A summary of the species of amphibians and reptiles of the state has been compiled, including their geographic distributions, habitats, and conservation statuses. The herpetofauna of San Luis Potosí consists of 41 species of amphibians and 141 species of reptiles. San Luis Potosí shares the highest number of species with Hidalgo and Tamaulipas, and the least number of species with Nuevo León. In San Luis Potosí, there are several taxa of particular conservation concern including salamanders, emydid and trionychid turtles, anguid and xenosaurid lizards, and natricid and colubrid snakes.

Checklist, Chihuahuan Desert, conservation status, herpetofauna, shared species, Sierra Madre Oriental

San Luis Potosí is a relatively small state (surface area = 63,068 km², 3.1% of the surface area of Mexico) located in the north-central part of Mexico, between 24°29' and 21°10'N and 98°20' and 102°18'W (see Figure 1; INEGI 2009). The climate of San Luis Potosí varies from the temperate, dry high plains to the warm, relatively humid coast (Lemos-Espinal and Dixon 2013). Several distinctive habitats are found within the boundaries of the state, including the Chihuahuan Desert in the western half and tropical perennial forests in the southeastern portion (= Huasteca Potosina). Three physiographic provinces that vary in their temperature and the moisture retention of their soils (INEGI 2009; Lemos-Espinal and Dixon 2013) are found in San Luis Potosí: the Sierra Madre Oriental, the North Gulf Coastal Plains, and the Central Plateau (Figure 1). The Tropic of Cancer crosses the northern part of the state, and to the east San Luis Potosí nearly reaches the Gulf of Mexico to the east. The elevation of San Luis Potosí varies from about 50 m above sea level to about 3,180 m in Cerro Grande (INEGI 2009). The variation in climate (Figure 2) and physiography of the state have created a mosaic of habitat and vegetation types in San Luis Potosí (Figure 3) that most likely affect the distribution and presence of amphibians and reptiles in the state (see Lemos-Espinal and Dixon 2013 for detailed description of these habitats and vegetation types).

Our understanding of the herpetofauna of San Luis Potosí still remains somewhat limited (see Lemos-Espinal and Dixon 2013 for a review of previous herpetological studies in San Luis Potosí). Our intent with this paper is to encourage others to continue studying the herpetofauna of the state by providing a summary of the species of amphibians and reptiles of the state, their geographic distributions, habitat, and conservation status. By placing all this information into one, easily accessible place, we hope to provide a starting place for further research on the herpetofauna of San Luis Potosí. In addition, a comparison of the amphibian and reptile species lists to those in the neighboring states is provided in an effort to identify unique aspects of the herpetofauna of San Luis Potosí, as well as shared species, with the aim to understand the potential conservation or management needs at the state or regional level.

Figure 1. 

Topographical map with physiographic provinces of the state of San Luis Potosí, Mexico. The thicker black lines delineate the major habitat types found in San Luis Potosí (from west to east): Central Plateau, Sierra Madre Oriental, and North Gulf Coastal Plains (INEGI 2009). Maps modified from Cervantes-Zamora et al. (1990); http://www.gifex.com/fullsize/2009-09-17-3/Mapa-de-Amrica.html; García E – Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (2008).

Figure 2. 

Climate map of the state of San Luis Potosí, Mexico (modified from García – CONABIO 1998).

Figure 3. 

Vegetation type map of the state of San Luis Potosí, Mexico (modified from Dirección General de Geografía – INEGI 2005).

This list of amphibians and reptiles of the state of San Luis Potosí was compiled from the following sources: (1) our own field work; (2) specimens from the Laboratorio de Ecología – UBIPRO (LEUBIPRO) collections; (3) a thorough examination of the available literature on amphibians and reptiles of the state; and (4) databases from the Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (National Commission for the Understanding and Use of Biodiversity; CONABIO), including records from the following 30 collections:

AMNH Collection of Herpetology, Herpetology Department, American Museum of Natural History

ANSP Collection of Herpetology, Herpetology Department, Academy of Natural Sciences of Philadelphia

BMNH Collection of Herpetology, Zoology Department, The Natural History Museum, British Museum (Natural History)

CAS Collection of Herpetology, Herpetology Department, California Academy of Sciences

CMNH Collection of Herpetology, Amphibians and Reptiles Section, Carnegie Museum of Natural History – Pittsburgh

