Introduction

Lizards are key organisms for studies in numerous fields in biology, such as evolution, behaviour, population, community ecology, and life-history theory (Mesquita et al., 2016), and have even contributed to medicine advances (Mahdavi et al., 2008). This successful group of reptiles inhabits virtually all places on Earth, except the highest and coldest ones (Pianka and Vitt, 2003). Over 6,000 lizard species have already been described, a number that represents more than half of all living reptiles known to date (Uetz and Hošek, 2016). In Brazil, more than 270 species are known to occur (Costa and Bérnils, 2018). However, there are still many taxonomic problems to be solved and knowledge gaps to be fulfilled, mainly in North and Northeast regions of the country.

The state of Paraíba is situated on the easternmost portion of the South American continent and much of its biodiversity is still unknown. Rodrigues (1986; 2000; 2003) demonstrated that samplings are still deficient in the region. The state comprises vegetation physiognomies that vary from coastal rainforests (Atlantic Forest domain) to semiarid and arid regions in the seasonally dry forests (Caatinga domain), including Caatinga enclaves’ moist forests, known as “brejos de altitude”, which are islands of moist forest surrounded by dry forests. Most of the original Atlantic Forest domain coverage has been lost due to anthropic occupation, and the Caatinga still suffers with the transformation of its natural landscapes into crops and pastures (Leal et al., 2005; Tabarelli et al., 2005).

Another worrying issue in this state is the extent of its protected areas. Less than 10% of its territory is legally protected, being only approximately 1.2% of the original Caatinga formation under some degree of legal protection (MMA, 2011). In the Atlantic Forest, this number is higher (6%) (MMA, 2011), but it is still under the goal of 17% of legally protected areas until 2020, defined in the Aichi Biodiversity Targets, a short term international plan to save biodiversity (Prates and Irving, 2015).

Herpetology Notes, volume 12: 749-763 (2019) (published online on 09 July 2019)

Lizard fauna from the state of Paraíba, northeastern Brazil: current knowledge and sampling discontinuities

Lissa Dellefrate Franzini1,*, Izabel Regina Soares da Silva1, Daniel Oliveira Santana1, Fagner Ribeiro Delfim, Gustavo Henrique Calazans Vieira1, and Daniel Oliveira Mesquita1

1 Programa de Pós-Graduação em Ciências Biológicas (Zoologia), Departamento Sistemática e Ecologia, Universidade Federal da Paraíba, João Pessoa, PB, Brazil.

* Corresponding author: lissa.dfranzini@gmail.com

Abstract. We compiled a list of lizard species from the state of Paraíba, Northeast Brazil to systematize and widen the knowledge about the lizard fauna in that state, which is still poorly known in most of its municipalities and has never been systematized in a species list. We considered data from the literature and the major scientific collections that house specimens from the state of Paraíba and we gathered a total of 2,767 records for 48 municipalities, corresponding to 36 lizard species of 27 genera and 12 families. The capital, municipality of João Pessoa, presented the highest number of species records. This number of records is a result of a high sampling effort, which is probably a consequence of the higher number of reptile specialists working in this municipality, which have for long contributed to the scientific knowledge on the region. Most of the records come from municipalities with protected areas near research centres, which are usually more accessible. In the remaining municipalities, sampling is still deficient, mainly in regions of the Caatinga domain. We calculated the evolutionary distinctiveness (ED) of the lizard species compiled in the study and verified that Diploglossus lessonae, Coleodactylus meridionalis, Lygodactylus kuglei, Iguana iguana, and Salvator merianae represent important evolutionary histories in the biodiversity of the state.

Keywords. Squamata, Caatinga, Atlantic Forest, species list

Lissa Dellefrate Franzini et al.750

The knowledge about the species that occur in a state is critical to their conservation. Regarding reptile and amphibian fauna from Paraíba, the few published species lists focused on small regions and conservation units (e.g., Arzabe et al., 2005; Santana et al., 2008). In this sense, we aim to systematize data concerning lizard species from the state of Paraíba. We consider data from the literature and the major scientific collections that house specimens from that state. Additionally, we provide a brief discussion about their conservation status and identify areas of poor sampling to highlight the need of broadening the knowledge of lizards from the region.

Material and Methods

In the present list, we compiled data from scientific collections and literature. We obtained data from deposited specimens on Coleção Herpetológica da Universidade Federal da Paraíba (CHUFPB), Coleção Herpetológica da Universidade de Brasília (CHUNB) and Museu de Zoologia da Universidade de São Paulo (MZUSP). Specimens from CHUFPB and MZUSP were analysed by FRD in order to avoid misidentification. In CHUNB, the identification of the specimens was performed by the collection curator, Dr. Guarino Rinaldi Colli, following the same guidelines. Complementary data was obtained from the online database Specieslink (2017). In this database, we obtained records from Museum of Comparative Zoology – Harvard University (MCZ) and Coleção de Répteis do Museu de Zoologia da Unicamp (ZUEC). Because we were not able to check the identifications obtained from Specieslink, we included only those species that, by the experience of the authors, could occur in the listed localities. A voucher per municipality per institution and the number of specimens deposited is provided by species in the Appendix.

We carried out the search on scientific literature using the terms: “lizards + Paraíba”; “herpetofauna + Paraíba”; “reptiles + Paraíba” in Google Scholar and Web of Science databases. Using the same criteria, we carried out a similar search to obtain data concerning the lizard fauna from the states of Alagoas, Ceará, Pernambuco, and Rio Grande do Norte. We consulted data from those states to compare the lizard fauna from Paraíba with nearby localities. We excluded from the compilation all articles that mentioned only records deposited in CHUFPB to avoid data redundancy, data from the other collections analysed in this study were not found in the literature and therefore we did not exclude

articles from any other collection from the analysis. Even though Serpentes is a clade placed among lizards, we decided not to include it in the present study due to particularities regarding snake biology (high diversity, low abundance). Also, a detailed study of the snakes of Paraíba is already available (Filho et al., 2017).

Taxonomic nomenclature and classification followed the Brazilian List of Reptile Species (Costa and Bérnils, 2018). We included the introduced species Hemidactylus mabouia (Moreau de Jonnès, 1818) in the analysis because it is well-established in Brazil and it is listed in the Brazilian List of Reptile Species (Costa and Bérnils, 2018). To evaluate the species’ conservation status, we consulted both the International Union for Conservation of Nature (IUCN, 2017) and the Brazilian red list of threatened species (MMA, 2014).

Even though more specific site information or exact coordinates were available for part of the data, all localities were listed as municipalities, since this was the sole information available for most records. We built a rarefaction curve based on individuals vs. species to evaluate the expected richness on our sampling. For this, we used the software EstimateS 9.1.0 (Colwell et al., 2012) with 1,000 randomizations, without reposition, and Chao1P as estimator of species richness (Lopez et al., 2012). We used an individual-based curve because our sampling design was also individual based, and because the individuals are the entities that carry taxonomic information (Gotelli and Colwell, 2001).

Another rarefaction curve was built considering only data deposited in CHUFPB, where each decade was considered as a sample. This analysis was performed in the same way as described for the individuals vs. species curve but with a Chao2P estimator, since it had a lower standard deviation. We evaluated only the species deposited over time for the CHUFPB to better understand the information provided by this collection, which concentrates most of the data about the lizard species from the state of Paraíba.

