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ZooKeys 1166: 287-306 (2023)
DOI: 10.3897/zookeys.1166.86986

Research Article

An updated list of the Mexican herpetofauna: with a summary of
historical and contemporary studies

Aurelio Ramirez-Bautista’™®, Lizzeth A. Torres-Hernandez', Raciel Cruz-Elizalde2®,
Christian Berriozabal-Islas*®, Uriel Hernandez-Salinas™®, Larry David Wilson>®,
Jerry D. Johnson”, Louis W. Porras’, Carlos Jesus Balderas-Valdivia™®,

Adriana J. X. Gonzalez-Hernandez’™®, Vicente Mata-Silva”®

1 Laboratorio de Ecologia de Poblaciones, Centro de Investigaciones Bioldgicas, Instituto de Ciencias Basicas e Ingenieria, Universidad Autonoma del Estado
de Hidalgo, Km 4.5 Carretera Pachuca-Tulancingo, 42184 Mineral de La Reforma, Hidalgo, Mexico

2 Laboratorio de Ecologia y Diversidad Faunistica, Facultad de Ciencias Naturales, Universidad Autonoma de Querétaro, Avenida de las Ciencias S/N, Santa Fe
Juriquilla, C. P 76230, Querétaro, Querétaro, Mexico

3 Programa Educativo de Ingenieria en Biotecnologia, Universidad Politécnica de Quintana Roo, Av. Arco Bicentenario, M 11, Lote 1119-33, Sm 255, 77500 Cancun,

Quintana Roo, Mexico

eo AN DTD oO

Instituto Politécnico Nacional, CIIDIR Unidad Durango, Sigma 119, Fraccionamiento 20 de Noviembre II, Durango 34220, Mexico

Centro Zamorano de Biodiversidad, Escuela Agricola Panamericana Zamorano, Departamento de Francisco Morazan, Tegucigalpa, Honduras

1350 Pelican Court, Homestead, Florida 33035-1031, USA

Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968-0500, USA

7705 Wyatt Earp Avenue, Eagle Mountain, Utah, 84005, USA

Direccion General de Divulgacion de la Ciencia, Zona Cultural de Ciudad Universitaria, Universidad Nacional Auténoma de México, Mexico City, Mexico

10 Coleccidn Nacional de Anfibios y Reptiles, Instituto de Biologia, Universidad Nacional Autonoma de México, Ciudad Universitaria, Coyoacadn, CP04510, Mexico
Corresponding author: Raciel Cruz-Elizalde (cruzelizalde@gmail.com)

OPEN Qrceess

Academic editor: Uri Garcia-Vazquez
Received: 26 May 2022

Accepted: 1 May 2023

Published: 12 June 2023

ZooBank: https://zoobank.
org/903A1DF4-5FED-4825-A03D-
4574CF93698A

Citation: Ramirez-Bautista A, Torres-
Hernandez LA, Cruz-Elizalde R,
Berriozabal-Islas C, Hernandez-Salinas
U, Wilson LD, Johnson JD, Porras
LW, Balderas-Valdivia CU, Gonzdlez-
Hernandez AJX, Mata-Silva V (2023)
An updated list of the Mexican
herpetofauna: with a summary of
historical and contemporary studies.
ZooKeys 1166: 287-306. https://doi.
org/10.3897/zookeys.1166.86986

Copyright: © Aurelio Ramirez-Bautista et al.

This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International - CC BY 4.0).

Abstract

The growth in our knowledge of the diversity of the herpetofauna of Mexico has oc-
curred over the period of approximately 445 years from the work of Francisco Hernan-
dez to that of a broad multinational array of present-day herpetologists. The work of this
huge group of people has established Mexico as one of the most significant centers of
herpetofaunal biodiversity in the world. This status is the result of a complex orography,
in addition to diverse habitats and environments and the biogeographic history of Mexi-
co. The current herpetofauna consists of 1,421 native and introduced species, allocated
to 220 genera, and 61 families. This figure is comprised of 1,405 native species and 16
non-native species (as of April 2023). The non-native species include two anurans, 13
squamates, and one turtle. The level of endemism is very high, presently lying at 63%,
with this level expected to increase with time. Species richness varies among the 32
federal entities in the country, from a low of 50 in Tlaxcala to a high of 492 in Oaxaca.
Amphibian species richness by state-level can be envisioned as comprising three lev-
els of low, medium, and high, with the lowest levels occurring in the Peninsula of Baja
California, a group of seven states in north-central and central Mexico, and a group of
three states in the Yucatan Peninsula, with the highest levels occupying the southern
states of Guerrero, Puebla, Veracruz, Oaxaca, and Chiapas, and the medium level in the
remaining states of the country. Reptile species richness also can be allocated to three
categories, with the lowest level occupying Baja California Sur, a group of central states,
and the states of the Yucatan Peninsula, and the highest level found in a cluster of the
states of Veracruz, Guerrero, Oaxaca, and Chiapas. Knowledge of the Mexican herpeto-
fauna will continue to grow with additional studies on systematics, conservation, and
the construction of checklists at various levels.