CNAR Colección Nacional de Anfibios y Reptiles, Instituto de Biología UNAM

EALC Ernest A. Liner Collection of Herpetology

ENCB Colección Herpetológica, Departamento de Zoología, Escuela Nacional de Ciencias Biológicas

ENEPI Colección Herpetológica, Departamento de Biología Experimental, Escuela Nacional de Estudios Profesionales, Unidad Iztacala, UNAM

FMNH Division of Amphibians and Reptiles, Field Museum of Natural History

FSM-UF Collection of Herpetology, Florida State Museum, University of Florida

FWMSH Fort Worth Museum of Sciences and History

LACM Collection of Herpetology, Herpetology Section, Natural History Museum of Los Angeles County

LSUMZ Collection of Herpetology, Museum of Zoology, Biological Sciences Division, Louisiana State University

MCZ Collection of Herpetology, Museum of Comparative Zoology, Harvard University Cambridge

MNHUK Museum of Natural History, Division of Herpetology, University of Kansas

MZFC-UNAM Colección Herpetológica, Museo de Zoología “Alfonso L. Herrera”, Facultad de Ciencias UNAM

MVZ Collection of Herpetology, Museum of Vertebrate Zoology, Division of Biological Sciences, University of California Berkeley

SDNHM Collection of Herpetology, Herpetology Department, San Diego Natural History Museum

TCWC Collection of Herpetology, Texas Cooperative Wildlife Collection, Texas A&M University

TNHC Collection of Herpetology, Texas Natural History Collection, University of Texas Austin

TU Collection of Herpetology, Biology Department, Tulane University, New Orleans

UAZ Amphibians and Reptiles Collection, University of Arizona

UCM Collection of Herpetology, University of Colorado Museum

UIMNH Collection of Herpetology, University of Illinois Museum of Natural History

UIUC Collection of Herpetology, Museum of Natural History, University of Illinois at Urbana-Champaign

UMMZ Collection of Herpetology, Museum of Zoology, University of Michigan Ann Arbor

USNM Collection of Herpetology, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution

UTAMM Merriam Museum, University of Texas Arlington

UTEP Collection of Herpetology, Laboratory of Environmental Biology, Biological Sciences Department, University of Texas - El Paso

Amphibian names follow Frost (2017) and AmphibiaWeb (2017) (http://amphibiaweb.org) and reptile names follow Uetz and Hošek (2017). Species were included in the list if they had confirmed records, either by direct observation or through documented museum records or vouchers. Species accumulation curves were created for the total herpetofauna, amphibians, and reptiles using the year of the first recorded observation for each species. Such species accumulation curves are likely to serve as good estimators of the potential species richness of amphibians and reptiles (see Raxworthy et al. 2012). In addition, the conservation status of each species was recorded based on three sources: 1) the IUCN Red List 2017; 2) Environmental Vulnerability Scores from Wilson et al. (2013a,b) and Johnson et al. (2015); 3) listing in SEMARNAT (2010).

The number of overlapping species with those neighboring states for which a recent checklist exists (Hidalgo: Lemos-Espinal and Smith 2015, Lemos-Espinal and Dixon 2016; Nuevo León: Lemos-Espinal et al. 2016; Tamaulipas: Farr 2015, Terán-Juárez et al. 2016; Querétaro: Dixon and Lemos-Espinal 2010) was determined, and hierarchical clustering analyses with single linkage and Euclidean distances using Systat 13 software (SYSTAT software, Chicago, IL) used to examine the similarities among the herpetofaunas of San Luis Potosí and its neighboring states (see Enderson et al. 2009 for a similar analysis). Lists were updated for Hidalgo (substituting Lampropeltis polyzona for L. triangulum, Ruane et al. 2014, Uetz and Hošek 2017); Nuevo León (adding Chiropterotriton miquihuanus, Campbell et al. 2014; Crotalus morulus, Bryson et al. 2014, Uetz and Hošek 2017; and substituting Lampropeltis annulata for L. triangulum, Ruane et al. 2014, Uetz and Hošek 2017); Querétaro (substituting substituting L. annulata for L. triangulum, Ruane et al. 2014, Uetz and Hošek 2017; adding Amastridium sapperi, Calzada-Arciniega 2014). The neighboring states of Guanajuato, Veracruz, and Zacatecas do not have recent, comprehensive checklists of amphibians and reptiles available so were not included in comparisons.