For phylogenetic comparisons, we used the hypothesis provided by Zheng and Wiens (2016), which is a time-calibrated phylogeny for Squamata based on 52 genes sampled for 4,162 species. We pruned their topology to obtain a tree where only the state of Paraíba’s lizards were retained, keeping the time of diversification in the pruned phylogeny and maintaining Sphenodon punctatus Gray, 1842 for comparative reasons. However, since some species from the state of Paraíba are not represented on that hypothesis, we replaced those species by the most related species in Zheng and Wiens’ tree. The replacements were: Diploglossus bilobatus

(O’Shaughnessy, 1874) by D. lessonae Peracca, 1890, Colobosauroides cearensis Cunha, Lima-Verde and Lima, 1991 by Dryadosaura nordestina Rodrigues, Xavier Freire, Pellegrino and Sites, 2005, Enyalius bilineatus Duméril and Bibron, 1837 by E. bibronii Boulenger, 1885, Enyalius leechii Boulenger, 1885 by E. catenatus Boulenger, 1885, Phyllodactylus xanti Cope, 1863 by Gymnodactylus darwinii Gray, 1845, Phyllodactylus unctus Cope, 1864 by Gymnodactylus geckoides Spix, 1825, and Tropidurus hygomi Reinhardt and Lütken, 1862 by Tropidurus semitaeniatus Spix, 1825. We recognize that such substitutions are not ideal, yet it is the only approach we could have taken, since virtually all the phylogenetic hypotheses provided for Squamata so far are not based on a complete (or nearly complete) sampling of lizard species. Moreover, most evolutionary changes in the tree of life of Squamata are nested in the deepest nodes (Mesquita et al., 2016), which potentially makes the replacement less harmful considering our purpose. The resulting tree was then used to calculate Evolutionary Distinctiveness (ED) statistic (Isaac et al., 2007) for each lizard species, which measures each species contributions to phylogenetic diversity, and is given in units of time before the present (in our case, millions of years – MY). The ED metric also permits us to discuss the putative taxonomic sampling gaps and phylogenetic uniqueness associated to the lizards within the boundaries of the state. By using only ED as a surrogate of the “originality” of species (Isaac et al., 2007), we can assume that, for the state of Paraíba, species with the highest ED values should be prioritized in conservation measures. The interpretation is pretty straightforward: if species with high ED values are lost within the state boundaries unique evolutionary histories will be lost in the phylogenetic diversity of the state. To obtain ED values we used the package caper (Orme et al., 2013) in R environment, software v 3.4.3 (R Core Team, 2017). Additional packages used to examine, prune, and plot phylogeny and associated data (ED values) were ape (Paradis et al., 2004), phyloseq (McMurdie and Holmes, 2013), and phytools (Revell, 2012). When pertinent, ED values are showed by the arithmetic mean of species ED values ± 1 standard deviation.

Unfortunately, because some species show no variation on their status of IUCN Red List (four species) or lack such status (Table 1), we could not calculate species EDGE values, a statistic that satisfy two requirements of conservation priority-setting approaches – the capture of biodiversity and robustness to uncertainty (Isaac et al., 2007). Thus, we also could not compare

ED values among the different species IUCN Red List status/categories and, since many species occur in both domains, we could not test for differences in ED values between Atlantic Forest and Caatinga, as Tonini et al. (2016) did.

Results

We gathered a total of 2,767 occurrence records, corresponding to 36 lizard species allocated in 27 genera of 12 families (Table 1; Fig.1; Fig. 2). Gymnophtalmidae was the richest family represented, with seven species from seven different genera occurring in the state. The families Anguidae and Sphaerodactylidae were represented by a single species each, Diploglossus lessonae and Coleodactylus meridionalis Boulenger, 1888, respectively. Dactyloidae was the only family that occurred exclusively in the Atlantic Forest, while other families were registered in both domains (Caatinga and Atlantic Forest). Nine species (26% of the species registered) occurred only in the Caatinga and twelve species (34%) occurred only in the Atlantic Forest (Table 1). For all the species listed here but four, the threat status according to IUCN is still not available (not classified yet). The remaining four species are assigned with the ‘least concern’ status (Table 1).

Ameivula ocellifera Spix, 1825 was the species present in more municipalities (27), followed by Tropidurus hispidus Spix, 1825, being recorded in 26 localities (Table 1). On the other hand, some species showed a narrow distribution on the state, being recorded in only two municipalities: Strobilurus torquatus Wiegmann, 1834, recorded in municipalities of João Pessoa and Santa Rita, in the Atlantic Forest, E. catenatus, recorded in municipalities of Mamanguape and Mataraca, also in the Atlantic Forest, and Hemidactylus brasilianus Amaral, 1935, recorded in municipalities of Cabaceiras and São José dos Cordeiros, in the Caatinga. We noticed that the species Copeoglossum nigropunctatum Spix, 1825 (formerly treated as Mabuya nigropunctatum) had been previously recorded in the state, but it was misidentified as Mabuya bistriata Spix, 1825 by Santana et al. (2008).

It is also worth mentioning the record of Stenolepis ridleyi Boulenger, 1887 for the state, which occurred exclusively in the municipality of Matureia, an enclave of moist forest within the Caatinga. And we point out that Coleodactylus meridionalis was recorded for Caatinga, but we point out that the species does not occur in Caatinga stricto sensu, but in more humid environments.

Lizard fauna from the state of Paraíba, northeastern Brazil 751

Lissa Dellefrate Franzini et al.752

Table 1. Lizard species from the state of Paraíba, Brazil and its distribution within the state. CA – Caatinga; AF – Atlantic Forest; BA – “Brejo de altitude”. I – Arzabe et al. (2005); II – Freire (1996); III – IBAMA (2015); IV – Magalhães-Júnior et al. (2014); V – Mendonça et al. (2014); VI – Ragner et al. (2014); VII – Queiroz et al. (2010); VIII – Vieira et al. (2012). CHUFPB – Coleção Herpetológica da Universidade Federal da Paraíba, CHUNB = Coleção Herpetológica da Universidade de Brasília, MCZ = Museum of Comparative Zoology – Harvard University, ZUEC = Coleção de Répteis do Museu de Zoologia da Unicamp; MZUSP = Museu de Zoologia da Universidade de São Paulo, Species topotypic from Paraíba are shown in bold, while *and **indicates IUCN Red List assessment as “Least Concern” and “not assessed yet”, respectively.

Taxon Municipalities Distribution Source

Anguidae

Diploglossus lessonae Peracca, 1890* Araruna, Bananeiras, Cabaceiras, Campina Grande, Gurinhém, Patos

CA; AF CHUFPB; MZUSP; I; VII

Dactyloidae

Dactyloa punctata (Daudin, 1802)** João Pessoa, Mamanguape, Rio Tinto AF CHUFPB; MCZ; MZUSP Norops fuscoauratus (D’Orbigny, 1837 in

Duméril e Bibron, 1837)** Caaporã, Cabedelo, João Pessoa AF CHUFPB; II

Norops ortonii (Cope, 1868)** Cabedelo, João Pessoa, Rio Tinto, Sapé AF CHUFPB; MZUSP; II

Gekkonidae

Hemidactylus agrius Vanzolini, 1978** Araruna, Cabaceiras, Cacimba de Dentro,Patos CA CHUFPB; CHUNB; MZUSP; I Hemidactylus brasilianus (Amaral, 1935)** Cabaceiras, São José dos Cordeiros CA CHUFPB; III

Hemidactylus mabouia (Moreau de Jonnès, 1818)**

Araruna, Areia, Caaporã, Cabaceiras, Cabedelo, Cacimba de Dentro, Campina Grande, Gurinhém, João Pessoa, Junco do Seridó, Mamanguape, Rio Tinto, São João do Cariri

CA; AF; BA CHUFPB; CHUNB; MZUSP; I; II

Lygodactylus klugei (Smith, Martin e Swain, 1977)**

Araruna, Cabaceiras, Cacimba de Dentro, Pombal, São João do Cariri, São José de Espinharas, São José dos Cordeiros

CA CHUFPB; CHUNB; MZUSP; I; III

Gymnophthalmidae

Acratosaura mentalis (Amaral, 1933)** Araruna, Cabaceiras, Cacimba de Dentro, São José dos Cordeiros

