A peer-reviewed open-access journal ZooKeys 1054: 127—153 (2021) SE Rea eae #ZooKeys https:/ / ZOO keys. pensoft.net Launched to accelerate biodiversity research Selva Zoque, Mexico: an important Mesoamerican tropical region for reptile species diversity and conservation José Luis Aguilar-Lépez', Ricardo Luria-Manzano’, Eduardo Pineda', Luis Canseco-Marquez? | Red de Biologia y Conservacién de Vertebrados, Instituto de Ecologia A. C., Carretera antigua a Coatepec 351, El Haya, Xalapa, C.P 91073, Veracruz, México 2. Departamento de Ecologia, Instituto de Biociéncias, Universidade de Séo Paulo, Rua do Matéo, Travessa 14, Cidade Universitaria, 05508-090, Séo Paulo, Sao Paulo, Brazil 3 Departamento de Biologia Evolutiva, Laboratorio de Herpetologta, Facultad de Ciencias, Uni- versidad Nacional Auténoma de México, A.P 70-399, C.P 04510, México City, México Corresponding author: Ricardo Luria-Manzano (ricardolm@ib.usp.br) Academic editor: Anthony Herrel | Received 27 April 2021 | Accepted 8 July 2021 | Published 3 August 2021 http://z00bank.org/F1D9569A-E45E-4B64-857F-9832 164D0DBO Citation: Aguilar-Lopez JL, Luria-Manzano R, Pineda E, Canseco-Marquez L (2021) Selva Zoque, Mexico: an important Mesoamerican tropical region for reptile species diversity and conservation. ZooKeys 1054: 127-153. https://doi.org/10.3897/zookeys.1054.67916 Abstract The Selva Zoque region is characterized by a great variety of ecosystems for which there is little informa- tion about reptile species diversity and their conservation status. This study is the first assessment of the species richness, composition, and conservation status of reptiles of this region. Additionally, this informa- tion is compared with that of seven other tropical regions in northern Mesoamerica. In total, 141 native reptile species belonging to 81 genera and 29 families are recorded for the Selva Zoque region. Sixty species (42% of the total) recorded in Selva Zoque are in high-risk categories according to the Mexican Ministry of the Environment, the highest number for the Mexican regions of Mesoamerica. According to the IUCN, six species are in high-risk categories, seven species are in Data Deficient, and 23 (16%) have not been evaluated yet. According to the Environmental Vulnerability Scores approach, 28 species (20%) are in the high vulnerability category. The Selva Zoque species composition is most similar to Los Tuxtlas and Lacandona regions, and most dissimilar to Sian Ka’an Biosphere Reserve. The reptilian fauna of Selva Zoque has a distinctive composition, with the highest number (11 species) of endemic reptiles in the northern Mesoamerican, and species from two biogeographic provinces: the Gulf of Mexico and Copyright José Luis Aguilar-Lépez et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 128 José Luis Aguilar-Lopez et al. / ZooKeys 1054: 127-153 (2021) the Mexican Pacific Coast. These results indicate that the Selva Zoque is the most diverse region in native reptile species in northern Mesoamerica, highlighting it as extremely important for the conservation of the reptile fauna at local (southern Mexico) and regional levels (northern Mesoamerica). Keywords Compositional similarity, conservation value, Data Deficient, reptile fauna, species composition, species richness Introduction The 25 biodiversity hotspots identified by Myers et al. (2000) share two characteristics: each one harbors endemic plant species representing at least 0.5% of the global total, and have lost = 70% of their primary vegetation. Of these regions, Mesoamerica ranks fifteenth in relation to the latter characteristic (80% of primary vegetation lost; Myers et al. 2000), and ranks third in deforestation rate among the 13 hotspots for which information is available (Brooks et al. 2002). Despite this scenario, some areas still remain covered by primary vegetation (FAO 2011), highlighting the ongoing conser- vation value of this region. One such region is the Selva Zoque, composed by the Uxpanapa-Chimalapas zone (UC) and El Ocote Biosphere Reserve, is located on the Isthmus of Tehuantepec in the states of Veracruz, Oaxaca, and Chiapas in southern Mexico. The Selva Zoque region is the second largest extension of well-conserved tropical forest in northern Mesoa- merica, is a Pleistocene refuge with high number of endemic species (Pérez-Farrera et al. 2016), contains a large variety of vegetation types (Peterson et al. 2003) and has a broad range of elevation spanning 100 to 2300 ma.s.l. Additionally, the Selva Zoque region, together with the protected area La Sepultura Biosphere Reserve, in the state of Chiapas, make up the La Selva Zoque-La Sepultura Priority Conservation Area (Ar- riaga et al. 2000b). Currently, detailed knowledge of the vertebrate species diversity in the entire Selva Zoque, or a large portion of it, is available only for birds (Peterson et al. 2003), mammals (Lira-Torres et al. 2012), and amphibians (Aguilar-Lépez et al. 2016a). For other groups such as reptiles, our knowledge of species richness, species composition, and conservation status is limited to El Ocote Biosphere Reserve, where the reptile diversity has been revised on several studies (Reynoso et al. 2011; Luna- Reyes et al. 2017; Mufioz-Alonso et al. 2017) and scarce in UC zone. Herpetological expeditions have been carried out in the UC zone since at least the middle of the last century (Taylor 1951; Duellman 1960; Lynch and Wake 1989) but have focused mainly on amphibians. Moreover, herpetofaunal collection has been limited to few localities, and extensive portions of the region remain with no information. Worldwide, habitat modification represents the most common threat to terrestrial reptile species, with one in five species included in high-risk categories of extinction (Vulnerable, Endangered, or Critically Endangered) by the IUCN. A further one in five species is listed in the Data Deficient category, and four of ten have not been Reptiles of the Selva Zoque region, Mexico 129 evaluated according to the criteria of the Red List (Uetz et al. 2018; IUCN 2021). In Mexico, it is estimated that 13% of reptile species are threatened and for another 16% there is insufficient information (in Data Deficient category) to evaluate its extinction risk level (UCN 2021). On the other hand, about half of the species are included in the high-risk of extinction categories on the species list compiled by the Mexican Ministry of the Environment (NOM-059-SEMARNAT-2010). Given the variety of tropical forests that cover the Selva Zoque region, along with its geographic location and environmental heterogeneity (Wendt 1987; de Teresa 2000; SEMARNAT 2001), a high reptile diversity may inhabit in this region, with a significant portion of species under high risk of extinction. Based on a comprehensive review of databases, scientific literature, and fieldwork, we provide the first assessment of species richness, species composition and distribu- tion, and conservation status of the reptile fauna inhabiting the Selva Zoque region. Additionally, we compared this information with other tropical regions on the north- ern end of Mesoamerica. Materials and methods Study site The Selva Zoque region is located in southern Mexico east of the Isthmus of ‘Te- huantepec (Fig. 1). Mountains and hills dominate the region (Wendt 1987; SEMAR- NAT 2001; Ortiz-Pérez et al. 2004). The region is covered by several vegetation types, the main ones are evergreen tropical forest (100-1000 m a.s.l.), semi-evergreen tropi- cal forest (600-1200 m a.s.l.), deciduous tropical forest (100-600 m a.s.l.), tropi- cal montane cloud forest (1100-1800 m a.s.l.) and pine-oak forest (1800-2300 m a.s.l.) (Wendt 1987; SEMARNAT 2001). Mean annual temperature ranges from 12 to 23 °C and mean annual rainfall ranges from 800 to 4400 mm (Vidal-Zepeda 1990; SEMARNAT 2001). We delimited the study area using the polygon set by Arriaga et al. (2000a) for the Selva Zoque-La Sepultura Priority Conservation Area but excluded La Sepultura Biosphere Reserve (Fig. 1). Data collection We consulted two databases between January 2017 and May 2021: the National In- formation System on Biodiversity (SNIB) curated by the National Commission for the Knowledge and Use of Biodiversity (CONABIO) and the Global Biodiversity In- formation Facility (GBIF; www.gbif.org). We also reviewed the scientific literature on reptiles from the study region (Navarro-Singiienza and Meave-Castillo 1998; Espinoza et al. 1999; Reynoso et al. 2011; Carmona-Torres 2013; Aguilar-Lépez et al. 2014; Canseco-Marquez and Ramirez-Gonzalez 2015; Scarpetta et al. 2015; Aguilar-Lépez et al. 2016b; Gray et al. 2016; Luna-Reyes et al. 2017; Mufioz-Alonso et al. 2017; 130 José Luis Aguilar-Lopez et al. / ZooKeys 1054: 127-153 (2021) Figure |. Location of the Selva Zoque region. The yellow line denotes our study area limits, the thin black line indicates state boundaries, and black circles denote localities in which reptiles have been recorded. Campbell et al. 2018; del Moral-Flores et al. 2019). Additionally, we conducted field- work during 2013 and 2014 in surroundings of Arroyo Zarco, Uxpanapa, in Verac- ruz (17°11'N, 94°28'W), and San Francisco La Paz (17°5'N, 94°8'W), La Fortaleza (17°9'N, 94°13'W), and La Esmeralda (17°9'N, 94°46'W), in Santa Maria Chimalapa, Oaxaca. We surveyed areas with evergreen tropical forest and semi-evergreen tropical forest using standard visual encounter survey techniques (Crump and Scott 1994) dur- ing the day and at night. Cumulative sampling effort was 3250 person-hours. We cross-checked records obtained from the three sources of information to avoid duplication, since the databases consulted might have records of the same specimens. Only records with precise geographic coordinates or detailed information about the collection and observation site were included, and from these records we compiled a general database. To compare the data we compiled for the Selva Zoque region with the data from seven other tropical regions, we obtained information about reptile species richness and species composition from: 1) Los Tuxtlas in Veracruz (Lépez-Luna 2017), 2) Las Choapas municipality in Veracruz (Aguilar-Lépez and Canseco-Marquez 2006), 3) La Sepultura Biosphere Reserve in Chiapas (Nufiez-Orantes and Mufoz-Alonso 2000; Reynoso et al. 2011; Clause et al. 2020a, b), 4) the Lacandona rainforest in Chiapas (Hernandez-Ordéfez et al. 2015), 5) the Calakmul Biosphere Reserve in Campeche Reptiles of the Selva Zoque region, Mexico 131 (Calderén-Mandujano et al. 2010; Colston et al. 2015), 6) the Sian Ka’an Biosphere Reserve in Quintana Roo (Calderén-Mandujano et al. 2008), and 7) the Mayan Forest in Guatemala (Lee 1996; Campbell 1998). Data processing and analysis To identify the spatial distribution of reptile records in the study region, we projected all geo-referenced records onto our study area polygon using ArcGIS software, ver- sion 10.0 (ESRI 2010). To determine the distribution and taxonomically standardize the data set of species that inhabit the Selva Zoque and the other tropical regions, we consulted the specialized literature documenting taxonomic changes and descriptions of new species (Wiister et al. 2005; Castoe et al. 2009; Kohler 2010; Linkem et al. 2011; Cadle and Savage 2012; Hedges and Conn 2012; Iverson et al. 2013; Porras et al. 2013; Kohler et al. 2014; Ruane et al. 2014; Blair et al. 2015; Meza-Lazaro and Nieto-Montes de Oca 2015; Card et al. 2016; Gray et al. 2016; Kohler et al. 2016; McCranie and Hedges 2016; Wallach 2016; Nieto-Montes de Oca et al. 2017; Campbell et al. 2018; Carbajal-Marquez et al. 2020; Jadin et al. 2020; McCranie et al. 2020; Reyes-Velasco et al. 2020; Ramirez-Reyes et al. 2021). Using the compiled data, we defined four distribution categories: species distributed outside Mesoamerica as widely distributed species (WD), species restricted to Mesoamerica (MA), spe- cies restricted to northern Mesoamerica (MAMx), and species restricted to one of the eight regions considered. We delimited Mesoamerica as suggested by Campbell (1999) and