San Luis Potosí is home to 182 species of amphibians and reptiles which represent 33 families and 98 genera (Table 1). These include 41 species of amphibians (six salamanders, 35 anurans) and 141 of reptiles (one crocodilian, seven turtles, 48 lizards, 85 snakes). The herpetofaunal account for the state published by Lemos-Espinal and Dixon (2013) listed a total of 181 species of amphibians and reptiles, including Dennis’ Chirping Frog (Eleutherodactylus dennisi), a species not included in this paper since the only record for this species (ENCB-14250 – E. dennisi; collected on August 13^th, 1989. 1 km N of Apetzco, municipality of Xilitla, SLP), seems to be a misidentified E. longipes. Amastridium sapperi was added based on Calzada-Arciniega (2014). Sceloporus cowlesi was substituted for S. consobrinus based on Leaché (pers. comm.), Scincella silvicola for S. caudaequinae based on Linkem et al. (2011) and Uetz and Hošek (2017), and Holcosus undulatus for H. amphigrammus based on Meza-Lázaro and Nieto-Montes de Oca (2015). Lampropeltis triangulum was substituted for L. annulata and L. polyzona based on Ruane et al. (2014) and Uetz and Hošek (2017). Xenosaurus newmanorum was regarded as endemic to Mexico but not to San Luis Potosí based on Nieto Montes de Oca et al. (2017). No species is endemic to the state, and four are introduced: the American Bullfrog (Rana catesbeiana), the Common Four-clawed Gecko (Gehyra mutilata), the Common House Gecko (Hemidactylus frenatus), and the Mediterranean House Gecko (H. turcicus).

A list of 17 species (nine amphibians, eight reptiles) potentially occurring in San Luis Potosí was compiled (Table 2), based on species for which undocumented observations in San Luis Potosí exist but for which museum or other records are not available, and on species that have not been recorded or observed in the state, but whose distributional ranges come close to the borders of San Luis Potosí.

The species accumulation curves for all species, amphibians, and reptiles suggest that the current list of species is close to being the likely species richness for San Luis Potosí (Figure 4). These curves show a dramatic increase in documents herpetofaunal species during the 1940’s and 1950’s, primarily associated with the work of Edward Taylor and Hobart Smith (Smith 1939, Smith and Taylor 1945, 1950; Taylor 1949, 1950, 1952, 1953). Taken together with the relatively limited number of potential additions to the herpetofauna of San Luis Potosí (see Table 2), it seems likely that, barring the discovery of multiple cryptic species, that this is a fairly complete list of the herpetofauna of San Luis Potosí.

Figure 4. 

Species accumulation curves for the total herpetofauna, amphibians, and reptiles from San Luis Potosí. Horizontal lines are estimated asymptotes for the species accumulation curves.

Comparisons with neighboring states

Overall, San Luis Potosí shares the most species with Hidalgo and Tamaulipas, and shares the least number of species with Nuevo León (Table 4). It is particularly interesting to note that for most taxa Hidalgo shares the highest proportion of species with San Luis Potosí whereas Tamaulipas shares the highest proportion of lizard species by a large margin and Nuevo León shares a very high proportion (≈ 90%) of phrynosomatid species (Table 4). It is likely that this reflects the more arid nature of Tamaulipas and Nuevo León (i.e., Chihuahuan Desert), compared to the more mountainous Querétaro and Hidalgo. Thus, the numbers and types of shared species among San Luis Potosí and its neighboring states reflects the pattern of habitat and vegetation types found in each neighboring state (see also Smith and Lemos-Espinal 2015, Lemos-Espinal and Smith 2016, Lemos-Espinal et al. 2017). However, the results of the cluster analysis are somewhat different. The cluster analysis found that San Luis Potosí is clustered with the pair of Hidalgo and Querétaro for all species together and reptiles (Figure 5A). In contrast, for amphibians San Luis Potosí clusters with Querétaro, and this pair clusters with the pair of Nuevo León and Hidalgo (Figure 5B). Thus, it appears that amphibians and reptiles show different affinities among these states, again perhaps reflecting the available habitats or environments in each state. It therefore appears that in addition to state-specific conservation and management plans, more integrated habitat specific conservation plans that allow inter-state efforts would be the best approach to preserve the herpetofauna of San Luis Potosí and its neighboring states. In addition, the results of the cluster analysis suggest that amphibians and reptiles will each require different inter-state collaborations (i.e., the states involved in such collaborations might differ between amphibians and reptiles based on the different patterns of clustering between these taxa).

