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Home , Acratosaura mentalis, Ameivula ocellifera, Anotosaura vanzolinia, Brasiliscincus, Cacimba de Dentro, Campina Grande

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

Lizard fauna from the state of , northeastern : 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

Abstract. We compiled a list of 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 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, and Salvator merianae represent important evolutionary histories in the biodiversity of the state.

Keywords. , Caatinga, Atlantic Forest, species list

Introduction The state of Paraíba is situated on the easternmost portion of the South American continent and much are key organisms for studies in numerous of its biodiversity is still unknown. Rodrigues (1986; fields in biology, such as evolution, behaviour, 2000; 2003) demonstrated that samplings are still population, community ecology, and life-history theory deficient in the region. The state comprises vegetation (Mesquita et al., 2016), and have even contributed physiognomies that vary from coastal rainforests to medicine advances (Mahdavi et al., 2008). This (Atlantic Forest domain) to semiarid and arid regions in successful group of inhabits virtually all places the seasonally dry forests (Caatinga domain), including on Earth, except the highest and coldest ones (Pianka Caatinga enclaves’ moist forests, known as “brejos de and Vitt, 2003). Over 6,000 lizard species have already altitude”, which are islands of moist forest surrounded by been described, a number that represents more than half dry forests. Most of the original Atlantic Forest domain of all living reptiles known to date (Uetz and Hošek, coverage has been lost due to anthropic occupation, and 2016). In Brazil, more than 270 species are known to the Caatinga still suffers with the transformation of its occur (Costa and Bérnils, 2018). However, there are still natural landscapes into crops and pastures (Leal et al., many taxonomic problems to be solved and knowledge 2005; Tabarelli et al., 2005). gaps to be fulfilled, mainly in North and Northeast Another worrying issue in this state is the extent of regions of the country. 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 1 Programa de Pós-Graduação em Ciências Biológicas (Zoologia), Departamento Sistemática e Ecologia, 2020, defined in the Aichi Biodiversity Targets, a short Universidade Federal da Paraíba, João Pessoa, PB, Brazil. term international plan to save biodiversity (Prates and * Corresponding author: [email protected] Irving, 2015). 750 Lissa Dellefrate Franzini et al.

The knowledge about the species that occur in a articles from any other collection from the analysis. state is critical to their conservation. Regarding reptile Even though Serpentes is a clade placed among lizards, and amphibian fauna from Paraíba, the few published we decided not to include it in the present study due to species lists focused on small regions and conservation particularities regarding snake biology (high diversity, units (e.g., Arzabe et al., 2005; Santana et al., 2008). In low abundance). Also, a detailed study of the snakes of this sense, we aim to systematize data concerning lizard Paraíba is already available (Filho et al., 2017). species from the state of Paraíba. We consider data Taxonomic nomenclature and classification followed from the literature and the major scientific collections the Brazilian List of Reptile Species (Costa and Bérnils, that house specimens from that state. Additionally, we 2018). We included the introduced species Hemidactylus provide a brief discussion about their conservation mabouia (Moreau de Jonnès, 1818) in the analysis status and identify areas of poor sampling to highlight because it is well-established in Brazil and it is listed in the need of broadening the knowledge of lizards from the Brazilian List of Reptile Species (Costa and Bérnils, the region. 2018). To evaluate the species’ conservation status, we consulted both the International Union for Conservation Material and Methods of Nature (IUCN, 2017) and the Brazilian red list of threatened species (MMA, 2014). In the present list, we compiled data from scientific Even though more specific site information or exact collections and literature. We obtained data from coordinates were available for part of the data, all deposited specimens on Coleção Herpetológica da localities were listed as municipalities, since this was Universidade Federal da Paraíba (CHUFPB), Coleção the sole information available for most records. We built Herpetológica da Universidade de Brasília (CHUNB) a rarefaction curve based on individuals vs. species to and Museu de Zoologia da Universidade de São Paulo evaluate the expected richness on our sampling. For (MZUSP). Specimens from CHUFPB and MZUSP were this, we used the software EstimateS 9.1.0 (Colwell analysed by FRD in order to avoid misidentification. et al., 2012) with 1,000 randomizations, without In CHUNB, the identification of the specimens was reposition, and Chao1P as estimator of species richness performed by the collection curator, Dr. Guarino Rinaldi (Lopez et al., 2012). We used an individual-based curve Colli, following the same guidelines. Complementary because our sampling design was also individual based, data was obtained from the online database Specieslink and because the individuals are the entities that carry (2017). In this database, we obtained records from taxonomic information (Gotelli and Colwell, 2001). Museum of Comparative Zoology – Harvard University Another rarefaction curve was built considering only (MCZ) and Coleção de Répteis do Museu de Zoologia data deposited in CHUFPB, where each decade was da Unicamp (ZUEC). Because we were not able to considered as a sample. This analysis was performed check the identifications obtained from Specieslink, in the same way as described for the individuals vs. we included only those species that, by the experience species curve but with a Chao2P estimator, since it of the authors, could occur in the listed localities. A had a lower standard deviation. We evaluated only the voucher per municipality per institution and the number species deposited over time for the CHUFPB to better of specimens deposited is provided by species in the understand the information provided by this collection, Appendix. which concentrates most of the data about the lizard We carried out the search on scientific literature species from the state of Paraíba. using the terms: “lizards + Paraíba”; “herpetofauna + For phylogenetic comparisons, we used the hypothesis Paraíba”; “reptiles + Paraíba” in Google Scholar and provided by Zheng and Wiens (2016), which is a time- Web of Science databases. Using the same criteria, we calibrated phylogeny for Squamata based on 52 genes carried out a similar search to obtain data concerning sampled for 4,162 species. We pruned their topology to the lizard fauna from the states of Alagoas, Ceará, obtain a tree where only the state of Paraíba’s lizards Pernambuco, and Rio Grande do Norte. We consulted were retained, keeping the time of diversification in data from those states to compare the lizard fauna from the pruned phylogeny and maintaining Sphenodon Paraíba with nearby localities. We excluded from the punctatus Gray, 1842 for comparative reasons. compilation all articles that mentioned only records However, since some species from the state of Paraíba deposited in CHUFPB to avoid data redundancy, data are not represented on that hypothesis, we replaced those from the other collections analysed in this study were not species by the most related species in Zheng and Wiens’ found in the literature and therefore we did not exclude tree. The replacements were: Diploglossus bilobatus Lizard fauna from the state of Paraíba, northeastern Brazil 751