287

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

Key words: Amphibians, Mexico, reptiles, species diversity

Introduction

Historically, our knowledge of the diversity of different biological groups
in Mexico has increased through the accumulation of local, regional, state,
and/or country level studies (mammals: Sanchez-Cordero et al. 2014, birds:
Navarro-Siglenza et al. 2014; amphibians: Wilson et al. 2010, 2013a; Par-
ra-Olea et al. 2014; and reptiles Wilson et al. 2010, 2013b; Flores-Villela and
Garcia-Vazquez 2014; Ramirez-Bautista et al. 2014). In studying history of
the Mexican herpetofauna, the amount of species richness in each area has
undergone two major stages: one in which it advanced rapidly, but slowed
after the publication of Smith and Taylor (1966), and the other involving the
integration of local studies that provided information for cataloging species
at the regional level. At the regional or state levels, however, the lists initially
were anecdotal, which resulted in a broad gap between the actual records
and the first official herpetofaunal catalogs (as reprinted in Smith and Tay-
lor 1966).

The contributions of Smith and Smith (1976, 1977, 1993) laid the foundation
for a series of studies aimed at determining the species richness of amphib-
ians and reptiles. Importantly, the growing number of amphibian and reptile
species and subspecies noted by these authors was highly significant. In re-
cent years, most of the species’ richness has been assessed at the species
level, based on the dubious legitimacy of subspecies as a formal taxonomic
hierarchical category (Flores-Villela et al. 2004; Wilson et al. 2010; Johnson et
al. 2015). Nonetheless, the number of species keeps increasing, based on the
rate of taxonomic changes, the description of new species, rediscoveries, and
the resurrection of species (Lara-Tufifo et al. 2016; Nicholson et al. 2018; Nie-
to-Montes de Oca et al. 2018; Mata-Silva et al. 2019). An increasing amount of
species richness has also resulted from the continuing amount of work being
conducted in several parts of the country. In addition to the information provid-
ed by Smith and Taylor (1966), Alvarez del Toro (1982) reported on the conser-
vation of biodiversity at both the regional and state levels (Flores-Villela et al.
2004). Prior to the publication of Smith and Taylor (1966), there were various
studies on the description of species and subspecies, but there were few stud-
ies aimed at listing species by region, and their contribution was the first to pro-
vide information on species richness (a list of species), keys for determining
species, synonymies, and information on geographic distribution. During this
period, an interest in these groups was minimal based on the low number of
herpetologists in Mexico, and because institutions that promoted field research
had limited (or non-existing) funds to print catalogs at regional and state levels
(Pérez-Higareda and Smith 1991).

Finally, between 1990 and 2000, a strong interest in biodiversity led to an
assessment of the herpetofauna of Mexico at both the regional and state lev-
els (Flores-Villela and Canseco-Marquez 2004; Wilson et al. 2013a, b). Sub-
sequently, information from these projects, along with long-term studies and
continued sampling, provided a more accurate number of the species involved.
These studies were followed by larger ones, which included a greater number

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 288

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

of states (Vazquez-Diaz and Quintero Diaz 1997, 2005; Lemos-Espinal and
Smith 2009; Dixon and Lemos-Espinal 2010; Lemos-Espinal and Dixon 2013;
Ramirez-Bautista et al. 2014; among others), as well as those from the entire
country (Flores-Villela and Canseco-Marquez 2004; Wilson and Johnson 2010;
Wilson et al. 2013a, b; Johnson et al. 2017). Presently, these studies form the
basis of our knowledge of the biodiversity of amphibians and reptiles in Mex-
ico (Wilson and Johnson 2010; Wilson et al. 2013a, b; Johnson et al. 2017;
Balderas-Valdivia and Gonzalez-Hernandez 2021; Balderas-Valdivia et al. 2022;
Smith and Lemos-Espinal 2022).