CA CHUFPB; MZUSP; I; III

Anotosaura vanzolinia Dixon, 1974** Cabaceiras, Campina Grande, São José da Mata, São José dos Cordeiros

CA CHUFPB; MZUSP; III; VII

Cercosaura ocellata Wagler, 1830** Alhandra, Conde, João Pessoa, Mamanguape, Rio Tinto

AF CHUFPB; CHUNB; MZUSP

Dryadosaura nordestina Rodrigues, Freire, Pellegrino e Sites, 2005**

Caaporã, Cabedelo, João Pessoa, Mamanguape, Rio Tinto

CA; AF CHUFPB; CHUNB; MZUSP

Micrablepharus maximiliani (Reinhardt e Luetken, 1862)**

Alhandra, Cabedelo, Cacimba de Dentro, Gurinhém, João Pessoa, Mamanguape, Mataraca, Rio Tinto

CA; AF CHUFPB; MZUSP; I; II

Stenolepis ridleyi Boulenger, 1887** Matureia BA MZUSP

Vanzosaura multiscutata (Amaral, 1933)** Cabaceiras, Cacimba de Dentro, Cajazeiras, Juazeirinho, Piancó, São João do Cariri, São José dos Cordeiros, Soledade, Umbuzeiro

CA CHUFPB; MZUSP; I; III

Iguanidae Taxon Municipalities Distribution Source

Iguana iguana (Linnaeus, 1758)** Araruna, Bananeiras, Caaporã, Cabaceiras, Campina Grande, Coremas, Gurinhém, Itabaiana, João Pessoa, Juazeirinho, Junco do Seridó, Mamanguape, Patos, Pocinhos, Rio Tinto, São João do Bonfim, São João do Cariri, São José dos Cordeiros

CA; AF; BA CHUFPB; MZUSP; ZUEC; I; III; V; VII

Leiosauridae

Enyalius bibronii Boulenger, 1885* Araruna, Areia, Gurinhém, São José dos Cordeiros, Sapé, Teixeira

CA; AF; BA CHUFPB; MZUSP; III

Enyalius catenatus (Wied, 1821)** Mamanguape, Mataraca AF CHUFPB

Mabuyidae

Brasiliscincus heathi (Schmidt e Inger, 1951)**

Alhandra, Arara, Araruna, Areia, Bananeiras, Cabedelo, João Pessoa, Mamanguape, Patos, Sapé, São José dos Cordeiros, Zabelê

CA; AF CHUFPB; CHUNB; MZUSP; I; III

Copeoglossum nigropunctatum (Spix, 1825)**

Cruz do Espírito Santo, João Pessoa, Mamanguape, Rio Tinto, Sapé

AF CHUFPB; MZUSP

Psychosaura agmosticha (Rodrigues, 2000)**

Cabaceiras, Cacimba de Dentro, Coremas, São José dos Cordeiros

CA CHUFPB; I; III; IV

Table 1. Continued.

Lizard fauna from the state of Paraíba, northeastern Brazil 753

Taxon Municipalities Distribution Source

Psychosaura macrorhyncha (Hoge, 1947)** Alhandra, Cabaceiras, Cabedelo, Mamanguape, São José dos Cordeiros, Sapé, Rio Tinto

CA; AF CHUFPB; MZUSP; II

Phyllodactylidae

Gymnodactylus darwinii (Gray, 1845)** Caaporã, Cabedelo, João Pessoa, Mamanguape AF CHUFPB; CHUNB; II

Gymnodactylus geckoides Spix, 1825** Araruna, Cabaceiras, Cacimba de Dentro, Campina Grande, João Pessoa, Mamanguape, Patos, Piancó, São João do Cariri, São José de Espinharas, São José dos Cordeiros

CA; AF CHUFPB; MCZ; MZUSP; ZUEC; I; III; VII

Phyllopezus periosus Rodrigues, 1986** Araruna, Cabaceiras, Cacimba de Dentro, Campina Grande, Itaporanga, Salgadinho, São João do Cariri, São José dos Cordeiros

CA CHUFPB; CHUNB; MCZ; MZUSP; I; III; VI

Phyllopezus pollicaris (Spix, 1825)** Araruna, Areia, Boa Vista, Cabaceiras, Cacimba de Dentro, Itaporanga, Junco do Seridó, Patos, Piancó, São João do Cariri, São José de Espinharas, São José dos Cordeiros

CA; BA CHUFPB; CHUNB; MZUSP; I

Polychrotidae

Polychrus acutirostris Spix, 1825** Areia, Cabaceiras, Gurinhém, João Pessoa, Mamanguape, Mataraca, São João do Cariri, São José dos Cordeiros, São José de Espinharas

CA; AF CHUFPB; MZUSP; III

Polychrus marmoratus (Linnaeus, 1758)** Areia, João Pessoa, Mamanguape, Rio Tinto AF; BA CHUFPB; MZUSP

Sphaerodactylidae

Coleodactylus meridionalis (Boulenger, 1888)**

Alhandra, Areia, Araruna, Caaporã, Cabedelo, Cruz do Espírito Santo, Jacaraú, João Pessoa, Mamanguape, Matureia, Rio Tinto

CA; AF; BA CHUFPB; CHUNB; MCZ; MZUSP; I; II

Teiidae

Ameiva ameiva (Linnaeus, 1758)** Alhandra, Araruna, Areia, Cabaceiras, Cabedelo, Gurinhém, João Pessoa, Junco do Seridó, Mamanguape, Mogeiro, Rio Tinto, São José dos Cordeiros, São José de Espinharas, Sapé

CA; AF; BA CHUFPB; CHUNB; MZUSP; I; III; II

Ameivula ocellifera (Spix, 1825)** Alhandra, Araruna, Areia, Cabaceiras, Cabedelo, Cacimba de Dentro, Coremas, Cruz do Espírito Santo, Desterro de Malta, Itaporanga, João Pessoa, Juazeirinho, Junco do Seridó, Mamanguape, Mataraca, Patos, Piancó, Pilões, Pocinhos, Prata, Rio Tinto, São João do Cariri, São José dos Cordeiros, São José de Espinharas, Sapé, Soledade, Umbuzeiro

CA; AF; BA CHUFPB; CHUNB; MZUSP; I; III; V

Taxon Municipalities Distribution Source

Kentropyx calcarata Spix, 1825** Caaporã, Cruz do Espírito Santo, João Pessoa, Mamanguape, Rio Tinto, Santa Rita, Sapé

AF CHUFPB; CHUNB; MZUSP

Salvator merianae (Duméril e Bibron, 1839)*

Cacimba de Dentro, Campina Grande, João Pessoa, Pocinhos, Rio Tinto, Santa Rita, São José dos Cordeiros

CA; AF CHUFPB; CHUNB; I; III; V; VII

Tropiduridae

Strobilurus torquatus Wiegmann, 1834** João Pessoa, Santa Rita AF CHUFPB

Tropidurus hispidus (Spix, 1825)** Alhandra, Araruna, Areia, Cabaceiras, Cabedelo, Cacimba de Dentro, Campina Grande, Coremas, Desterro de Malta, Gurinhém, João Pessoa, Junco do Seridó, Mamanguape, Mogeiro, Patos, Piancó, Pilões, Puxinanã, Rio Tinto, São João do Cariri, São José dos Cordeiros, São José de Espinharas, Sapé, Sousa, Teixeira, Umbuzeiro

CA; AF; BA CHUFPB; CHUNB; MZUSP; I; III; II; VII; VIII

Tropidurus semitaeniatus (Spix, 1825)* Araruna, Areia, Cabaceiras, Cacimba de Dentro, Campina Grande, Coremas, Gurinhém, Junco do Seridó, Piancó, Pilões, Puxinanã, São João do Cariri, São José de Espinharas, São José dos Cordeiros, Umbuzeiro

CA; BA CHUFPB; MZUSP; I; III; VII

Lissa Dellefrate Franzini et al.754

Figure 1. Lizards from the state of Paraíba: (A) Diploglossus lessonae, (B) Dactyloa punctata, (C) Norops fuscoauratus, (D) Hemidactylus brasilianus, (E) Brasiliscincus heathi, (F) Lygodactylus klugei, (G) Acratosaura mentalis, (H) Anotosaura vanzolinia, (I) Cercosaura ocellata, (J) Dryadosaura nordestina, (K) Micrablepharus maximiliani, (L) Vanzosaura multiscutata, (M) Iguana iguana, (N) Enyalius bibronii, (O) Brasiliscincus heathi. Photography by Daniel O. Mesquita, Fagner R. Delfim, Lucas Ribeiro, Bruno Halluan, and Izabel R. S. da Silva.

