest intensity of infestation in the four lizard species were the mite pockets. The spacial niche width of the chigger varied consistently among the four lizards species studied being highest for T. erytrocephalus and lowest for T. cocorobensis. We conclude that the distribution and intensity with which lizards of the genus Tropidurus are infested by Eutrombicula alfreddugesi larvae results from the interaction between aspects of host morphology (such as body size and the occurrence and distribution of mite pockets) and ecology (especially microhabitat use).     

Helminths of the exotic lizard Hemidactylus mabouia from a rock outcrop area in southeastern Brazil

The helminth fauna of 291 Hemidactylus mabouia (Lacertilia: Gekkonidae) from a rock outcrop area in the state of S?o Paulo, Southeastern Brazil, was studied. Five species were recovered, namely one unidentified species of centrorhynchid acanthocephalan (present only as cystacanths) and the nematodes Parapharyngodon sceleratus, P. largitor (Oxyuroidea: Pharingodonidae), Physaloptera sp. (Spiruroidea: Physalopteridae) and one indeterminate species of Acuariidae (Acuaroidea), with the latter two forms present only as larvae. Infection rates tended to increase with host size, but appeared to be unaffected by season. Hemidactylus mabouia shared most of its helminth fauna with two other sympatric lizard hosts, Mabuya frenata and Tropidurus itambere. The helminth assemblage of the H. mabouia population appears to have been entirely acquired by this exotic gecko from the local helminth species pool, rather than possessing any species from the parasite faunas of the original African populations.     

Body temperatures of Tropidurus torquatus (Squamata,Tropiduridae) from coastal populations: Do body temperatures vary along their geographic range?

We studied the geographic variation in body temperatures of the lizard Tropidurus torquatus in 10 restinga populations along approximately 1500 km of Brazilian coast. The mean activity body temperatures (30.8–36.2 °C) seems to remain constant along the populations and the differences recorded among them result from the adjustment of each lizard population to the local thermal environment (i.e. may express in part the local microhabitat temperatures occurring in each of the localities in that particular moment). Forested and open habitat tropidurine species have different mean activity body temperatures, probably as a consequence of the different thermal environments in these habitats.     

Feeding ecology of the lizard Tropidurus oreadicus Rodrigues 1987 (Tropiduridae) at Serra dos Carajás, Pará state, northern Brazil.

Tropidurus species commonly prey on arthropods, but they may also feed on vertebrates and plant material. The lizard Tropidurus oreadicus (Tropiduridae) is common in open vegetation habitats and generally has sexual dimorphism. In this study we analyzed the diet of T. oreadicus at Serra dos Carajás, Pará, in the north of Brazil. Snout-vent length (SVL) and jaw width (JW) were taken for 34 lizards. There was a significant difference in SVL and in JW, with males being larger than females. All lizards analyzed contained food in their stomachs. The diet of T. oreadicus at Serra dos Carajás was characterized by the consumption of a relative wide spectrum of food item categories (21 types of items), consisting of arthropods, part of one vertebrate and plant material, which characterizes the diet of a generalist predator. Volumetrically, the most important items in the diet of both sexes of T. oreadicus were flowers (M = 61.7%; F = 33%) and orthopterans (M = 1.7%; F = 3.5%). Ants were the most frequently consumed (100% for both sexes) and the most numerous (M = 94.5%; F = 89.4%) food item. Flowers also were frequently consumed (M = 91.7%; F = 54.5%), with their relative consumption differing significantly between sexes. There was not a significant sexual difference in prey volume, neither in number of preys per stomach, nor in type of prey ingested. There was no relationship between lizard jaw width and the mean volume of prey. The data showed that T. oreadicus is a relatively generalist lizard in terms of diet and that consumes large volumes of plant material, especially flowers of one species of genus Cassia.     