(O’Shaughnessy, 1874) by D. lessonae Peracca, 1890, ED values among the different species IUCN Red List Colobosauroides cearensis Cunha, Lima-Verde and status/categories and, since many species occur in both Lima, 1991 by Dryadosaura nordestina Rodrigues, domains, we could not test for differences in ED values Xavier Freire, Pellegrino and Sites, 2005, Enyalius between Atlantic Forest and Caatinga, as Tonini et al. bilineatus Duméril and Bibron, 1837 by E. bibronii (2016) did. Boulenger, 1885, Enyalius leechii Boulenger, 1885 by E. catenatus Boulenger, 1885, xanti Results Cope, 1863 by darwinii Gray, 1845, We gathered a total of 2,767 occurrence records, Phyllodactylus unctus Cope, 1864 by Gymnodactylus corresponding to 36 lizard species allocated in 27 genera geckoides Spix, 1825, and Tropidurus hygomi Reinhardt of 12 families (Table 1; Fig.1; Fig. 2). Gymnophtalmidae and Lütken, 1862 by Tropidurus semitaeniatus Spix, was the richest family represented, with seven species 1825. We recognize that such substitutions are not ideal, from seven different genera occurring in the state. yet it is the only approach we could have taken, since The families and Sphaerodactylidae were virtually all the phylogenetic hypotheses provided for represented by a single species each, Diploglossus Squamata so far are not based on a complete (or nearly lessonae and Coleodactylus meridionalis Boulenger, complete) sampling of lizard species. Moreover, most evolutionary changes in the tree of life of Squamata 1888, respectively. was the only family that are nested in the deepest nodes (Mesquita et al., 2016), occurred exclusively in the Atlantic Forest, while other which potentially makes the replacement less harmful families were registered in both domains (Caatinga considering our purpose. The resulting tree was then and Atlantic Forest). Nine species (26% of the species used to calculate Evolutionary Distinctiveness (ED) registered) occurred only in the Caatinga and twelve statistic (Isaac et al., 2007) for each lizard species, which species (34%) occurred only in the Atlantic Forest measures each species contributions to phylogenetic (Table 1). For all the species listed here but four, the diversity, and is given in units of time before the present threat status according to IUCN is still not available (not (in our case, millions of years – MY). The ED metric classified yet). The remaining four species are assigned also permits us to discuss the putative taxonomic with the ‘least concern’ status (Table 1). sampling gaps and phylogenetic uniqueness associated ocellifera Spix, 1825 was the species present to the lizards within the boundaries of the state. By using in more municipalities (27), followed by Tropidurus only ED as a surrogate of the “originality” of species hispidus Spix, 1825, being recorded in 26 localities (Isaac et al., 2007), we can assume that, for the state of (Table 1). On the other hand, some species showed a Paraíba, species with the highest ED values should be narrow distribution on the state, being recorded in only prioritized in conservation measures. The interpretation two municipalities: Strobilurus torquatus Wiegmann, is pretty straightforward: if species with high ED values 1834, recorded in municipalities of João Pessoa and are lost within the state boundaries unique evolutionary Santa Rita, in the Atlantic Forest, E. catenatus, recorded histories will be lost in the phylogenetic diversity of the in municipalities of Mamanguape and Mataraca, also state. To obtain ED values we used the package caper in the Atlantic Forest, and Hemidactylus brasilianus (Orme et al., 2013) in R environment, software v 3.4.3 Amaral, 1935, recorded in municipalities of Cabaceiras (R Core Team, 2017). Additional packages used to and São José dos Cordeiros, in the Caatinga. We noticed examine, prune, and plot phylogeny and associated data that the species Copeoglossum nigropunctatum Spix, (ED values) were ape (Paradis et al., 2004), phyloseq 1825 (formerly treated as nigropunctatum) (McMurdie and Holmes, 2013), and phytools (Revell, had been previously recorded in the state, but it was 2012). When pertinent, ED values are showed by the misidentified as Mabuya bistriata Spix, 1825 by arithmetic mean of species ED values ± 1 standard Santana et al. (2008). deviation. It is also worth mentioning the record of Unfortunately, because some species show no variation ridleyi Boulenger, 1887 for the state, which occurred on their status of IUCN Red List (four species) or lack exclusively in the municipality of Matureia, an enclave such status (Table 1), we could not calculate species of moist forest within the Caatinga. And we point out EDGE values, a statistic that satisfy two requirements that Coleodactylus meridionalis was recorded for of conservation priority-setting approaches – the Caatinga, but we point out that the species does not capture of biodiversity and robustness to uncertainty occur in Caatinga stricto sensu, but in more humid (Isaac et al., 2007). Thus, we also could not compare environments. 752 Lissa Dellefrate Franzini et al.