Lizard fauna from the state of Paraíba, northeastern Brazil 755

Figure 2. Lizards from state of Paraíba: (A) Copeoglossum nigropunctatum, (B) Psychosaura agmosticha, (C) Psychosaura macrorhyncha, (D) Gymnodactylus darwinii, (E) Gymnodactylus geckoides, (F) Phyllopezus periosus, (G) Polychrus acutirostris, (H) Polychrus marmoratus, (I) Coleodactylus meridionalis, (J) Ameiva ameiva, (K) Ameivula ocellifera, (L) Kentropyx calcarata, (M) Salvator merianae, (N) Strobilurus torquatus, (O) Tropidurus hispidus. Photography by Daniel O. Mesquita, Fagner R. Delfim, Lucas Ribeiro, Bruno Halluan, and Izabel R. S. da Silva.

Amongst the 223 municipalities in the state of Paraíba, 48 (21,52%) are listed in the literature or natural history museums as housing lizard species (Fig. 3) and 46 municipalities had at least one voucher specimen deposited at CHUFPB. João Pessoa municipality, the state’s capital, had the highest number of species registered (21), followed by Cabaceiras and Mamanguape municipalities, both with 19 species registered. João Pessoa and Mamanguape municipalities house legally protected areas and are both located in the eastern region of the state, which was originally covered by Atlantic Forest, while the city of Cabaceiras, which also houses a legally protected area, is located

west of these cities, inland of the state, region of the Caatinga domain. Most of the records listed in this study were obtained in municipalities near the coast and approximately 69% (1909) of the records were obtained in municipalities that house legally protected areas (Fig. 3).

The rarefaction curve obtained with the records from all the institutions and the literature was asymptotic (Fig. 4) and, specifically in CHUFPB, the number of species deposited per decade has not increased in the past years (Fig. 5). Lastly, ED values of the records ranged from 56.242 to 185.194 MY (Fig. 6), with a median of 68.764 MY, and mean value of 76.924 ± 25.287 MY (all those descriptive statistics associated to ED values are presented by excluding S. punctatus ED value). The highest ED value of the state Paraíba species was 185.195 MY (D. lessonae) whereas the lowest was 56.242 MY (Acratosaura mentalis Amaral, 1933 and S. ridleyi). C. meridionalis (130.735 MY), Lygodactylus klugei Smith, Martin and Swain, 1977 (118.846 MY), Iguana iguana Linnaeus, 1758 (99.024 MY), and Salvator merianae Duméril and Bibron, 1839 (98.579 MY) also presented high ED values.

Discussion

Our results indicate that the lizard fauna in the state of Paraíba is still poorly known for most of its municipalities (Fig. 3). Even though the species number sampled in some localities seems to be representative (Fig. 4), it is possible that the total number of lizard species for the state is still underestimated.

Lissa Dellefrate Franzini et al.756

Figure 4. Rarefaction curve (gray line) of lizard species considering total abundance of records in all sources of data and species richness estimator Chao1P (black line).

Figure 3. Number of lizard species record for municipalities in the state of Paraíba in literature and/or scientific museums. Dots represent legally protected areas.

Figure 5. Rarefaction curve of lizards deposited in the Herpetological Collection (CHUFPB) considering the abundance of lizards for each decade (gray line) and species richness estimator Chao2P (black line).

We verified that sampling is poor mainly in the inner part of the state (Fig. 3), an area that belongs to the Caatinga domain. For most municipalities, there were no records of lizards in natural history museums or scientific literature. The Atlantic coast of Paraíba concentrates the largest number of records whereas the western region, known as “Sertão Paraibano”, presents few records, with no species being recorded for most of the municipalities. It is estimated that 17 to 50% of the natural coverage of “Sertão Paraibano” municipalities have already been lost to deforestation, and this percentage is lower than the ones observed in the central area of country (“Borborema” region) and the Atlantic coast (INPE, 2014). However, the advance of livestock

production threatens the region of Sertão, which houses a small number of conservation areas (Fig. 3).

The “Borborema” region, in the centre of the state, houses the highest number of records in the Caatinga domain. However, those records are concentrated in the cities of Cabaceiras and São José dos Cordeiros, which are municipalities where there are conservation areas that have received researchers for several years. The lizard fauna in other municipalities of this region, where the loss of vegetation due to deforestation is the largest in the state (INPE, 2014), remain unknown (Fig. 3).

None of the species was considerable vulnerable or endangered by international (IUCN) and national (MMA) lists. However, we emphasize the presence of

Lizard fauna from the state of Paraíba, northeastern Brazil 757

Figure 6. Phylogenetic relationships among lizards of the state of Paraíba (left). Sphenodon punctatus (Rhynchocephalia: Sphenodontidae) is showed for comparative purposes only. From top to bottom, colored circles at the tips represent lizard families: dark orange = Dactyloidae, turquoise = Polychrotidae, brown = Leiosauridae, gray = Tropiduridae, green = Iguanidae, purple = Anguidae, black = Gymnophtalmidae, red = Teiidae, yellow = Mabuyidae, blue = Gekkonidae, light orange = Phyllodactylidae, light blue = Sphaerodactylidae. The barplot showed on the right depicts species ED values – notice the highest value for S. punctatus (277.601 MY) and the lowest for A. mentalis and S. ridleyi (56.242 MY). MYBP = millions of years before the present; MY = million years.

two of the species listed here in the list of endangered reptiles of the state of Pernambuco (Moura, 2017), a state which borders Paraíba and has some biotic and abiotic similarities with it. Cercosaura ocellata is a Gymnphtalmidae lizard that was recorded in five cities of the Atlantic coast of Paraíba and is considered vulnerable (VU) in Pernambuco. Strobilurus torquatus is a tropidurid lizard recorded only for the municipalities of João Pessoa and Santa Rita and it is also considered vulnerable in Pernambuco. The conservation status of these species in Paraíba is still not known and little is known about their ecology.

Regarding ED values, supposed losses would be more drastic in the case of D. lessonae, C. meridionalis, and I. iguana, since entire lizard families would be vanished from the state. Lygodactylus klugei, however, can be considered an interesting case, since it is only found in the Caatinga, even though other gekkonids also inhabits the biome. If the species becomes locally extinct, the family would still be represented in the state, but a particular evolutionary history (that pertaining to L. klugei branch) would be lost.

Evolutionary Distinctiveness values may be influenced by topology, branch lengths, sampling effort and the method used to generate time-calibrated phylogenies. Still, Isaac et al. (2007) point out that topological errors in some phylogenetic hypothesis do not affect the identity of high-ranking (ED or EDGE) species, especially when calculated for phylogenies containing 100 terminal taxa or more. Thus, the taxonomic substitutions we used here to obtain ED values should not have impacted our conclusions in a very deep way, however, possible divergences between the species in the analysis and the ones used as substitutes could be considered here. Although our phylogeny contains less than 100 terminals, by analysing Isaac and colleagues’ Figure 2, one can clearly notice that clades with more than 30 species is less prone to changes in ED values. We expect that with the development of conceptual and methodological approaches for species delimitation, with robust and comprehensive phylogenetic hypothesis for Squamata, and with detailed assessment of lizard conservation status (in both local, regional, and global scales) we will be able to achieve more reliable ED values for the lizards of the state of Paraíba.