considered northern Mesoamerica as the zone corresponding to Mexico. To determine the extinction risk category for each species, we consulted the list of Species at Risk published by SEMARNAT, updated in 2018 (NOM-059-SEMAR- NAT-2010), the Red List maintained by the International Union for the Conser- vation of Nature IUCN 2021), and the Environmental Vulnerability Score (EVS) proposed by Wilson et al. (2013). SEMARNAT’s categories are: Subject to Special Protection (Pr), Threatened (A), and Endangered (P). The IUCN’s three high-risk categories are: Vulnerable (VU), Endangered (EN) and Critically Endangered (CR); its low-risk categories are: Least Concern (LC) and Near Threatened (NT). We also included species in the Data Deficient (DD) category, and those Not evaluated (NE) by the IUCN. In addition, we consulted the EVS of Mexican reptile species that have been evaluated and assigned to one of three categories of vulnerability to environ- mental degradation: low (3-9), medium (10-13) and high (14-19). For the Mayan Forest in Guatemala, we were only able to assign the IUCN categories since the area lies outside of SEMARNAT’’s jurisdiction and there is not an evaluation of EVS for reptiles of Guatemala. We compared reptilian faunal composition between regions using Jaccard’s simi- larity index (Magurran 2004), which uses presence-absence data and is expressed as: 7 a atb+c J 132 José Luis Aguilar-Lopez et al. / ZooKeys 1054: 127-153 (2021) where a = the number of species shared between the two sites under comparison, 6 = number of species exclusive to the first site, and c = number of species exclusive to the second site. The index ranges from zero to one, with zero indicating that no species are shared between the sites being compared, and one indicating that all species are found in both sites. We plotted a dendrogram using PAST software version 2.17c (Hammer et al. 2001) to represent the relationship between sites in terms of their similarity in species composition according to the Jaccard index. For this analysis, we only included native species. Results Species richness, distribution, and conservation status A total of 141 native reptile species belonging to 81 genera and 29 families has been recorded at the Selva Zoque region. These comprise 62 species of lizards, 70 snakes, seven turtles, and two crocodilians (Appendix 1). The best represented families are Dipsadidae and Colubridae, with 29 and 25 species, respectively, followed by Dacty- loidae with 18 species, Phrynosomatidae with nine species, and Viperidae with seven species. The families with the fewest species in the region are Eublepharidae, Heloder- matidae, Mabuyidae, Phyllodactylidae, Scincidae, Boidae, Natricidae, Sybinophiidae, Leptotyphlopidae, Loxocemidae, Dermatemydidae, Emydidae, and Geoemydidae, with one species each one. Three non-native species have been recorded in the region, the lizards Anolis sagrei Duméril & Bibron Gehyra mutilata (Wiegmann) and Hemidac- tylus frenatus Duméril and Bibron. During our fieldwork (2013-2014), we recorded 48 species (Appendix 1), although all of them had been previously recorded. Reptiles have been recorded mostly on the periphery of the study region, notably on western, southern and eastern end portions. In the northwest, northeast, and west, reptile collections are located below 1000 m a.s.l., while in the southeast, most of col- lections are between 1000 and 2000 m a.s.l. The central portion of Chimalapas, the mountainous zone known as Espinazo del Diablo in Uxpanapa and a zone between El Ocote Biosphere Reserve and UC zones corresponding to the northwest extreme of Cintalapa in Chiapas, remain with no collection of reptiles (Fig. 1). The distribution of eleven of the 141 native species (8%) is restricted to the Selva Zoque region, 26 species (18%) are distributed in the northern part of Mesoamerica that corresponds to Mexico, 66 species (47%) are distributed on Mesoamerica, and 38 species (27%) have a wide distribution, extending beyond Mesoamerica (Appendix 1). Of the reptile species recorded in the Selva Zoque region, 60 species (42%) are in high-risk categories according to SEMARNAT: 39 species in the Subject to Special Protection category (Pr), 18 species are in the Threatened category (A) and three are Endangered (P). According to the IUCN Red List, six species (4%) are included in high-risk categories: three are Vulnerable (VU), two are Endangered (EN), and one is Critically Endangered (CR). Additionally, seven species are in the Data Deficient cat- Reptiles of the Selva Zoque region, Mexico 1:33 egory (DD) and 23 species have not been evaluated (NE). The remaining 105 species are in low-risk categories. According to the EVS system, 28 species (20%) are in the high vulnerability category (Appendix 1). Comparison of the richness, composition, and conservation status of the rep- tile species from the Selva Zoque region with that of other tropical regions With 141 native species recorded, the Selva Zoque region harbors the highest reptile species richness among the tropical regions considered in this study, surpassing Los Tuxtlas (113 species). The Mayan Forest ranks third (107 species), followed by La Lacandona (89 species), La Sepultura (79 species), Calakmul (73 species), Sian Ka’an (63 species), and finally Las Choapas (56 species; Table 1). The Los Tuxtlas Biosphere Reserve, with four species, has the highest diversity of non-native species, followed by Selva Zoque and Calakmul with three. La Sepultura, Sian Ka’an, and Las Choapas have two non-native species, and the Mayan Forest and Lacandona each have only one. The dendrogram indicates that La Sepultura has the most dissimilar species com- position of the nine regions (Fig. 2). This Biosphere Reserve is followed by Selva Zoque, Las Choapas and Los Tuxtlas in terms of their dissimilarity in relation to the remaining regions. The four remaining regions in turn form a group with a value of Jaccard index [C] = 0.5 (Fig. 2). The composition analysis by pairs indicates that