Syntopic species of Tropidurus found at the Reserva Particular do Patrimônio Natural Fazenda Pé da Serra, Serra do Arame, Ibotirama, Bahia, Brazil, and their respective habitats: (A, B) T. sertanejo, n. sp. (MZUSP 104274, allotype); (C, D) T. hispidus (MZUSP 104276); (E, F) T. pinima (MZUSP 104271).  

Syntopic species of Tropidurus found at the Reserva Particular do Patrimônio Natural Fazenda Pé da Serra, Serra do Arame, Ibotirama, Bahia, Brazil, and their respective habitats: (A, B) T. sertanejo, n. sp. (MZUSP 104274, allotype); (C, D) T. hispidus (MZUSP 104276); (E, F) T. pinima (MZUSP 104271).  

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Tropidurus Wied, 1825, is one of the most ubiquitous lizard genera distributed in open habitats of tropical and subtropical South America. Nevertheless, the broad representation of specimens of this group in scientific collections is hardly reflected in our knowledge of its taxo- nomic diversity. Most species currently assigned to Tropidurus began...

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... The Caatinga is a highly diverse region and an area of endemism for the genus Tropidurus, harbouring 13 of the 28 species described (Carvalho et al., 2013;Carvalho et al., 2016;Uetz et al., 2020). Tropidurus species use several habitat types (e.g., sandy soils, rocky outcrops, semi-arboreal, and open areas) and can have from very narrow (e.g., paleodunes from the São Francisco River, rock fields from Espinhaço Mountain Range) to broad geographic distributions (Mesquita et al., 2017;Rodrigues, 2005). ...
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Organisms adapted to open environments in South America have recently been used to understand the origins of the high Neotropical biodiversity. In the Caatinga, the largest continuous block of Seasonally Dry Tropical Forest in South America, phylogeographic studies have uncovered the role of historical climate changes and rivers (i.e., the São Francisco River, the largest perennial river in Caatinga), in promoting genetic differentiation and speciation of lizards and amphibians. We used mitochondrial data, demographic analyses, paleodistribution models, and landscape genetic methods to test the effects of spatial distances, historical climate fluctuations, and landscape heterogeneity on the genetic variation of the generalist lizard Tropidurus hispidus in the semi-arid Caatinga in northeastern Brazil. Four haplogroups with moderate geographical structure diverged in the Pleistocene, and exhibited high genetic diversity. Ecological niche models revealed large suitable climatic areas for T. hispidus in the past 790 thousand years, connecting the Caatinga and other regions via a narrow corridor. Part of the genetic differentiation in T. hispidus resulted from spatial distances among populations and isolation by resistance through climatic unsuitability areas in the Last Glacial Maximum (LGM), which probably reduced population connectivity and gene flow. Our findings highlight the role of the historical factors of the Caatinga, through LGM climate, and the generalist condition of species in shaping the genealogical histories of populations. Although the results are based on a single-locus approach, our study is a first step to shed light on the main drivers of the evolutionary history of T. hispidus, in a highly diverse and still poorly studied region. © The Trustees of the Natural History Museum, London 2022. All Rights Reserved.
... This analysis was also implemented in the CIPRES Science Gateway, version 3.3 (Miller et al., 2012). We used Microlophus quadrivittatus (Tschudi, 1845) to root the tree, with Plica plica (Linnaeus, 1758) and Tropidurus oreadicus Rodrigues, 1987 as additional outgroups, based on the most recent phylogenetic hypothesis for Tropiduridae, proposed by Carvalho et al. (2016). The outgroup sequences were obtained from data available on the GenBank database (Benson et al., 2016). ...