We could not compare our ED values to those of Tonini et al. (2016), because the results presented by them are not detailed for all the species they used on their study. The only comparison we can make is that all the lizard species from the state of Paraíba presented higher ED, but this may be a result of the pruning

process (“tips sampling”) we used here. However, as found by those authors, ED values obtained here are clustered, concentrated in geckos (C. meridionalis and L. klugei) and iguanas (I. iguana). Tonini et al. (2016) also discuss how the interpretation of ED may be complex, depending on factors such as endemism, extinction-risk, conservation status of biomes/regions, species and ecological traits (e.g., life history), and current species protection situation. A valuable factor we can address here is the (specimen) sampling bias associated to some lizard lineages: the data we used to build the rarefaction curve clearly shows that some lineages are easily sampled (e.g., Tropiduridae, Teiidae, and Gymnophtalimidae). The family Polychrotidae and Leiosauridae have species that move slowly in the environment, being more difficult to find. Concerning more exclusive clades, such as genera and species, there are also some examples: Hemidactylus (both H. brasilianus and H. agrius), some gymnophtalmid species (A. mentalis and C. ocellata), and the tropidurid S. torquatus. We also realized that the inventory that we obtained in this study (and other kind of studies that could provide us with species records within the state) are biased (see below), in a pattern very similar to that discussed by Hijmans et al. (2000) and Graham et al. (2004), that demonstrated that natural history and genebank collections can be biased by the fact the collectors tend to sample near main towns, on biological stations and along roads and rivers. Although we presently advocate that aspects of lizard biology may be of great influence on detecting (sampling) them (as discussed in Lovich, 2012 and Mushinsky and McCoy, 2016), we also acknowledge that some lizard lineages may be more prone to be represented in inventories. We can predict that lineages more prone to tolerate human modifications in natural habitats will be those more sampled.

Most of the records are near the coast, in the Atlantic Forest domain (Fig. 3). This region houses the cities of João Pessoa and Mamanguape, where the higher numbers of species (21 and 19, respectively) were found. João Pessoa is the capital of Paraíba and the most economically developed city in the state (IBGE, 2014). Campuses of two of the most important Universities in the state are found in this city, both with traditional undergraduate and graduate programs in Biological Sciences, which probably boosted herpetological studies in João Pessoa and nearby cities, such as Mamanguape and Rio Tinto (~62 Km from João Pessoa).

Another pattern about the geographic distribution of the records is that municipalities that showed high

Lissa Dellefrate Franzini et al.758

number of lizard species have legally protected areas in their territory (Fig. 3). The importance of the creation, maintenance and efficient management of protected areas is linked to the Aichi Target 11, established in the 10th Convention of Biological Diversity (COP, 2010) which states that, by 2020, 17% of terrestrial areas must be effectively protected. This percentage is far from being reached in Paraíba, where just 6.3% and 1.2% of its territory is designated to the conservation of Atlantic Forest and Caatinga biomes, respectively (IBGE, 2014).

Making a comparison between the state of Paraíba and nearby states located above the São Francisco River, one can observe that the state possesses the second lowest number of lizard species, presenting a few more species than Rio Grande do Norte, which has 33 species (Costa and Bérnils, 2018). Pernambuco (51 species) has the highest number of lizard species, followed by Ceará (44) and Alagoas (39). Also, Paraíba is the only one of these five states that does not have endemic or exclusive species. Considering only the Atlantic Forest (as a biogeographical region), Pernambuco and Alagoas are the states with the highest number of lizard species records (28), followed by Rio Grande do Norte (22) and Paraíba (25). Considering the Caatinga biogeographical regions (Velloso et al., 2002), Pernambuco also presents the highest number of lizard species records (30), seven of which are endemic or exclusive. The state of Ceará presents 25 species in the Caatinga domain, however, other 11 species can be found in Caatinga enclaves’ moist forests (“brejos de altitude”). In the states of Paraíba, Alagoas, and Rio Grande do Norte, there are 24 lizard species recorded for the Caatinga Biome in each one of the states.

Several species that were thought to be endemic or to have a narrow distribution in Northeast Brazil, in the Atlantic Forest and the Caatinga, have had their distribution expanded in the last few years due to new investigations and fieldwork. This demonstrates the possibility of discovering new records for the state of Paraíba by investing more sampling effort in the state. As an example, the species Strobilurus torquatus was first recorded in Paraíba after 10 years of fieldwork in two areas where it occurs (Rodrigues et al., 2013).

Widening the knowledge on lizard species that occur across the territory of Paraíba is important to a better understanding and conservation of lizard species. The number and distribution of species in the present list is probably underestimated. In this sense, academic studies and protected areas should play a major role in

providing subsidies for widening our knowledge about the lizards in remnants of Atlantic Forest and Caatinga.

Acknowledgments. We would like to thank all the employees in SUDEMA- PB and Dr. Bráulio A. Santos for information on protected areas in Paraíba. We thank all the photographers who allowed the use of their photos and all scientific museums and respective curators who provided data for this study. We also thank the referees that provided insightful comments and suggestions that improved the final version of this paper. LDF and IRSS thank Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for research fellowship and DOM and DOS thank Conselho Nacional de Pesquisa e Desenvolvimento (CNPQ) for research fellowship.

References

Arzabe, C., Skuk, G., Santana, G.G., Delfim, F. R., Lima, Y.C.C., Abrantes, S.H.F. (2005): Herpetofauna da área de Curimataú, Paraíba. In: Análise das variações da biodiversidade do bioma Caatinga: Suporte a estratégias regionais de conservação. Brasília: Ministério do Meio Ambiente, 264–280.

Colwell, R.K., Chao, A., Gotelli, N.J., Lin, S.-Y., Mao, C.X., Chazdon, R.L., Longino, J.T. (2012): Models and estimators linking individual-based and sample-based rarefaction, extrapolation and comparison of assemblages. Journal of Plant Ecology 5: 3–21.

COP (2010): Convention on biological diversity (CBD).Available at: https://www.cbd.int/2010-target/. Accessed on 04 August 2017

Costa, H.C., Bérnils, R.S. (2018): Répteis do Brasil e suas Unidades Federativas: Lista de espécies. Herpetologia Brasileira 7: 11–48.

Filho, G.A.P., Vieira, W.L.S, Alves, R.R.N., França, F.G.R. (2017): Serpentes da Paraíba: Diversidade e Conservação. 1ª Edição. João Pessoa – PB.

Freire, E.M.X. (1996): Ecologic and zoogeographic study on a fauna of lizards (sauria) from dunas of Natal, Rio Grande do Norte and from restinga de Ponta de Campina, Cabedelo, Paraíba, Brazil. Revista Brasileira de Zoologia 13: 903–921.

Gotelli, N.J., Colwell, R.K. (2001): Quantifying biodiversity: procedures and pitfalls in the measureament and comparison of species richness. Ecology letters 4: 379–391.

Graham, C.H., Ferrier, S., Huettman, F., Moritz, C., Peterson, A.T. (2004): New developments in museum-based informatics and applications in biodiversity analysis. Trends in Ecology and Evolution 19: 497–503.

Hijmans, R.J., Garret, K.A., Huamán, Z., Zhang, D.P., Schreuder, M., Bonierbale, M. (2000): Assessing the geographic representativeness of genebank collections: the case of Bolivian wild potatoes. Conservation Biology 14: 1755–1765.