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, , Cabaceiras, Campina CA; AF CHUFPB; MZUSP; I; Grande, Gurinhém, Patos VII Dactyloidae Dactyloa punctata (Daudin, 1802)** João Pessoa, Mamanguape, Rio Tinto AF CHUFPB; MCZ; MZUSP Norops fuscoauratus (D’Orbigny, 1837 in Caaporã, Cabedelo, João Pessoa AF CHUFPB; II Duméril e Bibron, 1837)** Norops ortonii (Cope, 1868)** Cabedelo, João Pessoa, Rio Tinto, Sapé AF CHUFPB; MZUSP; II Gekkonidae Hemidactylus agrius Vanzolini, 1978** Araruna, Cabaceiras, ,Patos CA CHUFPB; CHUNB; MZUSP; I Hemidactylus brasilianus (Amaral, 1935)** Cabaceiras, São José dos Cordeiros CA CHUFPB; III Hemidactylus mabouia (Moreau de Jonnès, Araruna, Areia, Caaporã, Cabaceiras, Cabedelo, CA; AF; BA CHUFPB; CHUNB; 1818)** Cacimba de Dentro, , Gurinhém, MZUSP; I; II João Pessoa, Junco do Seridó, Mamanguape, Rio Tinto, São João do Cariri Lygodactylus klugei (Smith, Martin e Araruna, Cabaceiras, Cacimba de Dentro, Pombal, CA CHUFPB; CHUNB; Swain, 1977)** São João do Cariri, São José de Espinharas, São MZUSP; I; III José dos Cordeiros Acratosaura mentalis (Amaral, 1933)** Araruna, Cabaceiras, Cacimba de Dentro, São José CA CHUFPB; MZUSP; I; dos Cordeiros III vanzolinia Dixon, 1974** Cabaceiras, Campina Grande, São José da Mata, CA CHUFPB; MZUSP; III; São José dos Cordeiros VII ocellata Wagler, 1830** Alhandra, Conde, João Pessoa, Mamanguape, Rio AF CHUFPB; CHUNB; Tinto MZUSP Dryadosaura nordestina Rodrigues, Caaporã, Cabedelo, João Pessoa, Mamanguape, CA; AF CHUFPB; CHUNB; Freire, Pellegrino e Sites, 2005** Rio Tinto MZUSP Micrablepharus maximiliani (Reinhardt e Alhandra, Cabedelo, Cacimba de Dentro, CA; AF CHUFPB; MZUSP; I; II Luetken, 1862)** Gurinhém, João Pessoa, Mamanguape, Mataraca, Rio Tinto Stenolepis ridleyi Boulenger, 1887** Matureia BA MZUSP Vanzosaura multiscutata (Amaral, 1933)** Cabaceiras, Cacimba de Dentro, Cajazeiras, CA CHUFPB; MZUSP; I; , Piancó, São João do Cariri, São José III dos Cordeiros, Soledade, TaxonIguanidae Municipalities Distribution Source Iguana iguana (Linnaeus, 1758)** Araruna, Bananeiras, Caaporã, Cabaceiras, CA; AF; BA CHUFPB; MZUSP; Campina Grande, Coremas, Gurinhém, Itabaiana, ZUEC; I; III; V; VII João Pessoa, Juazeirinho, Junco do Seridó, Mamanguape, Patos, Pocinhos, Rio Tinto, São João do Bonfim, São João do Cariri, São José dos Cordeiros Enyalius bibronii Boulenger, 1885* Araruna, Areia, Gurinhém, São José dos CA; AF; BA CHUFPB; MZUSP; III Cordeiros, Sapé, Teixeira Enyalius catenatus (Wied, 1821)** Mamanguape, Mataraca AF CHUFPB Mabuyidae Brasiliscincus heathi (Schmidt e Inger, Alhandra, Arara, Araruna, Areia, Bananeiras, CA; AF CHUFPB; CHUNB; 1951)** Cabedelo, João Pessoa, Mamanguape, Patos, Sapé, MZUSP; I; III São José dos Cordeiros, Zabelê Copeoglossum nigropunctatum (Spix, Cruz do Espírito Santo, João Pessoa, AF CHUFPB; MZUSP 1825)** Mamanguape, Rio Tinto, Sapé Psychosaura agmosticha (Rodrigues, Cabaceiras, Cacimba de Dentro, Coremas, São CA CHUFPB; I; III; IV 2000)** José dos Cordeiros

Lizard fauna from the state of Paraíba, northeastern Brazil 753

Table 1. Continued.