Flores-Villela (1993a) reported the number of species in Mexico based pri-
marily on the published literature and provided an analysis of the Mexican
herpetofauna based on Smith and Taylor (1966) and Smith and Smith (1976,
1977, 1993). Additionally, Flores-Villela (1993b) noted the number of amphib-
ian and reptile species as 291 and 706, respectively; these numbers eventual-
ly changed to 361 and 804 species, respectively (1,165 species; Flores-Villela
and Canseco-Marquez 2004). In time, Liner (2007) provided an updated list
that included both species and subspecies and indicated 393 amphibians
and 1,234 reptiles. The remarkable species richness reported in these stud-
ies was related to several factors, such as the complex physiography of Mex-
ico, the representation of both Nearctic and Neotropical regions in the coun-
try, as well as its diverse biogeographic regions and vegetational formations
(Morrone 2005). Subsequent knowledge of the herpetofauna was analyzed
in the context of its diversity, in which its components are the richness and
abundance of species (Ochoa-Ochoa et al. 2014), and how this might vary
according to environmental parameters (Cruz-Elizalde et al. 2016). Similar
numbers of amphibians (376) and reptiles (864) were reported by Parra-Olea
et al. (2014) and Flores-Villela and Garcia-Vazquez (2014), respectively; how-
ever, more recent studies showed an increase to 1,395 native species (394
amphibians, and 898 reptiles; Johnson et al. 2017; Balderas-Valdivia et al.
2022). These studies did not consider any introduced species and should
be updated with recent taxonomic changes and new species and records
(i.e., Hidalgo-Garcia et al. 2023). Additionally, increases in the richness and
diversity of the herpetofauna is constantly monitored (i. e. Herpetologia Mex-
icana 2023; Mesoamerican Herpetology Taxonomic List 2023) but they do
not provide a historical analysis of the richness or the proportion of state
herpetofaunal biodiversity.

The increase of species richness reflects the systematic studies conducted
by both Mexican and foreign herpetologists during the last three decades, in
addition to the impact of the previously mentioned historical studies. There-
fore, with the descriptions of new species, the knowledge of the biodiversity of
Mexico is enhanced, as well as attention to its conservation needs (Campbell
et al. 2018; Mata-Silva et al. 2019). This statement is supported by recent stud-
ies in which several species were described, e.g., the salamanders Chiroptero-
triton aureus or C. nubilus (Garcia-Castillo et al. 2018), the lizards Gerrhonotus
mcecoyi (Garcia-Vazquez et al. 2018a), and Xenosaurus fractus (Nieto-Montes
de Oca et al. 2018), snakes such as Rhadinaea nuchalis (Garcia-Vazquez et al.
2018b) and Geophis cansecoi (Griinwald et al. 2021), among others, and even
the descriptions of new snake genera such as Cenaspis (Campbell et al. 2018),
Metlapilcoatlus (Campbell et al. 2019) and Desertum (Blair et al. 2022).

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 289

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

Because of the efforts invested to study the Mexican herpetofauna by nu-
merous scientists with varying levels of experience, Mexico is known as one
of the richest countries with respect to this group of vertebrates (Wilson et al.
2010, 2013a, b; Johnson et al. 2017). Considering the above information, the
main goal of this study is to provide an updated list of the species that comprise
the Mexican herpetofauna (as of April 2023). The update of amphibian and rep-
tile species for Mexico (and by states) is necessary due to the dynamics that
have been taking place in the descriptions of new species for science, as well
as recent taxonomic changes and new reports of exotic species, which collec-
tively increase the number of species in these groups. The latest studies where
the lists of species presented by Balderas-Valdivia and Gonzalez-Hernandez
(2021) and Balderas-Valdivia et al. (2022) with only native species, however,
in the present study the most updated list is shown both in number of species
(native and introduced) and in taxonomic update.

Materials and methods

The updated species lists are based on the following:

1. Publications such as Herpetology of Mexico (Smith and Taylor 1966), and
studies at the regional (Pérez-Higareda and Smith 1991; Ramirez-Bautista
1994; Uribe-Pefia et al. 1999; among others), state (Alvarez del Toro 1982;
Vazquez-Diaz and Quintero-Diaz 1997, 2005; Lemos-Espinal and Smith
2009, 2016; Ramirez-Bautista et al. 2009, 2010, 2014; Dixon and Lemos-Es-
pinal 2010; Alvarado-Diaz et al. 2013; Lemos-Espinal and Dixon 2013; Car-
bajal-Marquez and Quintero-Diaz 2016; Lemos-Espinal et al. 2016, 2017,
2018a, b, 2019; Nevarez-de los Reyes et al. 2016; Teran-Juarez et al. 2016;
Woolrich-Pina et al. 2016, 2017; Cruz-Saenz et al. 2017; Gonzalez-Sanchez
et al. 2017; Palacios-Aguilar and Flores-Villela 2018; Ramirez-Bautista et
al. 2020; Rhodin et al. 2021; Barragan-Vazquez et al. 2022; Cruz-Elizalde
et al. 2022; Leyte-Manrique et al. 2022; among others), and country levels
(Flores-Villela 1993a; Flores-Villela and Canseco-Marquez 2004; Liner 2007;
Wilson and Johnson 2010; Wilson et al. 2013a, b; Flores-Villela and Garcia-
Vazquez 2014; Parra-Olea et al. 2014; Mata-Silva et al. 2015; Johnson et al.
2015, 2017; Balderas-Valdivia and Gonzalez-Hernandez 2021; Mata-Silva
et al. 2021; Balderas-Valdivia et al. 2022; Smith and Lemos-Espinal 2022);