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Wetlands are present in different parts of the globe, holding a significant portion of the local biodiversity, and have been under the constant influence of climatic changes worldwide on different time scales. Investigations with a phylogeographic approach have revealed the role of Quaternary climatic shifts on a global scale, favoring constant changes in the amplitude of floodplains and affecting the evolutionary history of multi-taxa. This pattern was observed in the Amazonian biota, but due to the complex geological history, some of these models alone are insufficient, especially in widespread species that live in specific regions of the biome. Herein, we investigated the species delimitation and biogeographic history of the semi-aquatic lizard Uranoscodon superciliosus (U. superciliosus), widely distributed in Amazonian floodplains, a habitat that was for a long time considered a corridor to gene flow. Our results support a high genetic diversity with five well-supported lineages within U. superciliosus: North of the Amazon River, East Xingu, Solimões, Purus, and Tapajós-Madeira, with the basal split within this group in the Miocene and others in the Plio-Pleistocene. These results corroborated a mixture of distinct processes that shaped the diversity of U. superciliosus with rivers as vicariant barriers and the already known role of paleoclimatic shifts during the Quaternary promoting diversification. Among tetrapods, our work is one of the first to describe the genetic structure in a widespread taxon along river-edge environments, thus supporting both patterns of diversification with the compartmentalized lineages along different Amazonian rivers (floodplains) and also the more common pattern found in upland forest species, with main rivers acting as vicariant barriers and playing a role in allopatric speciation.
... Terrestrial and diurnal. It has being observed among rocks on sandy trails covered by dry forests and caatinga vegetation (Carvalho et al. 2016). Diet based mainly on arthropods, no information about the preferred items on the diet of this species is known. ...
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The Caatinga is the largest seasonal dry tropical forest in South America and it has been historically neglected in terms of its bio- diversity. Regarding lizards, different studies led to the current knowledge of diversity and endemism in Caatinga, but detailed syntheses are scarce in the literature. We present the most detailed and up-to-date synthesis of knowledge about Caatinga lizards by providing a detailed (i) list of species; (ii) taxonomic richness patterns; (iii) knowledge gaps and spatial biases; and (iv) detailed distribution maps of all species that contain at least one occurrence record within Caatinga limits. We created a distribution database using occurrences of lizards in Caatinga based on scientific collections, field collection, and literature. We produce up-to-date dis- tribution maps, calculate the Extent of Occurrence and provide the environmental and bioclimatic profile for each species recorded. We draw taxonomic richness and sampling gap maps. Our database has 20,538 records of occurrence of lizards of the Caatinga. We recorded 93 lizard species (13 families), 52.7% of which are endemic. Forthy-four percent of the species present restricted distri- butions. We identified that 53% of the Caatinga area (or 70% of the municipalities) has no record of occurrence of lizards. The data presented are an important step towards synthesizing in detail the accumulated knowledge about Caatinga lizards and is crucial for accurate strategies for the conservation planning. It directs actions to advance our knowledge on Caatinga lizards: to concentrate inventories in sample void areas; continuous update of the species occurrence database, advance in the generation of autoecology data for species.
... Furthermore, some amphibian and reptile species have been recently described for this biome (e.g. Andrade et al., 2016;Carvalho et al., 2016;Franco et al., 2017), which reinforces that there is still a lack of checklists for some areas from the Caatinga. ...
... We measured ten adult males and ten adult females for each of the following species ( Tropiduridae species exhibit conspicuous external sexually dimorphic traits, which include the presence of ventral black spots in the hindlimbs and near the cloaca only in males (see Carvalho et al., 2016); we measured only adults and used these traits for sex differentiation. For each species, we assured that all individuals measured were collected at the same locality and during the same field campaign. ...
... We then reconstructed ancestral states of the occurrence, pattern of sexual dimorphism and the identity of the dimorphic digit using Maximum Likelihood; the evolutionary model used corresponded to 'equal rates model' [ER], implemented with the Ape package (see Paradis et al., 2004) and using the aforementioned coding system. Topology was constructed in Mesquite v.3.61, based on phylogenetic hypotheses from Pyron et al. (2013) and Carvalho et al. (2016). ...
... Following Lofeu et al. (2017), and given that body size and some other body traits are very sexually dimorphic in the Tropiduridae (see Kohlsdorf et al., 2006;Gomes & Kohlsdorf, 2011;Brandt & Navas, 2013;Brandt et al., 2016), we used SDI to evaluate phenotypic integration between sexual dimorphism in body traits and variation in digit lengths. Specifically, we used Phylogenetic Generalized Least Squares [PGLS implemented with the phytools package (Revell, 2012); topology modified from the phylogenetic hypotheses of Pyron et al. (2013) and Carvalho et al. (2016)] to test for correlations between SDI in body traits and SDI in the digits that were sexually dimorphic at least in one of the species studied here. ...