Taxon Municipalities Distribution Source Psychosaura macrorhyncha (Hoge, 1947)** Alhandra, Cabaceiras, Cabedelo, Mamanguape, CA; AF CHUFPB; MZUSP; II São José dos Cordeiros, Sapé, Rio Tinto Gymnodactylus darwinii (Gray, 1845)** Caaporã, Cabedelo, João Pessoa, Mamanguape AF CHUFPB; CHUNB; II Gymnodactylus geckoides Spix, 1825** Araruna, Cabaceiras, Cacimba de Dentro, Campina CA; AF CHUFPB; MCZ; Grande, João Pessoa, Mamanguape, Patos, Piancó, MZUSP; ZUEC; I; III; São João do Cariri, São José de Espinharas, São VII José dos Cordeiros Phyllopezus periosus Rodrigues, 1986** Araruna, Cabaceiras, Cacimba de Dentro, Campina CA CHUFPB; CHUNB; Grande, Itaporanga, Salgadinho, São João do MCZ; MZUSP; I; III; Cariri, São José dos Cordeiros VI Phyllopezus pollicaris (Spix, 1825)** Araruna, Areia, Boa Vista, Cabaceiras, Cacimba CA; BA CHUFPB; CHUNB; de Dentro, Itaporanga, Junco do Seridó, Patos, MZUSP; I Piancó, São João do Cariri, São José de Espinharas, São José dos Cordeiros acutirostris Spix, 1825** Areia, Cabaceiras, Gurinhém, João Pessoa, CA; AF CHUFPB; MZUSP; III Mamanguape, Mataraca, São João do Cariri, São José dos Cordeiros, São José de Espinharas Polychrus marmoratus (Linnaeus, 1758)** Areia, João Pessoa, Mamanguape, Rio Tinto AF; BA CHUFPB; MZUSP Sphaerodactylidae Coleodactylus meridionalis (Boulenger, Alhandra, Areia, Araruna, Caaporã, Cabedelo, CA; AF; BA CHUFPB; CHUNB; 1888)** Cruz do Espírito Santo, Jacaraú, João Pessoa, MCZ; MZUSP; I; II Mamanguape, Matureia, Rio Tinto (Linnaeus, 1758)** Alhandra, Araruna, Areia, Cabaceiras, Cabedelo, CA; AF; BA CHUFPB; CHUNB; Gurinhém, João Pessoa, Junco do Seridó, MZUSP; I; III; II Mamanguape, , Rio Tinto, São José dos Cordeiros, São José de Espinharas, Sapé (Spix, 1825)** Alhandra, Araruna, Areia, Cabaceiras, Cabedelo, CA; AF; BA CHUFPB; CHUNB; Cacimba de Dentro, Coremas, Cruz do Espírito MZUSP; I; III; V 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, Taxon Soledade,Municipalities Umbuzeiro Distribution Source Spix, 1825** Caaporã, Cruz do Espírito Santo, João Pessoa, AF CHUFPB; CHUNB; Mamanguape, Rio Tinto, Santa Rita, Sapé MZUSP Salvator merianae (Duméril e Bibron, Cacimba de Dentro, Campina Grande, João CA; AF CHUFPB; CHUNB; I; 1839)* Pessoa, Pocinhos, Rio Tinto, Santa Rita, São José III; V; VII dos Cordeiros Strobilurus torquatus Wiegmann, 1834** João Pessoa, Santa Rita AF CHUFPB Tropidurus hispidus (Spix, 1825)** Alhandra, Araruna, Areia, Cabaceiras, Cabedelo, CA; AF; BA CHUFPB; CHUNB; Cacimba de Dentro, Campina Grande, Coremas, MZUSP; I; III; II; VII; Desterro de Malta, Gurinhém, João Pessoa, Junco VIII 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 Tropidurus semitaeniatus (Spix, 1825)* Araruna, Areia, Cabaceiras, Cacimba de Dentro, CA; BA CHUFPB; MZUSP; I; Campina Grande, Coremas, Gurinhém, Junco do III; VII Seridó, Piancó, Pilões, Puxinanã, São João do Cariri, São José de Espinharas, São José dos Cordeiros, Umbuzeiro 754 Lissa Dellefrate Franzini et al.

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) , (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. 756 Lissa Dellefrate Franzini et al.

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 ± Figure 3. Number of lizard species record for municipalities 25.287 MY (all those descriptive statistics associated in the state of Paraíba in literature and/or scientific museums. to ED values are presented by excluding S. punctatus Dots represent legally protected areas. 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), Amongst the 223 municipalities in the state of Lygodactylus klugei Smith, Martin and Swain, 1977 Paraíba, 48 (21,52%) are listed in the literature or (118.846 MY), Iguana iguana Linnaeus, 1758 (99.024 natural history museums as housing lizard species MY), and Salvator merianae Duméril and Bibron, 1839 (Fig. 3) and 46 municipalities had at least one (98.579 MY) also presented high ED values. voucher specimen deposited at CHUFPB. João Pessoa municipality, the state’s capital, had the highest number Discussion of species registered (21), followed by Cabaceiras and Mamanguape municipalities, both with 19 species Our results indicate that the lizard fauna in the registered. João Pessoa and Mamanguape municipalities state of Paraíba is still poorly known for most of its house legally protected areas and are both located in municipalities (Fig. 3). Even though the species number the eastern region of the state, which was originally sampled in some localities seems to be representative covered by Atlantic Forest, while the city of Cabaceiras, (Fig. 4), it is possible that the total number of lizard which also houses a legally protected area, is located species for the state is still underestimated.

Figure 4. Rarefaction curve (gray line) of lizard species Figure 5. Rarefaction curve of lizards deposited in the considering total abundance of records in all sources of data Herpetological Collection (CHUFPB) considering the and species richness estimator Chao1P (black line). abundance of lizards for each decade (gray line) and species richness estimator Chao2P (black line). 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.