2. historical records (databases) compiled by the authors, which were
searched from 1940 to date;

3. records obtained in specialized journals, such as Mesoamerican Herpetol-
ogy, ZooKeys, Zootaxa, Revista Mexicana de Biodiversidad, Herpetologi-
cal Review, and Amphibian & Reptile Conservation, as well as publications
of newly described species for the country (until 30 April 2023);

4. databases generated from projects undertaken by the Comisi6n Nacional
para el Conocimiento y Uso de la Biodiversidad (CONABIO 2019; https://
www.gob.mx/conabio); and

5. records of amphibians and reptiles from Mexico obtained from the Global
Biodiversity Information Facility (GBIF 2019; https://www.gbif.org/), and
authenticated by geographic distribution records from previously pub-
lished literature.

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 290

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

All taxonomic changes and the identification of the species were based on
the most recent taxonomic revisions (Martinez-Méndez and Méndez-De la Cruz
2007; Poe 2013; Wilson et al. 2013a, b; Ruane et al. 2014; Meza-Lazaro and
Nieto-Montes de Oca 2015; Montanucci 2015; Rovito et al. 2015; Duellman
et al. 2016; Leavitt et al. 2017; Canseco-Marquez et al. 2018; Campbell et al.
2019; Campillo-Garcia et al. 2021; Gutiérrez-Rodriguez et al. 2021; Everson et
al. 2021; Hernandez-Jiménez et al. 2021; Kohler 2021; Blair et al. 2022; Hernan-
dez-Austria et al. 2022; Pérez-Ramos and Luja-Molina 2022; Reyes-Velasco et
al. 2022; Devitt et al. 2023; Pérez-Ramos 2023). The scientific names of am-
phibians were updated according to Amphibian Species of the World (Frost
2023, https://www.amnh.org), and for reptiles, based on the Reptile Database
website (Uetz et al. 2023; http://www.reptile-database.org/).

Results
Composition of the current Mexican herpetofauna

Our results show that the current native herpetofauna of Mexico is comprised
as follows: amphibians = 16 families, 58 genera, and 430 species (Suppl. mate-
rial 1); and reptiles = 44 families, 155 genera, and 975 species (Suppl. material
2). For both groups the totals = 60 families, 213 genera, and 1,405 species (see
Suppl. materials 1, 2; Table 1).

Table 1. Composition of the native herpetofauna of Mexico, including number of endemic species and percentage of

endemism.

Orders

Anura
Caudata
Gymnophiona
Subtotals
Crocodylia
Squamata
Testudines
Subtotals

Totals

Families

11
4

Percentage of

Genera Species Endemic Species Endemism (%)

37 270 168 62.2
19 157 134 83.4
5 3 1 33:3
58 | 430 300 69.8
2 | 3 = :

135 921 566 61.3
18 51 20 39:2

155 975 586 60.1

213 1405 886 63.1

Increase in the total size of the native Mexican herpetofauna from 1577
to the present

The size of the Mexican herpetofauna has increased ca. 20.1-fold in the period
from 1577 to the present (from 71 species to 1,421; see Fig. 1). During the en-
suing 319 years, the number of species has increased ca. 3.1-fold, from 71 to
219 species or ca. 310%. The increase over this three-century period was 148
species. The largest increase during the history of the study of the Mexican
herpetofauna transpired from 1896 to 1993 (97 years), when the number of
species rose from 219 to 995 (or 776 species), a 3.5-fold increase or ca. 350%.

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 291

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

Number of species

—_> _
oe N
o o&
o o&

Since 1993, however, the rate of increase has slowed somewhat. From 1993
(995 species) to 2004 (1,165 species), the increase was 170 species; from
2004 (1,165 species) to 2013 (1,227 species) 62 species were added; and from
2013 (1,227 species) to 2017 (1,292 species) there were 65 additional species.
Finally, from 2017 to the present (2023), the numbers increased from 1,292
species to 1,421, in this study 129 species are added (Fig. 1).