Los Tuxtlas had the species composition most similar to that of Selva Zoque, with 44% shared species (C’ = 0.44), followed by Lacandona (C,= 0.39), whereas the region with j j the least similar species composition to Selva Zoque was Sian Ka’an (C= 0.20). The regions with the most similar species composition were Lacandona and Mayan Forest in Guatemala (C= 0.72), followed by Sian Ka’an and Calakmul (C= 0.58). The least similar regions were Sian Ka’an and La Sepultura (C,= 0.14). The Selva Zoque and Los Tuxtlas regions have the highest number of endemic species to the regions, with eleven each, followed by La Sepultura (three species) and Mayan Forest with one endemic species while the rest of regions do not have endemic species (Appendix 1). Also, the Selva Zoque region has the highest number of species in high-risk categories of extinction according to the NOM-059 criteria, with 60 spe- Table |. Taxonomic composition of native reptile species recorded in the Selva Zoque region and seven other regions in northern Mesoamerica. Non-native species are not included in the taxonomic composi- tion data. Tropical region Orders Families Genera Lizards Snakes Turtles Crocodilians Native species Non-native species Selva Zoque 3 29 81 62 70 7 2 141 3 Los Tuxtlas 3 29 75 35 63 14 1 113 4 Mayan Forest 3 26 70 35 61 9 2 107 1 Lacandona 3 25 64 28 52 Z 2 89 1 La Sepultura 2 25 59 29 47 3 0 79 2 Calakmul ) 21 51 24 39 9 1 73 3 Sian Ka’an 3 23 51 25 26 10 2 63 2 Las Choapas 3 20 4] 20 28 Z 1 56 2 134 José Luis Aguilar-Lopez et al. / ZooKeys 1054: 127-153 (2021) Selva Zoque Los Tuxtlas Sian Ka’an Calakmul Lacandona Mayan Forest Las Choapas La Sepultura 0 01 O2 03 O04 O05 06 07 O08 0.9 1 Composition similarity (Jaccard index) Figure 2. Similarity in reptile species composition for eight tropical areas in northern Mesoamerica. cies, followed by Los Tuxtlas with 46 species. The other regions have from 33 species (La Sepultura) to 25 species (Las Choapas) in high-risk categories (Fig. 3A). Based on the IUCN Red List, the number of reptiles in high-risk categories of extinction is high- est in Los Tuxtlas, with ten species, and the other regions have from one to six species in these categories. The Selva Zoque region has the highest number of species classified as Not evaluated (NE) and with Data Deficient (DD) on the IUCN Red List with 30 species, followed by Los Tuxtlas (24 species); the remaining regions have from eight to 14 NE and DD species (Appendix 1; Fig. 3B). The Selva Zoque has the highest number of species (28) included in the high vulnerability category to environmental degradation, followed by Los Tuxtlas with 18 species; the remaining regions have be- tween five and ten species (Fig. 3C). Discussion Our results show that the Selva Zoque region harbors the greatest reptile species rich- ness in northern Mesoamerica, with a distinctive species composition and several species inhabit exclusively on Selva Zoque region. Furthermore, one in every two reptile species that inhabit in Selva Zoque is threatened, is highly vulnerable or there is not sufficient information to know its extinction risk level. All this underscore the importance of the Selva Zoque region for reptile conservation in Mexico and Mesoamerica. This region also offers opportunities to study unexplored well-preserved tropical forest areas, as well as species for which little is known about their biology, ecology, and conservation status. The Selva Zoque region is more diverse in native reptile species than even Los Tuxtlas, which has 28 fewer native species (L6pez-Luna 2017). In the Selva Zoque region three exotic species (Anolis sagrei, Gehyra mutilata, Hemidactylus frenatus) have Reptiles of the Selva Zoque region, Mexico 135 Selva Zoque Mayan Forest Los Tuxtlas Lacandona La Sepultura Las Choapas Calakmul Sian Ka‘an Selva Zoque Mayan Forest Los Tuxtlas Lacandona La Sepultura Las Choapas Calakmul Sian Ka’an Selva Zoque Mayan Forest Los Tuxtlas Lacandona La Sepultura Las Choapas Calakmul Sian Ka’an a) — So i) =) td So oy i=) vi i) oO oS Number of species Figure 3. Number of reptile species in high extinction-risk categories according to A NOM-059-SE- MARNAT-2010 B IUCN Red List, and C the Environmental Vulnerability Score, for eight tropical regions in northern Mesoamerica. The black bars in the three plots represent the number of species in high extinction-risk categories, and the grey bars in the graph B represent the number of species not evaluated or in the Data Deficient category by the IUCN. 136 José Luis Aguilar-Lopez et al. / ZooKeys 1054: 127-153 (2021) been recorded, fewer than other regions such as Los Tuxtlas, which is located near the coast. The occurrence of the highest diversity of native species in some portions of Selva Zoque over that of other tropical regions has been previously observed in amphibians (Aguilar-Lopez et al. 2016a). The 141 native reptile species recorded in the Selva Zoque region represent 16% of the 864 reptile species recorded in Mexico (Flores-Villela and Garcta- Vazquez 2014), 52% of the 270 reptile species recorded in south-eastern Mexico (Johnson et al. 2010), and 11% of the 1284 species recorded in Mesoamerica (Johnson et al. 2018). That said, the inventory of reptile species in the Selva Zoque region is far from complete, particularly in UC zones. In the last three years alone, three new species have been described (Anolis purpuronectes [Gray et al. 2016], Chersodromus australis [Canseco-Marquez et al. 2018] and Cenaspis aenigma [Campbell et al. 2018]), and more species await formal description. Furthermore, several zones within our study area lack reptile records entirely and remain unexplored, and hence additional surveys could reveal reptile species unrecorded for the region and perhaps altogether new spe- cies to science. As such, the species richness we report here is likely an underestimate, highlighting the need for additional sampling effort to complete the species inventory of the Selva Zoque region. The high species richness in the Selva Zoque