We verified that sampling is poor mainly in the production threatens the region of Sertão, which houses inner part of the state (Fig. 3), an area that belongs to a small number of conservation areas (Fig. 3). the Caatinga domain. For most municipalities, there The “Borborema” region, in the centre of the state, were no records of lizards in natural history museums houses the highest number of records in the Caatinga or scientific literature. The Atlantic coast of Paraíba domain. However, those records are concentrated in the concentrates the largest number of records whereas the cities of Cabaceiras and São José dos Cordeiros, which western region, known as “Sertão Paraibano”, presents are municipalities where there are conservation areas few records, with no species being recorded for most of that have received researchers for several years. The the municipalities. It is estimated that 17 to 50% of the lizard fauna in other municipalities of this region, where natural coverage of “Sertão Paraibano” municipalities the loss of vegetation due to deforestation is the largest have already been lost to deforestation, and this in the state (INPE, 2014), remain unknown (Fig. 3). percentage is lower than the ones observed in the central None of the species was considerable vulnerable area of country (“Borborema” region) and the Atlantic or endangered by international (IUCN) and national coast (INPE, 2014). However, the advance of livestock (MMA) lists. However, we emphasize the presence of 758 Lissa Dellefrate Franzini et al. two of the species listed here in the list of endangered process (“tips sampling”) we used here. However, as reptiles of the state of Pernambuco (Moura, 2017), found by those authors, ED values obtained here are a state which borders Paraíba and has some biotic clustered, concentrated in (C. meridionalis and abiotic similarities with it. Cercosaura ocellata and L. klugei) and (I. iguana). Tonini et al. is a Gymnphtalmidae lizard that was recorded in five (2016) also discuss how the interpretation of ED may cities of the Atlantic coast of Paraíba and is considered be complex, depending on factors such as endemism, vulnerable (VU) in Pernambuco. Strobilurus torquatus extinction-risk, conservation status of biomes/regions, is a tropidurid lizard recorded only for the municipalities species and ecological traits (e.g., life history), and of João Pessoa and Santa Rita and it is also considered current species protection situation. A valuable factor vulnerable in Pernambuco. The conservation status of we can address here is the (specimen) sampling bias these species in Paraíba is still not known and little is associated to some lizard lineages: the data we used known about their ecology. to build the rarefaction curve clearly shows that some Regarding ED values, supposed losses would be more lineages are easily sampled (e.g., Tropiduridae, Teiidae, drastic in the case of D. lessonae, C. meridionalis, and and Gymnophtalimidae). The family Polychrotidae I. iguana, since entire lizard families would be vanished and Leiosauridae have species that move slowly in the from the state. Lygodactylus klugei, however, can be environment, being more difficult to find. Concerning considered an interesting case, since it is only found in more exclusive clades, such as genera and species, the Caatinga, even though other gekkonids also inhabits there are also some examples: Hemidactylus (both the biome. If the species becomes locally extinct, the H. brasilianus and H. agrius), some gymnophtalmid family would still be represented in the state, but a species (A. mentalis and C. ocellata), and the tropidurid particular evolutionary history (that pertaining to L. S. torquatus. We also realized that the inventory that klugei branch) would be lost. we obtained in this study (and other kind of studies Evolutionary Distinctiveness values may be influenced that could provide us with species records within the by topology, branch lengths, sampling effort and the state) are biased (see below), in a pattern very similar method used to generate time-calibrated phylogenies. to that discussed by Hijmans et al. (2000) and Graham Still, Isaac et al. (2007) point out that topological et al. (2004), that demonstrated that natural history errors in some phylogenetic hypothesis do not affect and genebank collections can be biased by the fact the the identity of high-ranking (ED or EDGE) species, collectors tend to sample near main towns, on biological especially when calculated for phylogenies containing stations and along roads and rivers. Although we 100 terminal taxa or more. Thus, the taxonomic presently advocate that aspects of lizard biology may substitutions we used here to obtain ED values should be of great influence on detecting (sampling) them (as not have impacted our conclusions in a very deep way, discussed in Lovich, 2012 and Mushinsky and McCoy, however, possible divergences between the species in 2016), we also acknowledge that some lizard lineages the analysis and the ones used as substitutes could be may be more prone to be represented in inventories. We considered here. Although our phylogeny contains less can