IBAMA (2015): RPPN Fazenda Almas – Plano de Manejo. Recife – PE

IBGE (2014): Produto interno bruto a preços corrente e produto interno bruto per capita segundo as grandes regiões, as unidades da federação e os municípios – 2010–2013. Available at: http://www.ibge.gov.br/home/estatistica/economia/

Lizard fauna from the state of Paraíba, northeastern Brazil 759

pibmunicipios/2010_2013/default.shtm. Accessed on 07 August 2017

INPE (2014): Monitoramento da Caatinga. Available at: http://www.geopro.crn2.inpe.br/desmatamento.htm. Accessed on March 2018

Isaac, N.J.B., Turvey, S.T., Collen, B., Waterman, C., Baillie, J.E.M. (2007): Mammals on the EDGE: Conservation Priorities Based on Threat and Phylogeny. PLOS One 2: e296.

IUCN (2017): The IUCN red list of threatened species. Version 2017–1. Available at: http://www.iucnredlist.org. Accessed on 20 June 2017

Leal, I.R., Da Silva, J.M.C., Tabarelli, M., Lacher Jr., T.E. (2005): Mudando o curso da conservação da biodiversidade na Caatinga do Nordeste do Brasil. Megadiversidade 1: 139–146.

Lopez, L.C.S., Fracasso, M.P.D.A., Mesquita, D.O., Palma, A.R.T., Riul, P. (2012): The relationship between percentage of singletons and sampling effort: A new approach to reduce the bias of richness estimates. Ecological indicators 14: 164–169.

Lovich, R. E. (2012): Techniques for reptiles in difficult-to-sample habitats. In: Reptile Biodiversity: Standard Methods for Inventory and Monitoring, p. 167–196. Mcdiarmid, R.W., Foster, M.S., Guyer, C., Gibbons, J.W., Chernoff, N. Eds., Berkeley, USA, University of California Press.

Magalhães-Júnior, A.J.C., Pereira, L.C.M., Nicola, P.A., Ribeiro, L.B., Júnior, S.M.D.A. (2014): Distribuição geográfica de Psychosaura agmosticha (Rodrigues, 2000) (Squamata, Mabuyidae). Biotemas 27: 217–222.

Mahdavi, A., Ferreira, L., Sundback, C., Nichol, J.W., Chan, E.P., Carter, D.J., Bettinger, C.J., Patanavanich, S., Chignozha, L., Ben-Joseph, E. (2008): A biodegradable and biocompatible gecko-inspired tissue adhesive. Proceedings of the National Academy of Sciences 105: 2307–2312.

McMurdie, P.J., Holmes, S. (2013): phyloseq: an R package for reproducible interactive analysis and graphics of microbiome census data. PLoS One 8: e61217.

Mendonça, L.E.T., Vieira, W.L.S., Alves, R.R.N. (2014): Caatinga ethnoherpetology: Relationships between herpetofauna and people in a semiarid region of Northeastern Brazil. Amphibian & Reptile Conservation 8: 24–32.

Mesquita, D.O., Faria, R.G., Colli, G.R., Vitt, L.J., Pianka, E.R. (2016): Lizard life-history strategies. Austral Ecology 41: 1–5.

MMA (2011): Ministério do Meio Ambiente: Cadastro nacional de unidades de conservação. Available at: http://www.mma.gov.br/areas-protegidas/cadastro-nacional-de-ucs/consulta-por-uc. Accessed on 12 August 2017.

MMA (2014): Ministério do Meio Ambiente: Livro vermelho da fauna brasileira ameaçada de extinção. Available at: http://pesquisa.in.gov.br/imprensa/jsp/visualiza/index.jsp?jornal=1&pagina=121&data=118/112/2014. Accessed on 20 June 2017.

Moura, G. J. B. (2017): Compilação contínua das espécies de anfíbios e “répteis” do estado de Pernambuco. Available at: https://lehpufrpe.wixsite.com/lehpufrpe/especies-ameacadas-pe. Accessed on 02 February 2018.

Mushinsky, H. R., McCoy, E. D. (2016): Measuring microhabitats used by non-avian reptiles. In: Reptile Ecology and Conservation. A Handbook of Techiniques, p. 254-271. Dodd, C. K. Ed., Oxford, Oxford University Press.

Orme, D., Freckleton, R., Thomas, G., Petzold, T., Fritz, S.,

Isaac, N., Pearse, W. (2013): caper: Comparative Analyses of Phylogenetics and Evolution in R. R package version 0.5.2. Available at: https://CRAN.R-project.org/package=caper.

Paradis, E., Claude, J., Strimmer, K. (2004): APE: analyses of phylogenetics and evolution in R language. Bioinformatics 20: 289–290.

Pianka, E.R., Vitt, L.J. (2003): Lizards. Windows to the evolution of diversity. California, USA, University of California Press.

Prates, A.P.L., Irving, M.D.A. (2015): Conservação da biodiversidade e políticas públicas para as áreas protegidas no Brasil: Desafios e tendências da origem da CDB às Metas de Aichi. Revista Brasileira de Políticas Públicas 5: 28–57.

Queiroz, R.N.M.D., Alves, L.D.S., Oliveira, B.H.S., Albuquerque, H.N.D. (2010): Análise da herpetofauna do Complexo Aluízio Campos. Revista Brasileira de Informações Científicas 1: 22–28.

R Core Team. (2017): R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0. Url: http://www.R-project.org/.

Ragner, P., França, F., França, R., Silva, G. (2014): História natural do lagarto Phyllopezus periosus (Squamata: Phyllodactylidae) em um ambiente semi-árido no Nordeste do Brasil. Revista Biociências 20: 5–12.

Revell, L.J. (2012): phytools: An R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution 3: 217–223.

Rodrigues, K.C., Delfim, F.R., Castro, C.S.S.D., França, F.G.R., Filho, E.L., Mesquita, D.O., Oliveira, F.A.D., Santos, A.C.A.D., Ferrari, S.F., Valença-Montenegro, M.M. (2013): Strobilurus torquatus Wiegmann, 1834 (Squamata: Tropiduridae): New records from Brazilian state of Paraíba and a geographic distribution map. Check List 9: 614–617.

Rodrigues, M.T. (1986): Potencial faunístico da Caatinga. Anais do Simpósio sobre Caatingas e sua exploração racional. Brasília, EMBRAPA, DDT, 163–170.

Rodrigues, M.T. (2000): A new species of Mabuya (Squamata: Scincidae) from the semirarid Caatingas of Northeastern Brazil. Papéis Avulsos de Zoologia 41: 313–328.

Rodrigues, M.T. (2003): Herpetofauna da Caatinga. Ecologia e conservação da Caatinga 4: 181–236.

Santana, G.G., Vieira, W.L.S., Pereira-Filho, G.A., Delfim, F.R., Lima, Y.C.C., Vieira, K.S. (2008): Herpetofauna em um fragmento de floresta atlântica no estado da Paraíba, região nordeste do Brasil. Biotemas 21: 75–84.

Specieslink (2017): Eletronic database. Available at: http://splink.cria.org.br. Accessed on 20 June 2017.

Tabarelli, M., Pinto, L.P., Silva, J.M.C., Hirota, M.M., Bedê, L.C. (2005): Desafios e oportunidades para a conservação da biodiversidade na Mata Atlântica brasileira. Megadiversidade 1: 132–138.

Tonini, J.F.R., Beard, K.H., Ferreira, R.B., Jetz, W., Pyron, R.A. (2016): Fully-sampled phylogenies of squamates reveal evolutionary patterns in threat status. Biological Conservation 204: 23–31.

Uetz, P., Hošek, J. (2016): The reptile database. Available at: reptile-database.reptarium.cz. Accessed on 20 June 2017.