region may be the result of a series of factors. One is the notably complex orography with lowland zones, both on the Gulf of Mexico and on the Pacific versant, and also a series of mountain ranges of intermedi- ate elevation—the Sierra Atravesada, the Espinazo del Diablo, the Sierra Tres Picos, and Cerro La Colmena (Wendt 1987; Ortiz-Pérez et al. 2004; SEMARNAT 2001)—with elevations from 100 to 2300 m a.s.l. There is a range of 11 °C in mean annual tem- perature across this elevation gradient, and a difference of 3600 mm in mean annual rainfall across the Selva Zoque (Beard 1955; SEMARNAT 2001). In addition, at least seven vegetation types exist in the region (Arriaga et al. 2000b; SEMARNAT 2001). Together, these factors create a wide variety of habitats for a large diversity of reptile species with different eco-physiological requirements and evolutionary histories. Although the Selva Zoque did not result clustered with any of the regions in the similarity analysis, Los Tuxtlas and Lacandona Biosphere Reserves are the most similar tropical regions to the Selva Zoque. ‘This pattern has been observed for amphibians in a comparison between Uxpanapa-Chimalapas zone and the same tropical regions (ex- cept from La Sepultura) used in this study, and can be explained by the high number of recorded species in these three sites and the high number of species that they share. The Selva Zoque reptile fauna has a combined component of species from two differ- ent biogeographic provinces (Morrone 2005): species found in the province of Gulf of Mexico (e.g., Anolis sericeus Hallowell, Holcosus amphigrammus (Smith and Laufe), and those in the province of Mexican Pacific Coast (e.g., Loxocemus bicolor Cope, Por- thidium dunni (Hartweg and Oliver), Rhinoclemmys rubida (Cope)). The high percent- age (8%) of endemism of reptile species to the Selva Zoque region may be due to its stable Pleistocene climate conditions, which allowed for the diversification of different biological groups (Lira-Torres et al. 2012; Rodriguez-Gémez et al. 2013). A pattern Reptiles of the Selva Zoque region, Mexico 137 of high endemicity of the whole Selva Zoque region has been observed for mammals (Escalante 2003), but also in parts of Selva Zoque region like Uxpanapa-Chimalapas for other vertebrates as amphibians (Aguilar-Ldpez et al. 2016) or for Chimalapas zone in the case of birds (Peterson et al. 2003). Our results suggest that Selva Zoque is a priority conservation area for the reptile fauna of Mexico because a relatively high proportion (43%) of the species that inhabit there are in high-risk of extinction categories in the NOM-059; equivalent to 33% of all the Mexican reptile species included in this initiative (SEMARNAT 2010). Ad- ditionally, a moderate number of reptile species are in the high vulnerability category of environmental degradation (20%). In contrast, Selva Zoque does not harbor a high number of species in high-risk categories of extinction according to the IUCN. The differences in the number of species in high risk of extinction among initiatives is presumably because the NOM-059 does not consider the entire distribution of the species, it only takes into account the distribution of the species within the Mexican territory, which may lead to a restricted distribution, but only within Mexico, this does not necessarily reflect the entire distribution of the species. Additionally, populations of some reptile species that occur within Mexico may be scarce or may be declining, while populations of those species but outside of Mexico may be stable. Even so, the impor- tance of Selva Zoque region lies in the relatively high proportion (21%) of species that are classified as Not evaluated (NE) and Data Deficient (DD). Because of the conser- vation status of vegetation in some areas of the region, it represents an opportunity for gathering information on the biology of these species, and this could contribute to their being assigned a category. This is the case for Abronia bogerti Tihen, Anolis alva- rezdeltoroi Nieto Montes de Oca, Xenosaurus arboreus (Lynch and Smith), and Tanztilla briggsi Savitsky and Smith (Fig. 4A, B, E, G), endemic species to the Selva Zoque re- gion, or species with distribution in Mesoamerica as Trimorphodon biscutatus (Duméril, Bibron and Duméril), and Epictia phenops (Cope) (Fig. 4H, I). In any case, though widely distributed, some reptile species are considered threatened under three classifi- cation systems, such as Anolis pygmaeus Alvarez del Toro & Smith Bothriechis rowleyi (Bogert), Dermatemys mawii Gray and Crocodylus acutus (Cuvier) (Fig. 4C, J, K, L), all of which have been reported for the other regions (Appendix 1; Fig. 3). The extensive areas of well-preserved forest in Selva Zoque are mostly located in the Chimalapas zone and El Ocote Biosphere Reserve (Flamenco-Sandoval et al. 2007; Lira-Torres et al. 2012), and offer an opportunity not only for the conservation of rep- tile diversity, but also for all the biotic diversity that inhabits the Selva Zoque region. Conservation is under the aegis of community initiatives in Chimalapas and has been successful in recent decades (Monterrubio-Solis and Newing 2013); however, it is nec- essary to implement and subsequently strengthen conservation efforts in areas without any protection. For Uxpanapa and the intermediate zones between Uxpanapa and El Ocote Biosphere Reserve, the constant and accelerated transformation of the original vegetation cover into crops and pastureland (Flamenco-Sandoval et al. 2007; Hernan- dez et al. 2013) underscores the urgent need to stop the advance of the agricultural frontier (Arriaga et al. 2000b). 