Figure 5. 

Results of cluster analysis of the herpetofaunas of San Luis Potosí and its neighboring states (Hidalgo, Nuevo León, Querétaro, and Tamaulipas). The distances provided are Euclidean distances for A the entire herpetofauna and reptiles only, respectively and B amphibians only.

Table 4.

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Summary of the numbers of species shared between San Luis Potosí and neighboring Mexican states (not including introduced species). The percent of species from San Luis Potosí shared by a neighboring state are given in parentheses. – indicates either San Luis Potosí or the neighboring state has no species in the taxonomic group, thus no value for shared species is provided.

Taxa	San Luis Potosí	Hidalgo	Querétaro	Nuevo León	Tamaulipas
Class Amphibia	40	35 (87.5)	26 (65)	17 (42.5)	29 (72.5)
Order Caudata	6	5 (83.3)	4 (66.7)	0 (0)	4 (66.7)
Ambystomatidae	1	1 (100)	1 (100)	0 (0)	–
Plethodontidae	4	3 (75)	3 (75)	0 (0)	3 (75)
Salamandridae	1	1 (100)	–	–	1 (100)
Order Anura	34	30 (88.2)	22 (64.7)	17 (52)	25 (73.5)
Bufonidae	6	4 (67)	4 (67)	5 (83.3)	5 (83.3)
Craugastoridae	3	3 (100)	2 (67)	1 (33)	2 (67)
Eleutherodactylidae	5	3 (60)	3 (60)	3 (60)	4 (80)
Hylidae	9	9 (100)	6 (67)	2 (22.2)	5 (55.6)
Leptodactylidae	2	2 (100)	–	1 (50)	2 (100)
Microhylidae	2	1 (50)	1 (50)	2 (100)	2 (100)
Ranidae	4	4 (100)	3 (75)	1 (25)	2 (50)
Rhinophrynidae	1	1 (100)	1 (100)	–	1 (100)
Scaphiopodidae	2	2 (100)	2 (100)	2 (100)	2 (100)
Class Reptilia	138	98 (71.0)	92 (66.7)	75 (54.3)	100 (72.5)
Order Crocodylia	1	1 (100)	0 (0)	0 (0)	1 (100)
Crocodylidae	1	1 (100)	–	–	1 (100)
Order Testudines	7	5 (71.4)	3 (42.8)	2 (28.6)	5 (71.4)
Emydidae	2	1 (50)	–	0 (0)	1 (50)
Kinosternidae	4	4 (100)	3 (75)	1 (25)	3 (75)
Trionychidae	1	–	–	1 (100)	1 (100)
Order Squamata	130	92 (70.8)	73 (56.2)	73 (56.2)	94 (72.3)
Suborder Lacertilia	45	25 (55.6)	22 (48.9)	27 (60.0)	36 (80.0)
Anguidae	5	3 (60)	1 (20)	2 (40)	5 (100)
Corytophanidae	2	1 (50)	1 (50)	–	1 (50)
Crotaphytidae	1	–	–	1 (100)	1 (100)
Dactyloidae	2	2 (100)	1 (50)	–	1 (50)
Dibamidae	1	1 (100)	1 (100)	–	1 (100)
Eublepharidae	1	–	–	–	–
Iguanidae	1	1 (100)	–	–	1 (100)
Phrynosomatidae	19	9 (47.4)	10 (52.6)	17 (89.5)	15 (78.9)
Scincidae	5	3 (60)	3 (60)	4 (80)	5 (100)
Teiidae	3	2 (66.7)	2 (66.7)	2 (66.7)	3 (100)
Xantusiidae	4	2 (50)	3 (75)	1 (25)	2 (50)
Xenosauridae	1	1 (100)	0 (0)	–	1 (100)
Suborder Serpentes	85	67 (78.8)	51 (60.0)	46 (54.1)	58 (68.2)
Boidae	1	1 (100)	1 (100)	–	1 (100)
Colubridae	36	24 (66.7)	22 (61.1)	23 (63.9)	27 (75)
Dipsadidae	22	19 (86.4)	13 (59.1)	6 (27.3)	14 (63.6)
Elapidae	1	1 (100)	1 (100)	1 (100)	1 (100)
Leptotyphlopidae	3	2 (66.7)	1 (33.3)	2 (66.7)	2 (66.7)
Natricidae	12	12 (100)	7 (58.3)	7 (58.3)	6 (50)
Viperidae	10	8 (80)	6 (60)	7 (70)	7 (70)
TOTAL	178	133 (74.8)	118 (66.3)	92 (51.7)	129 (72.5)