Velloso, A.L, Sampaio, E.V.S.B., Pareyn, F.G.C. (2002): Ecorregiões propostas para o Bioma Caatinga. Instituto de

Lissa Dellefrate Franzini et al.760

Conservação Ambiental. The Nature Conservance do Brasil, Recife, APNE.

Vieira, W.L.S., Gonçalves, M.B.R., Nóbrega, R.P. (2012): Predation on Tropidurus hispidus (Squamata: Tropiduridae) by Lasiodora klugi (Araneae: Therapdosidae) in the semiarid Caatinga region of northeastern Brazil. Biota Neotropica 12: 263–265.

Zheng, Y., Wiens, J.J. (2016): Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species. Molecular Phylogenetics and Evolution 94: 537–547.

Appendix. Specimens included in analysis. One voucher per locality and collection is provided for each species. The total number of analysed specimens per locality and collection is presented after the voucher number.

Diploglossus lessonae: Bananeiras (CHUFPB3516; 1), Cabaceiras (CHUFPB3514; 3), Campina Grande (MZUSP95086; 1), Gurinhém (MZUSP68673; 1), Patos (CHUFPB3518; 1). Dactyloa punctata: João Pessoa (CHUFPB8510; 1), Mamanguape (CHUFPB3669;17, MCZ916; 1, MZUSP5620; 2), Rio Tinto (CHUFPB8393; 30). Norops fuscoauratus: Caaporã (CHUFPB15675; 4), Cabedelo (CHUFPB3895; 2), João Pessoa (CHUFPB55526; 16). Norops ortonii: Cabedelo (CHUFPB3885; 5), João Pessoa (CHUFPB3890; 9, MZUSP65988; 1), Rio Tinto (CHUFPB10324; 8), Sapé (CHUFPB3892; 1, MZUSP60902; 1). Hemidactylus agrius: Cabaceiras (CHUFPB1044; 1, MZUSP59118; 2), Patos (CHUNB61939; 1). Hemidactylus brasilianus: Cabaceiras (CHUFPB858; 5). Hemidactylus mabouia: Araruna (CHUPB9119; 4), Areia (CHUFPB6190; 11), Caaporã (CHUFPB 15782; 1), Cabaceiras (MZUSP71931; 1), Cabedelo (CHUFPB880; 2), Cacimba de dentro (CHUFPB9139; 1), Campina Grande (CHUFPB879; 1), Gurinhém (MZUSP 68653; 1), João Pessoa (CHUFPB869; 21, MZUSP59054; 2), Junco do Seridó (MZUSP44201; 1), Mamanguape (CHUFPB882; 4, MZUSP5402; 1, CHUNB56645; 1), Rio Tinto (CHUFPB 10332; 7), São João do Cariri (CHUFPB12629; 1,. Lygodactylus klugei: Araruna (CHUFPB3334; 5), Cabaceiras (CHUFPB6676; 9, MZUSP59128; 2, CHUNB50490;1), Pombal (CHUFPB3338; 1), São João do Cariri (CHUFPB3333; 20), São José de Espinharas (MZUSP44634; 1, CHUNB50474; 1), São José dos Cordeiros (CHUFPB6076; 2). Acratosaura mentalis: Cabaceiras (CHUFPB5962; 1, MZUSP60786; 1), São José dos Cordeiros (CHUFPB5962; 3). Anotosaura vanzolinia: Cabaceiras (MZUSP60633; 4), São José da Mata (CHUFPB5963; 2), São José dos Cordeiros (CHUFPB5965; 6). Cercosaura ocellata: Alhandra (CHUFPB9358; 2), Conde (MZUSP56959; 1), João Pessoa (CHUFPB3668; 1, MZUSP55525; 1), Mamanguape (CHUNB29005; 1), Rio Tinto (CHUFPB11017; 1). Dryadosaura nordestina: Caaporã (CHUFPB15688; 2), Cabedelo (CHUFPB3574; 3), João Pessoa (CHUFPB2480; 42, MZUSP60634; 1), Mamanguape (CHUFPB3561; 30, CHUNB56711; 1), Rio Tinto (CHUFPB11044; 26). Micrablepharus maximilianii: Alhandra (CHUFPB9339; 2), Gurinhém

Lizard fauna from the state of Paraíba, northeastern Brazil 761

(MZUSP65659; 1), João Pessoa (CHUFPB422; 4, MZUSP65991; 1), Mamanguape (CHUFPB419; 29, MZUSP6546; 1, CHUNB28987; 1), Mataraca (CHUFPB12632; 1), Rio Tinto (CHUFPB11016; 2). Stenolepis ridleyi: Matureia (MZUSP62784; 4). Vanzosaura multiscutata: Cabaceiras (CHUFPB3552; 13, MZUSP66232; 1), Cacimba de dentro (MZUSP9133; 1), Cajazeiras (CHUFPB14894; 1), Juazeirinho (MZUSP3551; 1), Piancó (MZUSP5590; 1), São João do Cariri (MZUSP394; 34), São José dos Cordeiros (CHUFPB801; 18), Soledade (MZUSP4724; 1), Umbuzeiro (MZUSP4723; 1). Iguana iguana: Bananeiras (CHUFPB15865; 1), Caaporã (CHUFPB15739; 1), Cabaceiras (CHUFPB753; 1, MZUSP65617; 1), Coremas (MZUSP5655; 1), Gurinhém (MZUSP65629; 2), Itabaiana (CHUFPB13208; 1), João Pessoa (CHUFPB2694; 13, MZUSP65617; 1), Juazeirinho (MZUSP4851; 1), Junco do Seridó (MZUSP44204; 1), Mamanguape (CHUFPB3128 1), Patos (MZUSP42717; 1), Rio Tinto (CHUFPB9127; 1), São João do Bonfim (CHUFPB2709; 1, ZUEC916; 1), São João do Cariri (CHUFPB2707; 1). Enyalius bibronii: Araruna (CHUFPB9092; 2), Areia (CHUFPB2603; 2, MZUSP66105; 1), Gurinhém (MZUSP65628; 1), São José dos Cordeiros (CHUFPB9329; 1), Sapé (CHUFPB2602; 1), Teixeira (CHUFPB2604; 1). Enyalius catenatus: Mamanguape (CHUFPB2640; 1), Mataraca (CHUFPB2639; 3). Brasiliscincus heathi: Alhandra (CHUFPB9355; 2), Arara (CHUFPB828; 1), Areia (CHUFPB3971; 1, MZUSP65896; 1), Bananeiras (MZUSP67397; 1), Cabedelo (CHUFPB3998; 1), João Pessoa (CHUFPB3979; 18, MZUSP65974; 2), Mamanguape (MZUSP5619; 1, CHUNB29026; 1), Patos (MZUSP42740; 1), Sapé (MZUSP60904; 2), Zabelê (CHUFPB11774; 1). Copeoglossum nigropunctatum: Cruz do Espírito Santo (CHUFPB9166; 2) João Pessoa (CHUFPB3926; 2), Mamanguape (CHUFPB12586; 1, MZUSP5618; 1), Sapé (CHUFPB3928; 1), Rio Tinto (CHUFPB9228; 8). Psychosaura agmosticha: Cabaceiras (CHUFPB3938; 207), Cacimba de dentro (CHUFPB9134; 2). Psychosaura macrorhyncha: Alhandra (MZUSP21480; 1), Cabaceiras (MZUSP60858; 1), Cabedelo (CHUFPB398; 1), Mamanguape (CHUFPB3958; 12), Sapé (MZUSP60891; 1), Rio Tinto (CHUFPB9234; 17). Gymnodactylus darwinii: Caaporã (CHUFPB5667; 1), Cabedelo (CHUFPB3470; 7), João Pessoa (CHUFPB9292; 11), Mamanguape (CHUNB29075; 1). Gymnodactylus geckoides: Araruna (CHUFPB9265; 1), Cabaceiras (CHUFPB3340; 29, ZUEC3563; 1, MZUSP59110; 2),