138 José Luis Aguilar-Lopez et al. / ZooKeys 1054: 127-153 (2021) - ai Figure 4. Reptile species in high extinction-risk categories by IUCN, NOM or EVS initiatives, or listed as Data Deficient or Not evaluated by the IUCN red list (see text) A Abronia bogerti B Anolis alvarezdeltoroi C Anolis pygmaeus D Lepidophyma tuxtlae E Xenosaurus arboreus F Xenosaurus rackhami G Tantilla briggsi Trimorphodon biscutatus | Epictia phenops J Bothriechis rowleyi K Dermatemys mawii and L Crocodylus acutus. The implementation of activities proven to be effective in the conservation of rep- tiles in transformed landscapes in the study region is desirable. These could include: 1) ecological restoration (Smith et al. 2015) to facilitate the recovery of the reptile com- munities over time, 2) the protection of vegetation remnants (Pulsford et al. 2017) that can harbor important components of overall species diversity, and 3) protection of buffer zones around water bodies (Semlitsch and Bodie 2003) that maintain popula- tions of several species, and also constitute biological corridors (Burbrink et al. 1999). Finally, the protection of Uxpanapa portion of Selva Zoque can contribute to the con- nection of natural protected areas of different governance types in the western (com- munity protected areas of the Chimalapas, in Oaxaca) and eastern (federal protected area El Ocote, in Chiapas) of Selva Zoque (Garcia-Bafuelos et al. 2019). This idea fits completely with the objective of the Mesoamerican Biological Corridor, an interna- Reptiles of the Selva Zoque region, Mexico 1:39 tional initiative that aims to maintain biological diversity, decrease habitat fragmenta- tion, improve the connectivity of the landscapes and of the ecosystems in Mesoamerica and to impulse social and economic development (Miller et al. 2001). Acknowledgements The authors thank A. Vinalay, C. Nochebuena, D. Aportela, L. Vazquez, L. Alcaide, J. Lépez, M. Oropeza, R. Flores, J. Pelayo, I. Caviedes, EF Vazquez, C. Hernandez, and U. Garcia for helping with the fieldwork. Policarpo Ronzén, L. Feria, and A. Vazquez provided logistical support for the trips to the field. The authors are grateful to M. Lépez-Luna for providing the photograph of Crocodylus acutus, to 1. Ahumada for providing the photograph of Bothriechis rowleyi, and to L. Badillo for providing the photograph of Epictia phenops. P. Baftuelos prepared the map and B. Delfosse and A. 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Molecular Ecology 14: 1095— 1108. https://doi.org/10.1111/j.1365-294X.2005.0247 1.x José Luis Aguilar-Lopez et al. / ZooKeys 1054: 127-153 (2021) 146 uInIpayy 41H uInIpayy uInIpsyy W3IH 41H MOT MOT wIMIpsyy uInIpsyy MOT MOT uIMIpsyy 43IH 41H 43IH 43IH 41H VN W3IH 41H SAq 243 0} Surprosor Aiyige -rouyna Jo Ar0Saye7) €1 IT SAT UIOOUOZ) ISeIT quaDyaq Req povenyeas 10N UIOOUOZ) ISeIT gyqeisuyn,, quaDyaq Beq UIOIUOT) ISPI'T UIOIUOT) ISPI'T UIOIUOT) SPOT UIOIUOT) 1SPI'T UIOIUOT) SPIT UIOOUOZ) ISeIT UIIOUOZ) ISeIT quaDyaq Req UIOOUOZ) ISeIT quaDyaq Req quaDyaq Beq quaDyaq Beq porenyead 10 porosuepuy quaroyaq wed LSTT dau NONI I Id 0 Id I I Id 0 0 Id 0 Id I Id I Id I I Id 0 Id I 0 Id 0 d 0 0 d 0 0 d 0 d 0 650 = sasOy -WON suede aH OO 2 Orns ©. 8S oo oo ococUcUWuWeMCcOlUwO ue vy EIS Se a oe oe oo a ) oo cocUcUcmlhmUlUCcUOllUO your 2PPrO VW XWVW a TAN VW XWVW XWVW oo oO 0900UCUCUCUOD a ee) SO. Oss a Oat Oe ee Or OS. re St eS Ett VW VW VW daw GTN aA OT Oo Oo -— Ont me oO ae aH Oo - OTF Oo CO aH Ont AT Oo XWVW VW IT XWVWN IT ST ZS onmroo°ococlUlcUcUlmlUr oo 7 oO tT OO oo co cocUcUcCOhUhcrrhUhcO oo ocoocUcmcWlUlcUCllUcO me OO TF OT I 0 0 0 0 ST 0 I 0 0 0 LT a8uvs uo “ngqryasiqd vinyyn = sepxny, -dageq = soy vuop sedeoyy onboz -uvdey] sey RAPS siaiag odv9 syouy JOUTWIOY], sauvisaduajnog soupy , (UUeUIZITY\ ) sugvo.0d1q syoupy , Jasus[Nog s1ayIaq syOUPy (Qpruryss) mayuvg syoupy 2 PIC AP SAIUOJ-O1DIN LoLoyapzaivayy soupy sepropApeg adory smgpssas snjgupUavT UURUIZIT YY Sadisuo] snjoupULavT » (WURUISaI\ ) 22sapupusay sauvydops0) (WIOTIOYAI) $gVIS149 saupg dousor) , UURWIZIT SNIVHIA snasyIsDg sepruvydoysso7y uuRUIsal A sujpgda20y] snJoUogLdar) - (yarurg) IVIJAZOL SNISIIY (Tjeqdurey 29 JopIayy) aupzssuz sngsajay (edoz) sSnNULUDAsvaUUa Snqsajay uOUURYS 29 JOpIay\ 7794 vIUuoLgy eqdurey zzaunups viuosqy Jpeqdureg zsvjauso viuosqy 20 op SOUOPI-OIDIN] 29 SoddY-VUNT ‘osney[D wI2uUasou VIUOLGY ITU 29 YUIG vz) vIUOLQY USUI 4.1ad0g viuoLqy sepmsuy BIPHIs9"T Japsoqns vieurenbs 1apsio VITILdda SSVTO so1sod¢ ‘(sepxny, so] ‘sedeoyy seq) saunuvig sdojgdtgopuy pur (emmyndas vq ‘sepxny soy) sma204n4 snpgovpiumazy ‘(ue ey] UkIg pur [NUTYZR]e> ‘sepxny soy ‘euopuvory ‘sedeoyy seq ‘onbo7z eayas) sugvuadf snjdqavpiuazy ‘(onbo7Z eajas) vyvjznue vidgar) ‘(1sdI0,J ULAR pue ue ey ueIs (nuryeyes ‘sepxny soy ‘onbo7 eajas) zaudvs soup “({NUyeTeD)) Supaqvyst49 SOUP :SUOTSAI Payenyead dy} UT papsodar satdods nox ‘woisds SA OY) UT posepisuos you saads = WN ‘onbo7 eajas = 79 ‘Asoo ueARPY = AY ‘sepPXny soT = [J “emmandas ey = sy] “yompyey sno Surmp popsosas saideds = v :suoneisaiqqy ‘sNI¥}S UOTIVAIISUOD puk ISULI UOTININSIp Way) “eoTIOWIOSATA UIJOYIIOU UT sUOTZaI JOYIO UdAds pue UOTsaI anbo7Z eaAjag dy} UT popsodas saidads ayndaz aaneu jo AsTT | xipuoddy 147 Reptiles of the Selva Zoque region, Mexico uIMIpsyy wIMIpsyy VN VN VN uInIpsyy MOT 43IH uIMIpsyy WH uIMIpsyy MOT MOT MOT uIMIpayy MOT VN MOT WnIpayA ystH 41H MOT 41H uInIpsyy MOT MOT 41H 41H 41H SAq 243 0} Surprosoe Aryige -rouyna Jo Ar0Sayqe7) SAD UIOIUOT) ISPI'T UIOIUOT) 1SPI'T poerenyesd ION poerenyeas ION poerenyeas ION UIOIUOT) SPIT UIOIUOT) ISPI'T UIOIUOT) ISPI'T UIOIUOT) ISBT eTqeroupnA, UIOIUOT) ISBT UIOIUOT) ISPI'T UIOIUOT) ISP9'T UIOIUOT) ISPI'T UIOIUOT) 1SPI'T UIOIUOT) ISP9'T poerenyess ION UIOIUOT) ISP9'T UIOIUOT) ISBT porosuepuy povenyeas 10N povenyeas 10N UIIOUOT) ISeI'T UIIOUOT) ISeI'T UIIDUOT) ISeIT povenyeas 10N quaDyaq Beq UIOOUOZ) ISeIT UIOIUOT) ISP9'T STITT Gaa NOOI Iq 0 Iq I 0 0 I Iq I Vv I Vv 0 Iq 0 d 0 Vv 0 Iq 0 Vv I I I I 0 0 T Iq 0 0 0 Vv 0 Vv 0 I I Iq 0 Iq 0 0 6S0 jsoIOT -WON uedepy on oe) om nt Or Oo oO oo oc oO 770 00° 0 ue vy Urls —- OOo oOo Oo Conn ornaeo$9’d oo coco oO 7 0 00 0 your oP oo Con se oo 7 oO CO CO O&O on on oOo nTr oOo oOo Oo vinjyn -dag ve] oo oO S on nr On ws CO Co On en et CO CO nH CO sepxny, SOT oo On es oOnmr Oo CON a oo oo TF oO Oo Co CO vuop = sedeoy) -uvoey] on Oo OC Corn onaeo$”’d ooo oo 7-r oO © SS seT (ae ee ee see