A phylogenetic analysis of the tropidurine lizards (Squamata: Tropiduridae), including new characters of squamation and epidermal microstructure

Characters of scale surface microstructure are combined with 'traditional'morphological characters in a phylogenetic analysis of the Tropidurini. Tropidurid lizards show variation in types of coarse and fine scale surface microstructure, in the anatomical distribution of different scale surface features, and in scale organ morphology and distribution. The morphology of the inner surface zone of scales is here described for the first time using scanning electron microscopy. Our phylogeny differs considerably from those proposed in earlier studies. New characters and frequency coding of polymorphic characters help resolve the problematic relationships of several species. Statistical confidence supports recognition of one large cis-Andean and one large trans-Andean clade of species. Based on our results, we synonymize Plesiomicrolophus with Microlophus and Uranoscodon with Tropidurus. The phylogenetic relationships of newly discovered Tropidurus are resolved: T. caltathelys is the sister species of T. melanopleurus; T. xanthochilus is the sister species of T. spinulosus. Tropidurus spinulosus is found to be more closely related to T. strobilurus and a clade of Amazonian species than to T. melanopleurus. The species previously placed in Uracentron are more closely related to species previously placed in Plica than to 77 strobilurus as previously thought.     

Phylogenetic relationships of South American lizards of the genus Stenocercus (Squamata: Iguania): A new approach using a general mixture model for gene sequence data

The South American iguanian lizard genus Stenocercus includes 54 species occurring mostly in the Andes and adjacent lowland areas from northern Venezuela and Colombia to central Argentina at elevations of 0-4000m. Small taxon or character sampling has characterized all phylogenetic analyses of Stenocercus, which has long been recognized as sister taxon to the Tropidurus Group. In this study, we use mtDNA sequence data to perform phylogenetic analyses that include 32 species of Stenocercus and 12 outgroup taxa. Monophyly of this genus is strongly supported by maximum parsimony and Bayesian analyses. Evolutionary relationships within Stenocercus are further analyzed with a Bayesian implementation of a general mixture model, which accommodates variability in the pattern of evolution across sites. These analyses indicate a basal split of Stenocercus into two clades, one of which receives very strong statistical support. In addition, we test previous hypotheses using non-parametric and parametric statistical methods, and provide a phylogenetic classification for Stenocercus.     

Sintopy of two Tropidurus lizard species (Squamata: Tropiduridae) in a rocky Cerrado habitat in central Brazil.

We studied the ecology of Tropidurus itambere and T. oreadicus that occur syntopically in rocky habitats of Cerrado vegetation in central Brazil during the dry season (April to September 2000). The two species are ecologically similar, but somewhat differentiated in vertical microhabitat use. The two species preferred rocky surface microhabitat. Both species demonstrated a unimodal activity pattern, with a peak between 10 and 15 h. Their diets were similar in composition and prey size. The most frequent item used by both species was ants, whereas the most important preys volumetrically were termites and ants. Small morphological differences observed between the two Tropidurus species could explain minor microhabitat divergence: T. itambere is slightly smaller, heavier, and more robust, and uses lower perches. T. oreadicus is larger, lankier, with longer extremities (tail, fore- and hindlegs), and uses a larger vertical microhabitat range. These ecological differences are slight, when compared with those observed between sympatric species of Tropidurus in spatially more heterogeneous landscapes. Considering the slight ecomorphological divergence between the two Tropidurus species and their high abundance in outcrops, we suggest that interspecific territoriality is the mechanism of coexistence.     

Extreme specialization to rocky habitats in Tropidurus lizards from Brazil: Trade‐offs between a fitted ecomorph and autoecology in a harsh environment

Ecomorphological theory indicates that different ecological requirements lead to different organismal designs. Given that species with equal requirements could not coexist, traits leading to more efficient use of resources may be selected to avoid competition among closely related syntopic species, generating specialized ecomorphs. We compared habitat use, diet, thermal biology and morphology among the syntopic Tropidurus semitaeniatus, T. helenae and T. hispidus in the Caatinga of Northeastern Brazil. Tropidurus semitaeniatus and T. helenae are flattened lizards specialized to rocks and rock crevices, whereas T. hispidus has a robust body and generalist habits. We aimed to test the hypothesis that morphological modifications observed in the flattened ecomorphs are related to modifications in diet and habitat use. Also, we hypothesized that specialization to habitat induces morphological modifications, which in turn may constrain lizard performance. Flattened species differed in habitat use, morphology and prey size when compared with the generalist ecomorph. Morphological modifications were related to specializations to rocky habitats and constrained the variety of prey items consumed. This phenotype also reduced their reproductive output when compared with a robust, generalist ecomorph.     