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Sexual dimorphism in digit lengths reflects phenotypic evolution mediated by developmental steroids. Differences in the identity of the sexually-dimorphic digit may evolve if the concentrations of sex-steroid receptors in the digit are easily modified and the initial changes have low impact on fitness. Accordingly, sexual dimorphism in digit lengths might initially originate under neutral selective regimes, being subsequently co-opted by embryonic hormonal effects on sensitive traits that are more likely to be targeted by selection. Correlated variation among sexually-dimorphic traits might therefore reflect pleiotropic hormonal modulation during development. Moreover, the identity and trend of the sexually-dimorphic digit might be evolutionarily labile even among closely-related species. We evaluated this model by assessing the identity of sexually-dimorphic digits among 11 Tropiduridae lizard species and inferring evolutionary patterns of sexual dimorphism. Assuming that digit lengths can be used as a proxy for hormonal modulation of steroid-sensitive traits during development, we tested for evolutionary associations among sexual dimorphism of digit lengths, body size and other traits that may be direct targets of sexual selection in the Tropiduridae. Sexual dimorphism in digit lengths is evolutionarily labile in the Tropiduridae, and diversity, instead of conservatism, seems to rule developmental mechanisms underlying the evolution of sexual dimorphism in digit ratios. ADDITIONAL KEYWORDS: developmental steroids-digit length-sexual dimorphism-Squamata-Tropiduridae.
... In recent years, new amphibians and reptiles species were described in Brazil (e.g. Fernandes and Hamdan, 2014;Lourenço et al., 2014;Magalhães et al., 2014;Pansonato et al., 2014;Colli et al., 2015;Maciel et al., 2015;Carvalho et al., 2016;Franco et al., 2017). However, without carrying out exhaustive checklists and fieldworks, many Brazilian species could become extinct before scientists even have a chance to identify them. ...
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Recent studies on Brazilian coastal zones and restinga environments revealed a high richness of amphibian and reptile species. However, there is still a lack of information about herpetofauna diversity in coastal zones of Northeastern Brazil. This study provides a checklist of amphibians and reptiles inhabiting the Environmental Protection Area (EPA) Delta do Parnaíba, Northeastern Brazil, suggesting conservation actions. To elaborate the checklist, we searched in seven electronic databases and check the following scientific collections: Zoological collection of Universidade Federal do Piauí (UFPI) and Herpetological collection of Universidade Regional do Cariri (URCA). In addition, we sampled 16 areas along the EPA Delta do Parnaíba close to the river branches and temporary ponds that compose the Parnaíba River Delta (December 2015 to April 2017) to fill gaps of information about herpetofauna in some regions from the EPA. We recorded 86 species (34 amphibians and 52 reptiles), including four anurans, one crocodilian, 14 snakes, 12 lizards and two amphisbaenians reported for the first time for the EPA Delta do Parnaíba. In addition, we added the first record of the snake Oxybelis fulgidus in Piauí state. The EPA Delta do Parnaíba shows high herpetofaunal richness; thus, we suggest that conservation actions should be taken to preserve the restingas environments in the Parnaíba River Delta and its high diversity of amphibians and reptiles.
... Molecular phylogenies are consistent with this hypothesis, showing that the exclusively psammophilous genera Calyptommatus, Nothobachia, Procellosaurinus and Psilops are part of a diverse gymnophthalmid radiation (Pellegrino et al. 2001) that includes other open-habitat genera such as Gymnophthalmus, Micrablepharus and Vanzosaura, which already show striking adaptations for a fossorial lifestyle such as the absence of eyelids, small body-size, and limb reduction (Rodrigues 1991c). The Tropidurus torquatus group, on the other hand, is a monophyletic group (Frost et al. 2001;Carvalho et al. 2016) in which several species are sandy-habitat specialists and show morphological adaptations associated with locomotion on sand (Rodrigues 1987;Kohlsdorf et al. 2004;Grizante et al. 2010). Relatives of the endemic species T. psammonastes and the sand-dwelling T. cocorobensis occur in distinct types of open habitats in Central and Northeastern Brazil (Carvalho et al. 2016). ...