Conservation status

Nearly 82% of the amphibians and reptile species that have been evaluated by the IUCN falls in the Least Concern category (does not include DD species; Table 5, Figure 6). However, only 60% are not listed by SEMARNAT (Table 5, Figure 6). The discrepancy between the IUCN and SEMARNAT listings are greater for reptiles than amphibians (Figure 6). The average EVS for all herpetofaunal species in San Luis Potosí that have been evaluated is 10.67. These overall numbers tend to obscure the presence of particular groups, or even individual species, that occur in San Luis Potosí that are at potential risk and that may warrant special attention. Some taxa of particular concern, based on their IUCN listing, SEMARNAT category, or their EVS include the salamanders in general, and Plethodontidae and Salamandridae in particular; the emydid and trionychid turtles, anguid and xenosaurid lizards, and natricid and colubrid snakes. These taxa reflect assessments at the global or country-level scale. It may be, and indeed it is likely, that there are multiple species of amphibians and reptiles that are more or less threatened at the state level than these larger scale assessments suggest. However, given the relative paucity of population-level studies and assessments on the herpetofauna of San Luis Potosí, such conservation or management needs are unknown.

Figure 6. 

Percent of A amphibians and B reptiles listed in protected categories on the IUCN Red List and SEMARNAT. Green is percent in Least Concern (IUCN) or Not Listed (SEMARNAT), Red is percent in protected categories. N is the total number of species assessed by each agency.

Table 5.

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Summary of native species present in San Luis Potosí by family, order or suborder, and class. Status summary indicates the number of species found in each IUCN conservation status in the order DD, LC, V, NT, E, CE (see Table 1 for abbreviations; in some cases species have not been assigned a status by the IUCN and therefore these may not add up to the total number of species in a taxon). Mean EVS is the mean Environmental Vulnerability Score, scores > 14 are considered high vulnerability (Wilson et al., 2013a,b) and conservation status in Mexico according to SEMARNAT (2010) in the order NL, Pr, A, P (see Table 1 for abbreviations).