predict that lineages more prone to tolerate human than 100 terminals, by analysing Isaac and colleagues’ modifications in natural habitats will be those more Figure 2, one can clearly notice that clades with more sampled. than 30 species is less prone to changes in ED values. Most of the records are near the coast, in the Atlantic We expect that with the development of conceptual and Forest domain (Fig. 3). This region houses the cities methodological approaches for species delimitation, of João Pessoa and Mamanguape, where the higher with robust and comprehensive phylogenetic hypothesis numbers of species (21 and 19, respectively) were for Squamata, and with detailed assessment of lizard found. João Pessoa is the capital of Paraíba and the most conservation status (in both local, regional, and global economically developed city in the state (IBGE, 2014). scales) we will be able to achieve more reliable ED Campuses of two of the most important Universities values for the lizards of the state of Paraíba. in the state are found in this city, both with traditional We could not compare our ED values to those of undergraduate and graduate programs in Biological Tonini et al. (2016), because the results presented by Sciences, which probably boosted herpetological studies them are not detailed for all the species they used on in João Pessoa and nearby cities, such as Mamanguape their study. The only comparison we can make is that and Rio Tinto (~62 Km from João Pessoa). all the lizard species from the state of Paraíba presented Another pattern about the geographic distribution higher ED, but this may be a result of the pruning of the records is that municipalities that showed high Lizard fauna from the state of Paraíba, northeastern Brazil 759 number of lizard species have legally protected areas in providing subsidies for widening our knowledge about their territory (Fig. 3). The importance of the creation, the lizards in remnants of Atlantic Forest and Caatinga. maintenance and efficient management of protected areas is linked to the Aichi Target 11, established in the Acknowledgments. We would like to thank all the employees 10th Convention of Biological Diversity (COP, 2010) in SUDEMA- PB and Dr. Bráulio A. Santos for information which states that, by 2020, 17% of terrestrial areas on protected areas in Paraíba. We thank all the photographers who allowed the use of their photos and all scientific museums must be effectively protected. This percentage is far and respective curators who provided data for this study. We from being reached in Paraíba, where just 6.3% and also thank the referees that provided insightful comments and 1.2% of its territory is designated to the conservation suggestions that improved the final version of this paper. LDF of Atlantic Forest and Caatinga biomes, respectively and IRSS thank Coordenação de Aperfeiçoamento de Pessoal de (IBGE, 2014). Nível Superior (CAPES) for research fellowship and DOM and Making a comparison between the state of Paraíba and DOS thank Conselho Nacional de Pesquisa e Desenvolvimento nearby states located above the São Francisco River, (CNPQ) for research fellowship. one can observe that the state possesses the second References lowest number of lizard species, presenting a few more species than Rio Grande do Norte, which has 33 species Arzabe, C., Skuk, G., Santana, G.G., Delfim, F. R., Lima, Y.C.C., (Costa and Bérnils, 2018). Pernambuco (51 species) Abrantes, S.H.F. (2005): Herpetofauna da área de Curimataú, has the highest number of lizard species, followed by Paraíba. In: Análise das variações da biodiversidade do bioma Caatinga: Suporte a estratégias regionais de conservação. Ceará (44) and Alagoas (39). 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(2001): Quantifying biodiversity: the Atlantic Forest and the Caatinga, have had their procedures and pitfalls in the measureament and comparison of distribution expanded in the last few years due to new species richness. Ecology letters 4: 379–391. investigations and fieldwork. This demonstrates the Graham, C.H., Ferrier, S., Huettman, F., Moritz, C., Peterson, A.T. (2004): New developments in museum-based informatics and possibility of discovering new records for the state of applications in biodiversity analysis. Trends in Ecology and Paraíba by investing more sampling effort in the state. Evolution 19: 497–503. As an example, the species Strobilurus torquatus was Hijmans, R.J., Garret, K.A., Huamán, Z., Zhang, D.P., Schreuder, first recorded in Paraíba after 10 years of fieldwork in M., Bonierbale, M. 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(2016): Measuring microhabitats 204: 23–31. used by non-avian reptiles. In: Reptile Ecology and Conservation. Uetz, P., Hošek, J. (2016): The . Available at: A Handbook of Techiniques, p. 254-271. Dodd, C. K. Ed., reptile-database.reptarium.cz. Accessed on 20 June 2017. Oxford, Oxford University Press. Velloso, A.L, Sampaio, E.V.S.B., Pareyn, F.G.C. (2002): Orme, D., Freckleton, R., Thomas, G., Petzold, T., Fritz, S., Ecorregiões propostas para o Bioma Caatinga. Instituto de Lizard fauna from the state of Paraíba, northeastern Brazil 761