... The Tropidurus torquatus group, on the other hand, is a monophyletic group (Frost et al. 2001;Carvalho et al. 2016) in which several species are sandy-habitat specialists and show morphological adaptations associated with locomotion on sand (Rodrigues 1987;Kohlsdorf et al. 2004;Grizante et al. 2010). Relatives of the endemic species T. psammonastes and the sand-dwelling T. cocorobensis occur in distinct types of open habitats in Central and Northeastern Brazil (Carvalho et al. 2016). Whiptail lizards of the Ameivula ocellifera group and the genus Glaucomastix (Goicoechea et al. 2016) are present in the dry diagonal of open habitats of South America and coastal plains, and are also known to occur frequently or exclusively on sandy-soil in open habitats (Dias and Rocha 2007;Colli et al. 2009;Arias et al. 2011). ...
... For comparative purposes, we retrieved mtDNA and nuDNA sequences from Genbank for Calyptommatus and related Gymnophthalmini (Goicoechea et al. 2016), Eurolophosaurus and Tropidurinae (Passoni et al. 2008;Carvalho et al. 2016) and Pseudoboini (Grazziotin et al. 2012;Pyron et al. 2013), and added molecular data from the sand-dwelling snake genus Rodriguesophis (Rodrigues 1993) and Eurolophosaurus (Appendix 2). We applied a Bayesian approach to concurrently infer phylogenetic relationships and divergence times, using molecular clock rates previously estimated for Neotropical snakes and lizards (Daza et al. 2009;Olave et al. 2015). ...
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In the middle course of the São Francisco River (SFR), inserted in the semiarid domain of the Caatinga, there is an extensive field of palaeodunes which represents an important testimony of former, drier climates in northeastern Brazil. The highly diverse sand-dwelling fauna of this dune region comprises several endemic reptiles with geographically structured distributions. Based on geomorphological, paleoclimatic, and distribution data of pairs of psammophilous reptiles, a paleolacustrine vicariant diversification scenario was proposed to explain the origin of this endemic fauna. In this model, after an endorheic phase during the last glacial maximum, when the SFR flowed into an interior lake, the SFR made its way out to the ocean through the dunes, separating ancestral populations of several groups of lizards and snakes on opposite margins. Phylogenetic data from endemic species are only partially concordant with this model, and highlight the contribution of both local and regional events as drivers of diversification. Estimates of divergence times suggest that speciation was not simultaneous, and that some events occurred prior to Pleistocene. Species that are widespread in the SFR dune region lack genetic structure between the two margins, contrasting with the pattern observed for endemic, habitat-specialist lineages. Both Pliocene and Pleistocene events appear to have promoted diversification in SFR dune endemic reptiles, and ecological interactions may have contributed to species turnover. The SFR may have acted both as a vicariant barrier and as a boundary delimiting secondary contact, reinforcing ongoing processes of speciation.
... A marked trait of this genus (and other genera) of this family is the presence of dark-coloured patches (Figure 1d,f) at ventral thighs and pre-cloacal flaps of adult males (Bruinjé, Coelho, Paiva, & Costa, 2019;Bruinjé, Moura, et al., 2019;Frost et al., 2001;Stebbins, Lowenstein, & Cohen, 1967). A primary premise of its function is that its colouration would play a role in visual communication (Pinto, Wiederhecker, & Colli, 2005), although it was also suggested that coloured patches could be just a consequence of chemical secretions produced by glandular scales (Carvalho et al., 2016;Harvey & Gutberlet, 1998;Kuns & Borges-Martins, 2013). However, the function of these ventral patches, including roles in visual or chemical signalling, remains largely unexplored. ...
Article
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.
... Even when in the middle of a sugarcane plantation, over 3 km far from the freeway, I could see lagartixas only in proximity to artificial structures (e.g., concrete wall of small dams). Carvalho et al. (2016) noted that the occurrence of a population of T. hispidus, in one southern area of its distribution, apparently was opportunistic and associated with man-made structures. This raises the possibility that lagartixas might profit from the anthropogenic built landscape by experiencing not only substantial population growth, but also range expansion. ...