Spatial and trophic niche differentiation in two sympatric lizards (Tropidurus torquatus and Cnemidophorus ocellifer) with different foraging tactics

Abstract In a ‘restinga’ habitat of southeastern Brazil, we studied the food habits and the microhabitat use of two lizards with distinct foraging modes: the tropidurid Tropidurus torquatus, a sit-and-wait predator, and the teiid Cnemidophorus ocellifer, a wide forager. The diet of the two species differed strongly, indicating a low level of similarity in their trophic niche. The sit-and-wait predator fed mainly on mobile prey, whereas the wide forager fed mainly on sedentary prey (larvae). The spatial niche breadth of T. torquatus was larger than that of C. ocellifer. Despite interspecific differences, the two species overlapped greatly in micro-habitat use. The data indicate that at Linhares the two lizard species differed more in food resources than in microhabitat, and that most of the food differences reflect the foraging patterns of the species.     

Chemical signalling behaviour in intrasexual communication of lizards lacking femoral pores

Signals that are used in animal communication may have multiple sensorial channels and functions. Animal communication integrates very distinct mechanisms such as behaviour, morphology and physiological secretions. Chemical signals occur in several contexts and are known to have a role in sexual selection. In many lizards, pheromones secreted through femoral pores are used to attract females and demark territory dominance. In lizard species without femoral pores, however, study of chemical signalling is negligible. Lizards of the genus Tropidurus have no ventral pores, but express melanic patches in their ventral thighs and cloacae (body regions linked to chemical communication in other lizards), which may play a role in both visual signalling and chemical signalling. Here, we describe the occurrence of pelvic rubbing—a chemical signalling behaviour—in two Tropidurus species, displayed in intraspecific agonistic intrasexual staged encounters. In addition, we compile a list of currently reported species that display this behaviour, checking for its social contexts and presence/absence of secretion pores. We analysed behaviours in conspecific trials between males of Tropidurus semitaeniatus and of T. hispidus during their breeding season. In T. semitaeniatus, displays of pelvic rubbing elicited aggressive responses from the opponents. Tropidurus hispidus, however, did not react to these conspecific displays. Pelvic rubbings were also partially linked to defecation in both species, which is likely due to secretions from internal urodeal glands expelled during defecation. Also, high rates of tongue flick behaviours during the encounters support the hypothesis that these lizards make use of chemical communication. Our observations corroborate previous suggestions that glandular scales — scales covered by generation glands produced at the epidermis across the lizards’ shedding cycles — are present in their melanic ventral patches. Our work also highlights the potential of characterizing such ventral patches as multimodal signalling badges. Further, we investigated the contexts in which pelvic rubbing is displayed adding to our understanding of the role played by chemical signalling in lizards without femoral pores.     

Helminth community structure of Tropidurus torquatus (Squamata: Tropiduridae) in a rocky outcrop area of Minas Gerais state, southeastern Brazil

The helminth community of Tropidurus torquatus (Squamata: Tropiduridae) from a rocky outcrop area located in the state of Minas Gerais, southeastern Brazil, was studied. Ninety-two of the 110 individuals examined (83.6%) harbored helminths. Five species were found, including 3 nematodes ( Physaloptera lutzi , Parapharyngodon bainae , and Oswaldofilaria chabaudi ), 1 unidentified cestode species, and 1 acanthocephalan cystacanth also not identified. Only the nematode species had sufficient data to perform ecological analysis, with P. lutzi exhibiting the highest prevalence (67.3%). Prevalence between male and female hosts differed only for Oswaldofilaria chabaudi , with males exhibiting the highest values. The intensities of infection by P. lutzi and O. chabaudi were different among male and female hosts, with males also exhibiting the highest values. The host body size was positively related to intensity of infection for all nematode species. Local seasonality had some influence on the helminth community structure. Host diet, sexual dimorphism, and behavior (territorialism, forage strategy) represented important factors for the structure of this parasite community. In general, the helminth community was species poor, depauperate, and non-interactive, representing a typical structure observed in lizard hosts.     

Escape tactics by a neotropical montane lizard: a comparison of flight responses against natural and nonnatural predators

Prey must judge the level of threat related to a given predator attack to employ the most efficient escape strategy at a low cost. In this context, the capacity of prey to correctly assess the threat from predators might optimise their decision on the strategy employed to flee. We examined the escape behaviour of the montane lizard Tropidurus montanus in response to different types of predatory stimuli. Lizards stimulated by a model of a natural predator ran shorter distances straight towards shelter and spent less time in flight. On the other hand, when exposed to a model of a nonnatural predator, T. montanus performed longer flights in distance and time, running across open and exposed substrates in the view of predators. These results showed that the lizard T. montanus made different escape decisions according to the type of predatory stimulus.     