Class	Order/ Suborder	Family	Status Summary	Mean EVS	SEMARNAT
Amphibia	Caudata		0,1,1,1,1,1	13.33	0,4,1,1
		Ambystomatidae	0,1,0,0,0,0	10	0,1,0,0
		Plethodontidae	0,0,1,1,1,1	14.5	0,3,1,0
		Salamandridae	0,0,0,0,1,0	12	0,0,0,1
	Anura		0,24,4,3,2,0	8.93	22,9,2,1
		Bufonidae	0,5,0,0,0,0	7.6	5,1,0,0
		Craugastoridae	0,1,1,1,0,0	12.33	1,2,0,0
		Eleutherodactylidae	0,2,3,0,0,0	13.2	4,1,0,0
		Hylidae	0,7,0,1,1,0	7.44	7,1,1,0
		Leptodactylidae	0,2,0,0,0,0	5.5	2,0,0,0
		Microhylidae	0,2,0,0,0,0	6.5	1,1,0,0
		Ranidae	0,2,0,1,1,0	11.75	0,2,1,1
		Rhynophrynidae	0,1,0,0,0,0	8	0,1,0,0
		Scaphiopodidae	0,2,0,0,0,0	4.5	2,0,0,0
	Subtotal		0,25,5,4,3,1	9.59	22,13,3,2
Reptilia	Crocodylia		0,1,0,0,0,0	13	0,1,0,0
		Crocodylidae	0,1,0,0,0,0	13	0,1,0,0
	Testudines		0,3,0,1,0,0	13.14	2,5,0,0
		Emydidae	0,0,0,0,0,0	16	2,0,0,0
		Kinosternidae	0,2,0,1,0,0	11.25	0,4,0,0
		Trionychidae	0,1,0,0,0,0	15	0,1,0,0
	Squamata		4,84,5,0,7,0	10.82	80,26,22,1
	Lacertilia		1,29,3,0,2,0	11.84	28,10,6,1
		Anguidae	1,2,1,0,0,0	14	3,1,0,1
		Corytophanidae	0,1,0,0,0,0	10.5	0,2,0,0
		Crotaphytidae	0,1,0,0,0,0	13	0,0,1,0
		Dactyloidae	0,0,0,0,0,0	8.5	2,0,0,0
		Dibamidae	0,1,0,0,0,0	10	0,0,1,0
		Eublepharidae	0,1,0,0,0,0	9	0,0,1,0
		Iguanidae	0,0,0,0,0,0	12	0,1,0,0
		Phrynosomatidae	0,16,0,0,1,0	12	16,1,2,0
		Scincidae	0,3,0,0,0,0	10.0	4,1,0,0
		Teiidae	0,2,0,0,0,0	11.7	3,0,0,0
		Xantusiidae	0,2,2,0,0,0	13.25	0,3,1,0
		Xenosauridae	0,0,0,0,1,0	15	0,1,0,0
	Serpentes		3,55,2,0,5,0	10.29	52,16,16,0
		Boidae	0,0,0,0,0,0		1,0,0,0
		Colubridae	0,25,0,0,2,0	9.42	27,2,6,0
		Dipsadidae	3,10,0,0,2,0	9.73	14,8,0,0
		Elapidae	0,1,0,0,0,0	11	1,0,0,0
		Leptotyphlopidae	0,2,0,0,0,0	13	3,0,0,0
		Natricidae	0,9,2,0,1,0	11	4,0,8,0
		Viperidae	0,8,0,0,0,0	12,9	2,6,2,0
	Subtotal		4,88,5,1,7,0	10.95	82,32,22,1
TOTAL			4,109,10,4,10,1	10.64	105,45,25,3

The conservation status of the reptiles and amphibians in each habitat type was examined. For amphibians, the percentage of species in protected IUCN categories (VU, NT, EN, CE) varied among the habitat types. Twenty-nine percent of amphibians in the Chihuahuan Desert were listed in IUCN categories, 72% in the Sierra Madre Oriental, 46% in the subtropics of the Sierra Madre Oriental, and 8% of the generalists. For SEMARNAT categories, 57% of amphibians in the Chihuahuan Desert, 72% in the Sierra Madre Oriental, 46% of the subtropics of the Sierra Madre Oriental, and 23% of the generalists were listed. Thus, for amphibians, species found in the Sierra Madre Oriental are the most threatened whereas the generalists were least threatened. Reptiles showed a slightly different pattern. For the IUCN listings, all habitat types had relatively few species in the protected categories (Chihuahuan Desert, 8%; Sierra Madre Oriental, 18%; subtropics of the Sierra Madre Oriental, 10%; and generalists, 9%). However, for SEMARNAT, 28% of reptiles in the Chihuahuan Desert, 50% from the Sierra Madre Oriental, 50% from the subtropics of the Sierra Madre Oriental, and 42% of the generalist species were in the protected categories. For reptiles, the conservation status of the species in each habitat type is more evenly distributed across the habitat types than in amphibians.

Hopefully, by establishing this list of herpetofaunal species with their global and country-level conservation statuses will prompt further investigations into the amphibians and reptiles of this state, which could provide the needed information to allow for state specific, or even habitat type, conservation measures to be undertaken. Specific threats known to be present in San Luis Potosí are deforestation and habitat loss (Miranda-Aragón et al. 2012, Reyes Hernández et al. 2013, Ramos-Lara and Koprowski 2014), industrial pollutants and heavy metals (Alcalá-Jáuregui et al. 2014, Pérez-Vázquez et al. 2016a, b), mining (Razo et al. 2004, Chapa-Vargas et al. 2010, Espinosa-Reyes et al. 2014), and overexploitation of water resources (Esteller et al. 2012).

We thank J. Sigala and J. Penner for very helpful comments that greatly improved the manuscript. Support for this study was provided by Dirección General de Asuntos del Personal Académico, Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (DGAPA-PAPIIT) through the Project IN215418. We are grateful to Alejandra Núñez Merchand from the National Commission for the Understanding and Use of Biodiversity (CONABIO) for kindly creating and providing the topographic, climate, and vegetation maps used in this publication.