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Abstract Urbanization, with its cohort of environmental stressors, has a dramatic effect on wildlife, causing loss of biodiversity and decline in population abundance customarily associated with increasing levels of impervious surface and fragmentation of native habitats. Some studies suggest that faunal species from open habitats, and with higher abundance in natural environments, seem more likely to tolerate and live in urban environments. Here I evaluate how the level of urbanization affects lagartixas (Tropidurus hispidus) one of the most common lizards found in open vegetation ecosystems in NE Brazil. I surveyed a total of 47 transects across sites that differed in proportion of impervious surface (high, mild, peri‐urban, and rural). I also collected specific biotic (herbaceous cover, tree, and arthropod abundance) and abiotic (amount of shelters and impervious surfaces) factors that could affect lagartixas abundance. Ants were the most common arthropod both in the rural and urban environment. Lagartixas thrive in urban environments, and trees and shelter were key predictors of their abundance. Lagartixas show a clear association with use of artificial structures. The low densities of lagartixas in rural areas and higher density in urbanized areas suggest that they colonized urban areas due to the hard surfaces and they probably are not exploiting a novel habitat, but somewhat responding to conditions resembling those in which they evolved. Finally, lagartixas are extremely common in tropical cities, they have a suite of features that are associated with selective pressures in cities and they might play a key functional role in urban ecosystems making this lizard an excellent system for the study of ecology and adaptation to the urban environments.
... Future steps would be to explore the physiological mechanisms responsible for the production of color of the chest and ventral patch areas. Coloration of ventral patches in Tropidurus species is supposed to be linked to the occurrence of glandular scales (Carvalho et al. 2016), which was not studied yet. Hormone levels might be responsible for mediating high melanin in the tissue. ...
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Behavior can help to establish dominance in intrasexual interactions, preventing more costly aggressive interactions and improving access to mates. Distinct color morphs often correlate with behavior, driving differential reproductive success between them. The lizard Tropidurus semitaeniatus usually expresses two male color morphs, Yellow or Black. It is likely that morphs play a role in reproduction, which is still unexplored. Here, we test whether there is morph-related dominance during intrasexual interactions. We used ex situ behavioral trials to test whether a particular morph shows dominance, gathering dominance by attributing scores to aggressive/evasive behaviors. We also tested whether winner individuals show higher performance (sprint speed), and whether spectrophotometric measures of body coloration predict winners of male encounters. Morphs showed differences in behaviors suggesting alternative behavioral tactics: Black males behave more aggressively and less evasively while Yellow males show the opposite sets of behavior. Black males also tend to be dominant, but dominants do not show higher sprint speeds than submissive males. Chest coloration, often displayed during encounters, highly predicts winnings (particularly high yellow chroma and low lightness and UV). Our results show that lizards signal competitive condition by behaviorally exposing their chest. Males displaying more head bobs and with darker chests are more likely to win encounters. Our results suggest that Yellow males might undertake a sneaker tactic, preventing aggression costs by evasiveness even though they might perform similarly to Black males. Further studies should address whether female preference is biased in relation to male morphs and its colorations, which would suggest selective processes towards costly signals and morph maintenance. Significance statement In the struggle for survival and reproduction, often, there is no single best strategy and multiple distinct tactics may evolve. Behavior, color signaling, and performance are frequently correlated with distinct color morphs, which can coexist as alternative mating tactics. However, studies that are able to integrate all these traits are scarce. Here, we test whether different color morphs of the lizard Tropidurus semitaeniatus show different behaviors and dominance patterns. We also test whether these color morphs differ in their performances (sprint speed) and visual signaling (behavioral displays and intensity of coloration). We demonstrate that Black and Yellow-morph males adopt distinct behavioral tactics: an aggressive (Black-morph) and an evasive (Yellow-morph) tactic. We also show that dominance is highly correlated to chest’s intensity of dark yellow colorations and that lizards signal their dominance status through displays of head bob bouts.