we Oo = aS we CO er vr Ont Tr onboz PAPPS VW a PANN XWVW XWVW VW dx GTN VW VW VW VW a TANN VW VW VW VW XWVW VW VW XWVW ZS VW ZS VW VW VW IT XWVW XWVWN a8uvs uo “ngqrysiqd adory o1sy vuosouksg ey sepnewosoudryg (udnnno}) vpnvodin snjovpvoaq], opAey, SnUSDUL smypdgovpoghg aeprAoeporAyd sospoH 29 smoynepy oruRIQ sy vucoquds veosv yy sospoH 29 smoyel lURIQOI\] Yjosuy BLosLuDT seprAnqeyy » (shaeuUry) vupnst vuvnsT (Aerxy) syyzuets nenpsouary (uuewsar DN) mypunsad vinvsouaq’) (mevys) wengzuvoy vinvsouay’) (ado) sosuafap xhugov) oepruens] (UURLISII\) WnpLLLoY vULsapo]aLy sepneursapopay (J]oMoTTeH) snso1sads snugvgsqdouuls seprmyeyydouurssy , Aerxy suvsaja xhuoajoy sepreydayqny Apasan, 29 YOY sumwgopiun syouy adory szusofiun syouy sIaiag sngouopido.s syoupy (YarUIS 2g CIOL Jap zareayy) szdzqsojzds syouy » JPMOTTEH] Swaotuas soupy » MNOs0g MZans1upod sOUy 2 WNW 29 OIOL Jap zareayy snaviusd soupy » BO op sau0yy -O1DIN 29 90g ‘OrezET-eZay ‘Avy saqsauosndind syouy ynosog uséajad syoupy YUNG 29 CIOL, Jap zareapy sagvpnoutsiasvd syouy YU zvpnywu syouy , odor snutinuma soupy (WURUISaI\\) Seguaazaayy syOUPy UOSIOpUdP] 29 LIU] suvupanp syoupy yprurg sauzadns syouy , ISIN 2Q YWUIS supnvoissarduos syouy so1sod¢ 127-153 (2021) José Luis Aguilar-Lopez et al. / ZooKeys 1054 148 uInIpayy uInIpsyy uIMIpsyy VN uIMIpayy WINIpayA ysIy wINIpayA MOT uIMIpsy[ uINIpsyy wInIpsyy wInIpsyy wInIpey| uInIpsyy uIMIpsyy MOT MOT uIMNIpsy[ wIMIpsyy uInIpayy MOT uIMIpsyy 43IH umMIpayy MOT W3IH MOT W3IH uInIpayy 43IH uIMIpayy uIMIpsyy SAq 243 0} Surprosse Aiyige -rouyna Jo Ar0Saye7) SAT povenyeas 10N povenyeas 10N UIOOUOZ) ISeIT quapyaq Beq povenyeas 10N UIOOUOZ) ISeIT UIIOUOT) ISeIT UIOOUOZ) ISeI'T UIIOUOZ) ISeIT UIOIUOT) ISPIT UIOIUOT) SPIT UIOIUOT) SPIT UIOIUOT) SPIT UIOIUOT) 1SP9'T UIOIUOT) ISBT UIOIUOT) SPIT UIOIUOT) 1SPI'T UIOIUOT) SPIT UIOIUOT) SPIT UIOIUOT) 1SP9'T UIOOUOZ) ISeIT UIIOUOZ) ISeIT UIOOUOZ) ISeI'T UIOOUOZ) ISeI'T UIOOUOZ) ISeIT UIIOUOT) ISeI'T quaDyaq eeq UIOOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOT) ISeI'T UIOOUOT) ISeI'T UIOOUOZ) ISeI'T UIOIUOT) ISP9'T STITT Gada NONI me OOF CO OF TF Oo 4d I 4d 0 = se Or OF CO OS 4d — >) Yn uINIps O1 oO aia ~ Qa te) n 431H LI = wINIpsy| IT pas) S 4st LI lad wINIpsy IT MOT 8 uINIpsy €1 VN VN 431H 91 MOT 8 wINIpsy €1 SAq op? 0} Surprosoe Aryige -rouyna Jo Ar0Saye7) SAH UIIOUOT) ISeIT UIOOUOT) ISeI'T quaDyaq Bed UIOOUOZ) ISeIT UIOIUOT) ISP9'T UIOIUOT) ISP9'T UIOIUOT) ISBT UIOIUOT) 1SP9'T UIOIUOT) ISPI'T UIOIUOT) ISPI'T UIOIUOT) SPIT UIOIUOT) 1SP9'T poerenyeasd ION poerenyesd ION UIOIUOT) ISPI'T UIOIUOT) SPOT UIOIUOT) SPIT qusDyaq Beq UIOIUOT) SPIT poerenyesd ION poerenyess ION poerenyesd ION quaDyaq Beq UIOOUOZ) ISeIT UIOIUOT) SPIT peueievoty TRON] porosuepuy UIOIUOT) 1SP9'T poerenyead ION STITT Gada NONI Id 0 0 0 i) I I 0 T T I 0 T 0 0 i) 0 I Id 0 T 0 0 0 V 0 Id 0 Id 0 I V 0 Id I 0 650 =: sasOy -WON) uede oo Co. 0oO0UcrrhUhcCOUhUcOO - OOo Oo FF oo o9c0[UcUCDUlUOD i ue ey Urls Oe) m- OOo Oo oo oO fC Oo fo oo o090c[UcWmClhUOD your oP Con ne nH OH CO oe 6 oo = S&S ame OO fo CG oo o.cmUcoOoO FT Oo vinjyn -dag ve] Se ee ee ee oe > on EE a a) Co ot et Oo OS ooonr Oo sepxny, SOT COR He HH Oo we =a OF Oo Oo & om oO Tt Oo Oo i a) -uvoey] Onn wnwnoo oo oc Oo S&S Co Om Oo oOo... = COCOOoOn Oo evuop = sedvoy) sey Conon wnnwr Oo SS Det cS: aS or oOo fF = = Ss oe ON ON onboz PAPPS VW VW IT VW VW VW VW a TAN da VW VW a TAN ZS ZS VW VW VW ST a TAN IT VW ZS XWVWN VW XWVW VW ZS VW VW a8uvs uo “ngqrysiqd (adox)) sazavfiaasg susdigr siaiag snqvyqia stqdouo’) yprurg 29 vunq-zadoyq “eparesipy-z019qg /2m10u siqdouor ([HpuNG 2 UoOIqig ‘fuguUINC) smsvaur srqdouoy) Aaqreg mpuugas sauvgdo1u0y (pwnd 2 woIqig ‘fupuINd) sasvyyzaanbuinb sauvgdo1u0) adory sugzataard sauvgdoiu0y » (PIED 29 preg) syvisadut sauvgdoiuo0’) » GayIUN) suaprssef sauvgdoruoy (oyun) sagvyoundig sauvgdoruoy 2 (adod) vuyuros vy any (uIpneq]) 7749 749) Jeqdures 29g zayezuo0ry -zaryurey ‘Zanbieyy-ooasury synsny snUuLospossay’) IPH 29 Yrus qpeqduresy veuszuan srdspuary » (ouIay\) waddys wnipusisvupy (adoy) wnuzuosta songdapy uel wngvdizatponb songdapy ypu 29 youd] wngvrasufiny songdapy seprpesdiq urpneg “ojmadus vog aeplog sayuedsag Joproqns UOUURYS 29 JO[IAN SISUIUTZAVULUDS snAanvsoury wenig UuyYyIvL snanvsoUuay (yarurs 2g youd) snasoguy snanvsouay aeplimesousyx , UOUURYS 29 JOON avpexng vuutydoprdaT ynosog sg42us viudgdopidaT 2 Japa sisuauvdylod vuukgdopidaT Azag avkvu vudgdopidaT O10, [Op Zorearyy 2 ys z27ady viudgdopidaT Jpeung wnyyjnsvunanpy, vucdgydoprdaT oeprisnjyuey (2[GON 29 Inogseg) snasd snsoopy so1sed¢ 127-153 (2021) José Luis Aguilar-Lopez et al. / ZooKeys 1054 150 uINIpsy[ VN uIMIpsyy wIMIpsyy MOT wInIpsyy wIMIpsyy uInIpsyy uIMIpsyy MOT 41H uInIpayy uInIpsyy VN wInIpsyy MOT wIMIpsyy 48tH MOT MOT wIMIpsyy MOT MOT MOT MOT uINIpsyy MOT uINIpsyy MOT MOT 43IH uInIpsyy wmnIpey MOT MOT SAd 243 0} Surprosoe Aiyige -rouyna Jo Ar0Saye7) SAT UIOIUOT) 1SP9'T poerenyeas ION UIOIUOT) SPIT UIOOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOZ) ISeI'T UIOOUOT) ISeIT UIOOUOZ) ISeIT UIIOUOZ) ISeIT quaDyaq Beq UIOOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOZ) ISeI'T qusDyaq Beq UIOOUOZ) ISeIT UIIOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOZ) ISeIT UIOOUOT) ISeI'T povenyeas 10N UIIOUOZ) ISeIT UIOOUOZ) ISeI'T UIOOUOZ) ISeIT UIONUOZ) ISeI'T UIOOUOZ) ISeI'T UIOOUOT) ISeIT UIOOUOZ) ISeI'T povenyeas 10N UIOOUOT) ISeIT gyqeisuynA, UIOOUOZ) ISeI'T UIOIUOT) ISP9'T LSTT daa NONI I I V I I Id I 0 0 i! I I 0 I 0 I Id 0 I I I I I 0 0 I 0 Id 0 i! Id 0 Id 0 Id I Id I 0 Id 0 0 0 0 650 =: sasoy -WON) uede Oo ooo oO oO nr TO OoOUWUOmUcrTCUCOUCOUCcOUCcCOUCcOUCUcOcrTW CLC OoOCUcrrFWCOOCUCOCUCOUCcOUCcCCOUCc CWO oO nr Tr oO OT Oo ue ey els oo coco 0. OFF yr or ocmUmUDCUCUODUCUWTCOUcUrWDCUCUCUOOCUCcrMCOCUCOUCmcWUCLcCOUcUODUCcUOULURDCUcUTDCLCUC OURMCUCOOUcUMLhUc your oro Oo moo cocl.o Fr OTF OTF OoOOCOoOlUmUrrDrCmUTVCOOCUCcCcOCUCUcCOUOmUrrTCOCUCcCOCUCcOCUCcOCUCcOUCcOUCcOUUcOUcUOUcrTCUCcCOUCOhUhO Ca as ee ee! eingyn sepxny -dageq = soy Oo eto Oo. Oo mt OOo ot Oo OS OL! oS ee Ot OOO