Elevational gradients of diversity for lizards and snakes in the Hengduan Mountains,China

Comparing elevational gradients across a wide spectrum of climatic zones offers an ideal system for testing hypotheses explaining the altitudinal gradients of biodiversity. We document elevational patterns of lizard and snake species richness, and explore how land area and climatic factors may affect species distributions of lizards and snakes. Our synthesis found 42 lizard species and 94 snake species known from the Hengduan Mountains. The lizards are distributed between 500 and 3500 m, and the snakes are distributed between 500 and 4320 m. The relationship between species richness and elevation for lizards and snakes is unimodal. Land area explains a significant amount of the variation in lizard and snake species richness. The cluster analysis reveals pronounced distinct assemblages for lizards and snakes to better reflect the vertical profiles of climate in the mountains. Climatic variables are strongly associated with lizard and snake richness along the elevational gradient. The data strongly implicate water availability as a key constraint on lizard species richness, and annual potential evapotranspiration is the best predictor of snake species richness along the elevational gradient in the Hengduan Mountains.     

Low temperature-acclimation impairs cellular migration in the adult cerebral cortex of the tropical lizard, Tropidurus hispidus (Spix, 1825) (Squamata: Tropiduridae)

The effects of different temperature-acclimations on cellular proliferation and migration were studied in the cerebral cortex of the tropical lizard, Tropidurus hispidus. Lizards were divided in two groups: warm-acclimated lizards (WALs), maintained at the temperature and photoperiod conditions of their natural habitat (mean temperature 26°C; 12:12 light:dark) and the cold-acclimated lizards (CALs), maintained at the same cycle of illumination and a mean temperature of 16°C. Animals were injected with the proliferative marker 5-bromodeoxyuridine (BrdU) and euthanized fifteen or thirty days later for the immunostaining. There was no difference in the number of BrdU-positive nuclei between the experimental groups in any of the cortical layers. In CALs, the positive nuclei were found mostly close to the ependyma, whereas in WALs many positive nuclei were also found in the plexiform and cellular layers of the cortex. In CALs, BrdU-positive nuclei appeared grouped (of 2-3 nuclei), a characteristic not seen in the other group. These data suggest that temperature affects the migrating capability of the newly generated neurons in the lizard cortex, but appears not to interfere with its generation.     

Chromosome mechanisms of sex determination,G- and C-band patterns and nucleolus organizer regions in Tropidurus torquatus (Sauria,Iguanidae)

The mitotic and meiotic chromosomes of two groups of the iguanid lizard Tropidurus torquatus were studied after conventional staining, G- and C-banding and silver staining of nucleolar-organizer regions. Karyotypic variations involving the microchromosomes were observed in both groups and are, probably, related to the mechanisms of sex determination of the X₁X₂Y/X₁X₁X₂X₂ type in the one group, and of the XY/XX type in the other.     

Pollination and seed dispersal of Melocactus ernestii Vaupel subsp. ernestii (Cactaceae) by lizards: an example of double mutualism

Recent studies show that the mutualistic role of lizards as pollinators and seed dispersers has been underestimated, with several ecological factors promoting such plant–animal interactions, especially on oceanic islands. Our aim is to provide a quantitative assessment of pollination and seed dispersal mutualisms with lizards in continental xeric habitats. We carried out focal observations of natural populations of Melocactus ernestii (Cactaceae) in the Caatinga, a Brazilian semiarid ecosystem, in order to record the frequency of visits, kind of resource searched and behaviour of visiting animals towards flowers and/or fruits. We made a new record of the lizard Tropidurus semitaeniatus foraging on flowers and fruits of M. ernestii. During the search for nectar, T. semitaeniatus contacted the reproductive structures of the flowers and transported pollen attached to its snout. Nectar production started at 14:00 h, with an average volume of 24.4 μl and an average concentration of solutes of 33%. Approximately 80% of the seeds of M. ernestii found in the faeces of T. semitaeniatus germinated under natural conditions. The roles of T. semitaeniatus as pollinator and seed disperser for M. ernestii show a clear relationship of double mutualism between two endemic species, which may result from the environmental conditions to which both species are subject. Seasonality, low water availability and arthropod supply in the environment, high local lizard densities, continuous nectar production by the flower and fruits with juicy pulp may be influencing the visits and, consequently, pollination and seed dispersal by lizards in this cactus.     