Conservação Ambiental. The Nature Conservance do Brasil, Appendix. Specimens included in analysis. One voucher Recife, APNE. per locality and collection is provided for each species. Vieira, W.L.S., Gonçalves, M.B.R., Nóbrega, R.P. (2012): The total number of analysed specimens per locality and on Tropidurus hispidus (Squamata: Tropiduridae) by Lasiodora collection is presented after the voucher number. klugi (Araneae: Therapdosidae) in the semiarid Caatinga region of northeastern Brazil. Biota Neotropica 12: 263–265. Diploglossus lessonae: Bananeiras (CHUFPB3516; Zheng, Y., Wiens, J.J. (2016): Combining phylogenomic and 1), Cabaceiras (CHUFPB3514; 3), Campina Grande supermatrix approaches, and a time-calibrated phylogeny for (MZUSP95086; 1), Gurinhém (MZUSP68673; 1), squamate reptiles (lizards and snakes) based on 52 genes and 4162 species. Molecular Phylogenetics and Evolution 94: 537– Patos (CHUFPB3518; 1). Dactyloa punctata: João 547. 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 762 Lissa Dellefrate Franzini et al.

(MZUSP65659; 1), João Pessoa (CHUFPB422; 4, Cacimba de dentro (CHUFPB9128; 5), João Pessoa MZUSP65991; 1), Mamanguape (CHUFPB419; 29, (CHUFPB 3366; 1), Mamanguape (CHUFPB 3367; 4, MZUSP6546; 1, CHUNB28987; 1), Mataraca MZUSP59101; 1), Piancó (MCZ59272; 1), Patos (CHUFPB12632; 1), Rio Tinto (CHUFPB11016; 2). (MZUSP56958; 1), São João do Cariri (CHUFPB 9050; Stenolepis ridleyi: Matureia (MZUSP62784; 4). 1), São José de Espinharas (MZUSP44528; 2), São José Vanzosaura multiscutata: Cabaceiras (CHUFPB3552; dos Cordeiros (CHUFPB 9330; 2). Phyllopezus 13, MZUSP66232; 1), Cacimba de dentro (MZUSP9133; periosus: Araruna (CHUFPB9118; 1), Cabaceiras 1), Cajazeiras (CHUFPB14894; 1), Juazeirinho (CHUFPB1930; 28, MCZ172930; 1, CHUNB50455; 1, (MZUSP3551; 1), Piancó (MZUSP5590; 1), São João MZUSP59120; 50), Campina Grande (MZUSP90113; do Cariri (MZUSP394; 34), São José dos Cordeiros 1, CHUNB49501; 1), Itaporanga (CHUNB61922; 1), (CHUFPB801; 18), Soledade (MZUSP4724; 1), São João do Cariri (CHUFPB1955; 8), São José dos Umbuzeiro (MZUSP4723; 1). Iguana iguana: Cordeiros (CHUFPB9327; 1). Phyllopezus pollicaris: Bananeiras (CHUFPB15865; 1), Caaporã Araruna (CHUFPB9093; 5), Areia (CHUFPB1981; 1), (CHUFPB15739; 1), Cabaceiras (CHUFPB753; 1, Boa Vista (CHUFPB1979; 1), Cabaceiras (CHUFPB MZUSP65617; 1), Coremas (MZUSP5655; 1), 1957; 31, MZUSP59121; 37), Itaporanga Gurinhém (MZUSP65629; 2), Itabaiana (CHUNB61919; 1), Junco do Seridó (CHUFPB1980; 1, (CHUFPB13208; 1), João Pessoa (CHUFPB2694; 13, MZUSP44200; 1, CHUNB61944; 1), Patos MZUSP65617; 1), Juazeirinho (MZUSP4851; 1), Junco (CHUNB61928; 1), Piancó (MZUSP5589; 1), São João do Seridó (MZUSP44204; 1), Mamanguape do Cariri (CHUFPB1982; 13), São José de Espinharas (CHUFPB3128 1), Patos (MZUSP42717; 1), Rio Tinto (MZUSP44527; 1), São José dos Cordeiros (CHUFPB9127; 1), São João do Bonfim (CHUFPB2709; (CHUFPB9328; 1). Polychrus acutirostris: Areia 1, ZUEC916; 1), São João do Cariri (CHUFPB2707; 1). (MZUSP65894; 1), Cabaceiras (CHUFPB2594; 2), Enyalius bibronii: Araruna (CHUFPB9092; 2), Areia Gurinhém (CHUFPB2596; 1, MZUSP65646; 2), João (CHUFPB2603; 2, MZUSP66105; 1), Gurinhém Pessoa (CHUFPB2597; 3, MZUSP8389; 2), (MZUSP65628; 1), São José dos Cordeiros Mamanguape (CHUFPB2595; 1), Mataraca (CHUFPB9329; 1), Sapé (CHUFPB2602; 1), Teixeira (CHUFPB12634; 1), São João do Cariri (CHUFPB2593; (CHUFPB2604; 1). Enyalius catenatus: Mamanguape 1), São José de Espinharas (MZUSP44579; 1). Polychrus (CHUFPB2640; 1), Mataraca (CHUFPB2639; 3). marmoratus: Areia (MZUSP65890; 1), João Pessoa Brasiliscincus heathi: Alhandra (CHUFPB9355; 2), (MZUSP59056; 2, CHUFPB2586; 4), Mamanguape Arara (CHUFPB828; 1), Areia (CHUFPB3971; 1, (CHUFPB2590; 2), Rio Tinto (CHUFPB9245; 1). MZUSP65896; 1), Bananeiras (MZUSP67397; 1), Coleodactylus meridionalis: Alhandra (CHUFPB9357; Cabedelo (CHUFPB3998; 1), João Pessoa 3), Areia (CHUFPB3213; 9, MZUSP89294; 2), Caaporã (CHUFPB3979; 18, MZUSP65974; 2), Mamanguape (CHUFPB15669; 9), Cabedelo (CHUFPB3214; 5), (MZUSP5619; 1, CHUNB29026; 1), Patos Cruz do Espírito do Santo (CHUFPB9171; 5), Jacaraú (MZUSP42740; 1), Sapé (MZUSP60904; 2), Zabelê (CHUNB29071), João Pessoa (CHUFPB3219; 29, (CHUFPB11774; 1). Copeoglossum nigropunctatum: MZUSP59045; 1), Mamanguape (CHUFPB3248; 45, Cruz do Espírito Santo (CHUFPB9166; 2) João Pessoa MCZ 59279; 1, MZUSP59082; 2, CHUNB29069; 1), (CHUFPB3926; 2), Mamanguape (CHUFPB12586; 1, Matureia (MZUSP65605; 2), Rio Tinto (CHUFPB11045; MZUSP5618; 1), Sapé (CHUFPB3928; 1), Rio Tinto 58). Ameiva ameiva: Alhandra (CHUFPB 12614; 1), (CHUFPB9228; 8). Psychosaura agmosticha: Araruna (CHUFPB 9116; 3), Areia (CHUFPB12478; 1, Cabaceiras (CHUFPB3938; 207), Cacimba de dentro MZUSP65886; 2) Cabaceiras (CHUFPB2668; 4), (CHUFPB9134; 2). Psychosaura macrorhyncha: Cabedelo (CHUFPB2660; 5), Gurinhém (MZUSP68667; Alhandra (MZUSP21480; 1), Cabaceiras 2), João Pessoa (CHUFPB2647; 10), Junco do Seridó (MZUSP60858; 1), Cabedelo (CHUFPB398; 1), (MZUSP44205; 1), Mamanguape (CHUFPB2650; 13, Mamanguape (CHUFPB3958; 12), Sapé (MZUSP60891; CHUNB29057; 1), Mogeiro (MZUSP6075; 1) , Rio 1), Rio Tinto (CHUFPB9234; 17). Gymnodactylus Tinto (CHUFPB10313; 9), Sapé (CHUFPB2665; 3), darwinii: Caaporã (CHUFPB5667; 1), Cabedelo São José de Espinharas (MZUSP44619; 1). Ameivula (CHUFPB3470; 7), João Pessoa (CHUFPB9292; 11), ocellifera: Alhandra (CHUFPB9336; 3), Araruna Mamanguape (CHUNB29075; 1). Gymnodactylus (CHUFPB9095; 8), Areia (MZUSP65888; 1), geckoides: Araruna (CHUFPB9265; 1), Cabaceiras Cabaceiras (CHUFPB654; 109, MZUSP59179; 1), (CHUFPB3340; 29, ZUEC3563; 1, MZUSP59110; 2), Cabedelo (CHUFPB3704; 13), Cacimba de dentro Lizard fauna from the state of Paraíba, northeastern Brazil 763