Cacimba de dentro (CHUFPB9128; 5), João Pessoa (CHUFPB 3366; 1), Mamanguape (CHUFPB 3367; 4, MZUSP59101; 1), Piancó (MCZ59272; 1), Patos (MZUSP56958; 1), São João do Cariri (CHUFPB 9050; 1), São José de Espinharas (MZUSP44528; 2), São José dos Cordeiros (CHUFPB 9330; 2). Phyllopezus periosus: Araruna (CHUFPB9118; 1), Cabaceiras (CHUFPB1930; 28, MCZ172930; 1, CHUNB50455; 1, MZUSP59120; 50), Campina Grande (MZUSP90113; 1, CHUNB49501; 1), Itaporanga (CHUNB61922; 1), São João do Cariri (CHUFPB1955; 8), São José dos Cordeiros (CHUFPB9327; 1). Phyllopezus pollicaris: Araruna (CHUFPB9093; 5), Areia (CHUFPB1981; 1), Boa Vista (CHUFPB1979; 1), Cabaceiras (CHUFPB 1957; 31, MZUSP59121; 37), Itaporanga (CHUNB61919; 1), Junco do Seridó (CHUFPB1980; 1, MZUSP44200; 1, CHUNB61944; 1), Patos (CHUNB61928; 1), Piancó (MZUSP5589; 1), São João do Cariri (CHUFPB1982; 13), São José de Espinharas (MZUSP44527; 1), São José dos Cordeiros (CHUFPB9328; 1). Polychrus acutirostris: Areia (MZUSP65894; 1), Cabaceiras (CHUFPB2594; 2), Gurinhém (CHUFPB2596; 1, MZUSP65646; 2), João Pessoa (CHUFPB2597; 3, MZUSP8389; 2), Mamanguape (CHUFPB2595; 1), Mataraca (CHUFPB12634; 1), São João do Cariri (CHUFPB2593; 1), São José de Espinharas (MZUSP44579; 1). Polychrus marmoratus: Areia (MZUSP65890; 1), João Pessoa (MZUSP59056; 2, CHUFPB2586; 4), Mamanguape (CHUFPB2590; 2), Rio Tinto (CHUFPB9245; 1). Coleodactylus meridionalis: Alhandra (CHUFPB9357; 3), Areia (CHUFPB3213; 9, MZUSP89294; 2), Caaporã (CHUFPB15669; 9), Cabedelo (CHUFPB3214; 5), Cruz do Espírito do Santo (CHUFPB9171; 5), Jacaraú (CHUNB29071), João Pessoa (CHUFPB3219; 29, MZUSP59045; 1), Mamanguape (CHUFPB3248; 45, MCZ 59279; 1, MZUSP59082; 2, CHUNB29069; 1), Matureia (MZUSP65605; 2), Rio Tinto (CHUFPB11045; 58). Ameiva ameiva: Alhandra (CHUFPB 12614; 1), Araruna (CHUFPB 9116; 3), Areia (CHUFPB12478; 1, MZUSP65886; 2) Cabaceiras (CHUFPB2668; 4), Cabedelo (CHUFPB2660; 5), Gurinhém (MZUSP68667; 2), João Pessoa (CHUFPB2647; 10), Junco do Seridó (MZUSP44205; 1), Mamanguape (CHUFPB2650; 13, CHUNB29057; 1), Mogeiro (MZUSP6075; 1) , Rio Tinto (CHUFPB10313; 9), Sapé (CHUFPB2665; 3), São José de Espinharas (MZUSP44619; 1). Ameivula ocellifera: Alhandra (CHUFPB9336; 3), Araruna (CHUFPB9095; 8), Areia (MZUSP65888; 1), Cabaceiras (CHUFPB654; 109, MZUSP59179; 1), Cabedelo (CHUFPB3704; 13), Cacimba de dentro

Lissa Dellefrate Franzini et al.762

(CHUFPB9268; 7), Coremas (MZUSP6260; 1), Cruz do Espírito Santo (CHUFPB9170; 1), Desterro de Malta (CHUFPB3742; 2), Itaporanga (CHUNB61926; 1), João Pessoa (CHUFPB3717; 11, MZUSP65904; 1), Juazeirinho (MZUSP4857; 1), Junco do Seridó (MZUSP44238; 1), Mamanguape (CHUFPB3745; 22, MZUSP6266; 2), Mataraca (CHUFPB3728; 11), Patos (CHUNB61935; 1), Piancó (MZUSP5591; 1), Pilões (CHUFPB12598; 3), Prata (CHUFPB 9486; 1), São João do Cariri (CHUFPB3744; 10), São José de Espinhares (MZUSP44580; 1), Sapé (CHUFPB3738; 2, MZUSP60899; 1), Soledade (MZUSP6078; 1), Rio Tinto (CHUFPB10320; 6), Umbuzeiro (MZUSP4863; 1). Kentropyx calcarata: Caaporã (CHUFPB15663; 3), Cruz do Espírito Santo (CHUFPB9168; 2), João Pessoa (CHUFPB11775; 1, MZUSP56804; 3), Mamanguape (CHUFPB2517; 51, MZUSP56804; 1, CHUNB29047; 1), Rio Tinto (CHUFPB10922; 78, CHUNB29046; 1), Santa Rita (CHUFPB17517; 1), Sapé (CHUFPB2571; 15, MZUSP60885; 2). Salvator merianae: João Pessoa (CHUFPB4196; 4) Rio Tinto (CHUFPB11015; 1), Santa Rita (CHUNB29008; 1). Strobilurus torquatus: João Pessoa (CHUFPB10761; 1), Santa Rita (CHUFPB9163; 1). Tropidurus hispidus: Alhandra (CHUFPB9335; 1), Araruna (CHUFPB2820; 16), Areia

(CHUFPB2840; 3, MZUSP65861; 2), Cabaceiras (CHUFPB585; 95, MZUSP59176; 2), Cabedelo (CHUFPB2792; 5), Cacimba de dentro (CHUFPB9136; 2), Campina Grande (MZUSP55844; 1), Coremas (MZUSP5646; 1), Desterro de Malta (CHUFPB2826; 12), Gurinhém (MZUSP65638; 2), João Pessoa (CHUFPB349; 350, MZUSP5136; 1, CHUNB04191; 1), Junco do Seridó (MZUSP44211; 1), Mamanguape (CHUFPB2825; 9, MZUSP5603; 2, CHUNB29097; 1), Mogeiro (MZUSP5116; 2), Patos (CHUFPB2841; 2), Piancó (MZUSP5583; 1), Pilões (CHUFPB12597; 1), Puxinanã (MZUSP5137; 2), Rio Tinto (CHUFPB10293; 21), São João do Cariri (CHUFPB2808; 22), São José de Espinharas (MZUSP44633; 1), Sapé (CHUFPB2838; 2, MZUSP60898; 1), Teixeira (CHUFPB2821; 3), Umbuzeiro (MZUSP5128; 1). Tropidurus semitaeniatus: Araruna (CHUFPB2728; 8), Areia (CHUFPB12475; 3, MZUSP4630; 3), Cabaceiras (CHUFPB2729; 11, MZUSP65625; 1), Cacimba de dentro (CHUFPB9138; 1), Coremas (MZUSP5647; 1), Gurinhém (MZUSP65631; 1), Junco do Seridó (MZUSP44513; 1), Piancó (MZUSP5584; 1), Pilões (CHUFPB12591; 6), Puxinanã (MZUSP4623; 1), São João do Cariri (CHUFPB2719; 38), São José de Espinharas (MZUSP44532; 1), Umbuzeiro (MZUSP4624; 1).

Accepted by Fábio Hepp

Lizard fauna from the state of Paraíba, northeastern Brazil 763