Implications of climate change on the habitat shifts of tropical lizards

The effect of temperature on the distributions of ectothermic vertebrates is well documented. Despite the increase of 6°C expected in the next 60 years in South America, numerous vertebrates are still considered as ‘Least Concern’ species by the IUCN due to their large distribution, insufficient widespread threats and insignificant population decline. One example is the lizard Tropidurus torquatus (Squamata: Tropiduridae), commonly found thermoregulating in anthropic environments throughout the Brazilian Cerrado, but restricted to gallery forests in the equator‐ward localities. The urban areas in this warmer region have been colonised by other closely related congeners (e.g. Tropidurus oreadicus). This study aimed to understand this divergence of habitat selection by these tropirudids that may explain some of the species responses to past and future climate warming. We collected body temperatures (T_b), micro‐environmental temperatures (T_a) and operative (T_e) temperatures in four sites along a latitudinal gradient: a pole‐ward and two central sites where T. torquatus inhabit urban areas and one equator‐ward site where T. torquatus and T. oreadicus occur in the gallery forest and in urban microhabitats, respectively. All three populations of T. torquatus present similar T_b (35.5–36°C) and shared microhabitats with a similar T_a (34–37.3°C). The T_e in the equator‐ward urban site was considerably higher than in the gallery forest. Tropidurus oreadicus T_b was 38.2 °C (30.1–41.3°C) and was active at a T_a of 30.5–42.3°C. The overlap between the genus T_b, T_a and T_e highlights a decrease in the hours of activity that lizards would experience under climate warming. The reduction of hours of activity together with the devastation of natural habitats represents threats and an alarming scenario especially for the equator‐ward populations.     

Local richness and distribution of the lizard fauna in natural habitat mosaics of the Brazilian Cerrado

We investigate local lizard richness and distribution in central Brazilian Cerrado, harbouring one of the least studied herpetofaunas in the Neotropical region. Our results are based on standardized samplings at 10 localities, involving 2917 captures of 57 lizard species in 10 families. Local richness values exceeded most presented in earlier studies and varied from 13 to 28 species, with modal values between 19 and 28 species. Most of the Cerrado lizard fauna is composed of habitat‐specialists with patchy distributions in the mosaic of grasslands, savannas and forests, resulting in habitat‐structured lizard assemblages. Faunal overlap between open and forested habitats is limited, and forested and open areas may act as mutual barriers to lizard distribution. Habitat use is influenced by niche conservatism in deep lineages, with iguanians and gekkotans showing higher use of forested habitats, whereas autarchoglossans are richer and more abundant in open habitats. Contrary to trends observed in Cerrado birds and large mammals, lizard richness is significantly higher in open, interfluvial habitats that dominate the Cerrado landscape. Between‐localities variation in lizard richness seems tied to geographical distance, landscape history and phylogenetic constraints, factors operating in other well‐studied lizard faunas in open environments. Higher richness in dominant, open interfluvial habitats may be recurrent in Squamata and other small‐bodied vertebrates, posing a threat to conservation as these habitats are most vulnerable to the fast, widespread and ongoing process of habitat destruction in central Brazil.     

Microhabitat Variation Explains Local‐scale Distribution of Terrestrial Amazonian Lizards in Rondônia,Western Brazil

We investigate the role of ecology and phylogeny in the association between lizard abundance and microhabitat variables in an Amazon rain forest site. Using pitfall trap arrays, we collected data from 349 individuals belonging to 23 lizard species. After accounting for spatial autocorrelation and using a canonical correspondence analysis (CCA), we found that lizard captures were significantly associated with microhabitat variables, which accounted for 48 percent of the observed variation. Furthermore, a canonical phylogenetic ordination (CPO) indicated that microhabitat variables are more important in determining the distribution of lizard species than phylogenetic relationships among species. Termite nests, canopy openness, and tree circumference were strongly associated with the number of captures of certain lizard species. Our results confirm autecology studies of individual lizard species for which data are available. We suggest that maintaining heterogeneous forested microhabitats should be a central goal for sustaining a high lizard biodiversity in Amazon rain forests.