OO Ot OO OO oe -uvdey] ooo oO OF COCO OF TF OO Tr OOO cUrTrthUcUTIChUcTrCLUC OCUcrrTFrCOOCUCOUCcCOcUCOUCcCOCUCcOUUCcOUUcOCUcUOUcUOULUMLhULc vuop = sedeoys) sey Ce es eo a a a i! onboz RAPS VW VW daw a TANN VW VW a TAN VW VW a TANN IT VW VW VW ZS ETN da GTN VW VW XWVW dx VW VW VW VW GTA XWVWN dx a TANN ZS XWVW LT VW EPA a8uvs uo “ngqrsiqd » Wg Logzu7a uorpiqdospuaq adearg 29 3[peD wnsourusagyns uompigdospuaq SNIBUUTT 40792445U09 4aqn] 0’) sepliqnyo’) (pormnan-paty) smvqgdasopqus uopouay adory ms0avs sosdipopidosy (ro8uatnog) zaqasyf svsdipoprdosy rayluny wyviasuf svsdipoprdosy ado snaqnjoss1u snurgsounqasy (ado-)) smporuuns uogis (snoeuurT) snqvjngau uogig ypu 29 vunq-zodoyq ‘eporesipy-z019q szvauy uogis e (JOYIUN) saqvipiuip uogis (IoyUND) suvupos vyauipoyy (Teqdures 29 yaTUIS) Piasogavur DyaUIpLgYy pueqosury] 29 yNUIS 7/VsnopsvU vavupYYy » (OYUN) vynL0Iap vavuipyyy ador saprodyja snosa20 , (snavuurq) saievjoqad sndogslxeE » (Heung 29 voIqrg ‘qgumNd) avgas viuIyy » PAeIIL) 29 plleg wywwapvip viUuI\ (uel) zuvugnd sidajouvpy (nosruusy) syvuorsuasdas vatapoqdaT » (IaUN) MousKod vatapordaT roylunty wyvisvfoss1u varapordaT (]emoTfeH) vwjnavut vazapoqdaT (adoz) vipuaLf patapordaT (snoeuury) yywjnuup vuapordaT (ador)) snwtssznuay sapoqupuy (edor) SNIVAISIULUID SOPOJUDUT » (SNovUUTT) Yogouas sapoquvUT , ‘Aou ‘ds stqdoar SUPT 29 YIUIS sujouz92q07 stg doar) vuny-zodoyq 29 yiurg ‘eparesipy-z019g svyynl srqdoar) qrenig susouzsva siqdoary (ado) sanbsogiamyl snynug so1ood¢ 151 Reptiles of the Selva Zoque region, Mexico wIMIpsy[ MOT uInIpayy uInIpayy MOT VN ystH MOT MOT VN uni psy/\y VN ystH ystH 4st MOT MOT MOT MOT W3IH uInIpayy MOT uIMIpeyy MOT VN MOT 481H uIMIpsyy MOT MOT MOT 41H MOT MOT MOT MOT SAd 243 0} Surprosoe Aiyige -rouyna Jo Ar0Saye7) OI SAT UIOIUOT) 1SP9'T povenyeas 10N UIOOUOZ) ISeIT UIOOUOT) ISeIT UIOOUOZ) ISeIT qusDyaq Beq quaDyaq Be UIOOUOT) ISeIT UIOOUOZ) ISeI'T UIOOUOZ) ISeIT gyqeisun,, UIIOUOZ) ISeIT quaDyaq Beq UIOOUOZ) ISeIT UIIOUOZ) ISeI'T UIOOUOZ) ISeI'T UIIOUOT) ISeI'T UIIOUOZ) ISeIT UIIOUOZ) ISeIT UIIOUOZ) ISeI'T UIIOUOT) ISeI'T UIOOUOZ) ISeIT UIIOUOZ) ISeIT UIIOUOZ) ISeI'T povenyeas 10N UIIOUOZ) ISeIT UIIOUOT) ISeI'T UIIOUOZ) ISeIT UIOOUOZ) ISeI'T UIOOUOZ) ISeI'T UIOOUOZ) ISeIT quspyaq Beq UIOOUOZ) ISeI'T UIIOUOZ) ISeI'T UIIOUOZ) ISeI'T UIOIUOT) 1SP9'T STITT Gaa NOOI 0 Vv 0 Iq I I Iq 0 T Iq 0 T Iq 0 I 0 Id I Vv 0 Iq 0 Iq 0 I i! I I Iq 0 i! 0 i! I i! Vv I Vv 0 Vv I i! I Vv I 0 i! I 0 I 6S0 jsoIOT -WON_ uedeyy a OO ce Oo Oo mt Oe Oo ost Om OO CO Oo oO COO oH OC OO Ooo Ol O.UoO!. lO Ol O.CU-O ue ey els m= Onmr nr OO FTF Tr OTR Tr rT OTF Or nr oOo oOloOomcrVCOCOCUCOCcCOUmcCUCcCOCcCUCOUCUcCCOCUCUCcCOUCcCOCcOoOUlULrrtrLhULcrT LCcO your oro =S Ss we CO COCO FB ae et OR eR eR eR OO RR OR Oe Or CUCU OlUlCUlcrClUlUcrrCOUCOOUC OUCUlcrFlLUCUC;OCUMrOFOlCUC)OUOCOUCUCOUC rrOrlChUCOUCOlUClCrr eingyn sepxny -dag ve] SS Se SS SS SS Se On eK KS eS OF Ow wrr oo oO Cc Cc CO CO Oo Se St CO Co fo eS eK soT = Oe = OC Se Se wt we OCOwnrwrt rw wv Ot CO wee wr we OO COCO CO COO OT OCOUCCO CU OlClUCOlUrK CO vuop = sedeoys) -uvdey] meorwrnmt Onnrn ocrcoc coc coco eo ceo co oe ee ololo Ol OOO OO OS seT SS SS Ot Ut COON Ur Ur Ur COCO UN UN UN Ur rr rs rs rs cr Cc re DC OCUrUCcU ND rvwWrCOCUCOUUCcCOU Ur rRHCOUC OUCUCcOUC COU COUCH onboz RAPS dx VW VW VW VW dW IT VW VW VW VW VW ZS VW XWVWN VW EVAN dé q&a VW VW VW VW GTN XWVW GTN XWVWN GTN VW da da XWVWN VW da VW EPA a8uvs uo “ngqrsiqd (Tesoypep]) “fiquiogs vyposayy seppiney ({HpuING 2 WoIqIg ‘fupuUIN) smqvjnIs1qg Uopoydsoutsy 2 (WATWIS) suoruyy vIn pIULT (adod) yynuva vinytuny (JoJARL) vuutsstaasg vINTIUYT wrung 2g yoqdured visa1 vypsauny YU 29 YU “eporeSTPY-za19q ssUaay]s YIPLIUYT, (aIN090g) YsOIS1GIS VIITAUYT. adog magne vIIUDT (JOWIUN)) YpszOU VIPLIUYT, (oyun) sunl yyeuny YAUIS voyjnIIUNI HYLUYT yarurg 29 Ayzararg 199524q ypsUYT (a8) avdvut snug duds adory uLnuLoysomna snug duds ({HpuING 2 WoIqig ‘JupuUIN) sayprauzmadf vuLgLsouats v (PJOYIq) IIpavgyuasap DULGALOUITS » (snavuurT) snypynd sagoqtds (ado>) szdsviag stpoatquas (adod) vivasruma, viopvayys (ado) vfnranyf agdvjapnasy (edo) syposrsvauy srqdonaig (oyun) sngouop1sa0d xDUoUllgg (urpneq) smpi5ynf syaqtxO (r0]ke) sesuatsogod syaqhxC JHeung 29 UoOIqig ‘uguNg snuvoixau siqdoydaT (yun) szdoopdip srqdordaT , (snavuury) vymavgn stqdordaT (jmno0s0g) vuouqv styadosquvT v (odor) SNULOJOUIAUL svdLposUIsv yy ([HpuNG 2 woIqig ‘JupuUING) smzuvaoquau s1qdosusvy (OYUN) wywsatava vIUuIILT adog vygnd viuz1s17 (eSa[YyOS) sauafinsvdivu sniqouhiq (adoa) snazzos0,49 snigowdaq 2 (pun 29 voiqrg [lIpuING) smunuvjau uogosvuLdaq so1ood¢ 127-153 (2021) José Luis Aguilar-Lopez et al. / ZooKeys 1054 152 41H uInIpsyy VN umMIpe| VN VN uIMIpayy uIMIpayy uIMIpsyy 41H 43IH wIMIpsyy uImMIpsyy uImMIpsyy wIMIpsyy wIMIpsy[ MOT W3IH uIMnIpsyy MOT MOT VN uIMIpsyy MOT uIMIpsyy MOT SAq 243 0} Surprosoe Aryige -rouyna Jo Ar0Sayqe7) SI cl VN IT VN Il OI SAD UIOIUOT) ISP9'T UIOIUOT) SPIT UIOIUOT) ISPI'T UIOIUOT) ISBT poerenyesd ION poerenyeas ION UIOIUOT) SPIT poerenyeas ION UIOIUOT) 1SP9'T eyTqedouyn, lA UIOIUOT) 1SP9'T peueievoty TRON] UIOIUOT) ISP9'T UIOIUOT) ISP9'T UIOIUOT) 1SP9'T poerenyess 1ON perenyead ION UIOIUOT) SPIT UIOIUOT) ISPI'T UIOIUOT) ISP9'T UIOIUOT) ISPI'T poerenyeas ION UIOIUOT) SPIT UIOIUOT) 1SP9'T UIOIUOT) ISPI'T UIOIUOT) ISPI'T LSTT daa NONI Vv 0 I I 0 0 0 0 I I Iq 0 Vv 0 Iq I 0 I Iq 0 0 0 Iq 0 Iq 0 Iq 0 Iq 0 T I Vv I Vv 0 Vv 0 650 ——-sas0g -WON uedepy ont Oo Oo OSs Ou eh O02 a ) ue vy Urls ont Oo oO mw OO olcUrTCLUCOCUDO —a OOo oOo Oo your oP ooo Oo =a oe CO CO wm OO om oo Oo vinjyn -dag ve] oo oOo oo Coc ONTO CO oon TW sepxny, SOT -— Oo nr © oo ont TF OO a SS. tS vuop = sedeoy -uvoey] ooo Oo oo Cc On oO CO Oe St -S seT ooo Co OF Ne eS SS (a I ee oe onboz PAPPS VW VW XWVW VW IT XWVW VW a TANN a TAN XWVW VW VW VW VW VW XWVW VW IT VW VW VW XWVW dx GTA ETN EPA a8uvs uo “ngqrysiqd (eY~pNIIITZ -eI[n{ 29 YTUIG “eporeSIEY-z9I9q) Jaujo smpyvomidypayy (fpuING 2g WoIqig ‘[IgUING) smuvoixau smpvomdyay Jaqnepy] uvrgvz4 snjyjoLy apporae’] smueis snqygody pqRyW -IMOYYep] 29 ONsed-HIN ‘sory-zouyepy