(CHUFPB9268; 7), Coremas (MZUSP6260; 1), Cruz (CHUFPB2840; 3, MZUSP65861; 2), Cabaceiras do Espírito Santo (CHUFPB9170; 1), Desterro de Malta (CHUFPB585; 95, MZUSP59176; 2), Cabedelo (CHUFPB3742; 2), Itaporanga (CHUNB61926; 1), (CHUFPB2792; 5), Cacimba de dentro (CHUFPB9136; João Pessoa (CHUFPB3717; 11, MZUSP65904; 1), 2), Campina Grande (MZUSP55844; 1), Coremas Juazeirinho (MZUSP4857; 1), Junco do Seridó (MZUSP5646; 1), Desterro de Malta (CHUFPB2826; (MZUSP44238; 1), Mamanguape (CHUFPB3745; 22, 12), Gurinhém (MZUSP65638; 2), João Pessoa MZUSP6266; 2), Mataraca (CHUFPB3728; 11), Patos (CHUFPB349; 350, MZUSP5136; 1, CHUNB04191; (CHUNB61935; 1), Piancó (MZUSP5591; 1), Pilões 1), Junco do Seridó (MZUSP44211; 1), Mamanguape (CHUFPB12598; 3), Prata (CHUFPB 9486; 1), São (CHUFPB2825; 9, MZUSP5603; 2, CHUNB29097; 1), João do Cariri (CHUFPB3744; 10), São José de Mogeiro (MZUSP5116; 2), Patos (CHUFPB2841; 2), Espinhares (MZUSP44580; 1), Sapé (CHUFPB3738; 2, Piancó (MZUSP5583; 1), Pilões (CHUFPB12597; 1), MZUSP60899; 1), Soledade (MZUSP6078; 1), Rio Puxinanã (MZUSP5137; 2), Rio Tinto (CHUFPB10293; Tinto (CHUFPB10320; 6), Umbuzeiro (MZUSP4863; 21), São João do Cariri (CHUFPB2808; 22), São José 1). Kentropyx calcarata: Caaporã (CHUFPB15663; 3), de Espinharas (MZUSP44633; 1), Sapé (CHUFPB2838; Cruz do Espírito Santo (CHUFPB9168; 2), João Pessoa 2, MZUSP60898; 1), Teixeira (CHUFPB2821; 3), (CHUFPB11775; 1, MZUSP56804; 3), Mamanguape Umbuzeiro (MZUSP5128; 1). Tropidurus semitaeniatus: (CHUFPB2517; 51, MZUSP56804; 1, CHUNB29047; Araruna (CHUFPB2728; 8), Areia (CHUFPB12475; 3, 1), Rio Tinto (CHUFPB10922; 78, CHUNB29046; 1), MZUSP4630; 3), Cabaceiras (CHUFPB2729; 11, Santa Rita (CHUFPB17517; 1), Sapé (CHUFPB2571; MZUSP65625; 1), Cacimba de dentro (CHUFPB9138; 15, MZUSP60885; 2). Salvator merianae: João Pessoa 1), Coremas (MZUSP5647; 1), Gurinhém (CHUFPB4196; 4) Rio Tinto (CHUFPB11015; 1), (MZUSP65631; 1), Junco do Seridó (MZUSP44513; Santa Rita (CHUNB29008; 1). Strobilurus torquatus: 1), Piancó (MZUSP5584; 1), Pilões (CHUFPB12591; João Pessoa (CHUFPB10761; 1), Santa Rita 6), Puxinanã (MZUSP4623; 1), São João do Cariri (CHUFPB9163; 1). Tropidurus hispidus: Alhandra (CHUFPB2719; 38), São José de Espinharas (CHUFPB9335; 1), Araruna (CHUFPB2820; 16), Areia (MZUSP44532; 1), Umbuzeiro (MZUSP4624; 1).

Accepted by Fábio Hepp