1421

ee Ne

1165

1577. 1896 1993 2004 2013 2017 #42023
Year

Figure 1. Cumulative curve of the herpetofauna recorded in Mexico. The years correspond to studies carried out by
Hernandez (1577, published in 1959), Dugés (1896), Flores-Villela (1993a), Flores-Villela and Canseco-Marquez (2004),
Wilson et al. (2013a, b), Johnson et al. (2017), and this study.

Increase in the size of the non-native herpetofauna in Mexico

Currently, 16 species of non-native or exotic amphibians and reptiles have been
reported from Mexico (Gonzalez-Sanchez et al. 2021; Tepos-Ramirez et al.
2022), including two amphibians (Eleutherodactylus planirostris and Xenopus
laevis) and 14 reptiles (Anolis allisoni, A. carolinensis, A. cristatellus, A. sagrei,
Gehyra mutilata, Hemidactylus frenatus, H. garnotii, H. mabouia, H. turcicus,
Lepidodactylus lugubris, Tarentola mauritanica, Sphaerodactylus argus, Virgoty-
phlops braminus, and Graptemys pseudogeographica).

Endemism of the current Mexican herpetofauna

The percentage of species endemism of the Mexican herpetofauna is very high
at 63.1% (Table 1). By groups, amphibians represent the highest percentage of
endemism, with 69.8% (with 300 species), and a somewhat lower number in
reptiles at 60.1% (586 species).

Species richness among states

Currently, of the 32 federal entities, three states in southeastern Mexico (Oax-
aca, Veracruz, and Chiapas) maintain the highest richness levels (Fig. 2), i.e.,
492 (34.6% of Mexican herpetofauna), 367 (25.8%), and 364 (25.6%), respec-
tively (Figs 3, 4, Suppl. material 3). These states are followed by Guerrero (291,
20.5%), Puebla (278, 19.6%), Jalisco (259, 18.2%), Michoacan (231, 16.3%),

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 292

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

600

500

300

Number of species

200

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12.00
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os A & Cee £y RS FX oy oe o ee if ‘s OS * ree G ss oe
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States

Figure 2. Number of herpetofaunal species in Mexico by state (bars), and proportion of territorial surface (line).

Sonora (208, 14.6%), Tamaulipas (204, 14.4%), San Luis Potosi (202, 14.2%)
and Hidalgo (200, 14.1%), whereas the states with the lowest number of spe-
cies are Zacatecas (118, 8.3%), Baja California Sur (108, 7.6%), Guanajuato (98,
6.9%), Aguascalientes (98, 6.9%), Ciudad de México (82, 5.8%), and Tlaxcala
(50, 3.5%; Figs 2-4, Suppl. material 3).

Areal extension by state and species richness

The size of the 32 federal entities in Mexico varies considerably from that of
Ciudad de México (1,494.6 km?) to that of Chihuahua (247,412.5 km?), of which
the latter is 165.5 times larger than the former (Suppl. material 3). Obviously,
the percentage of the territorial area occupied by these 32 entities also varies
considerably (from 0.08 to 12.62%). These percentage values fall into a number
of categories, including 0.08-1.00% (Aguascalientes, Colima, Ciudad de Méxi-
co, Morelos, Querétaro, Tlaxcala), 1.01-3.00 (Campeche, Guanajuato, Hidalgo,
Estado de México, Michoacan, Nayarit, Puebla, Quintana Roo, Sinaloa, Tabasco,
and Yucatan), 3.01-6.00 (Baja California, Baja California Sur, Chiapas, Guerre-
ro, Jalisco, Nuevo Le6n, Oaxaca, San Luis Potosi, Tamaulipas, Veracruz, and
Zacatecas), and 6.01 and above (Chihuahua, Coahuila, Durango, and Sonora).
By territorial area, the largest states are Chihuahua, Sonora, Coahuila, and Du-
rango, all located in northern and north-central Mexico (Fig. 2). These states,
however, exhibit relatively low species richness (197, 208, 150, and 179, respec-
tively) compared to several smaller states like Oaxaca, Chiapas, Guerrero, Vera-
cruz, Puebla, and Jalisco (see values in section above; Fig. 2; Suppl. material 3).

The lowest values for amphibian species (7-29) are found in the two states
of the Peninsula of Baja California, a north-central swath of states that include
Coahuila, Nuevo Leon, Zacatecas, Aguascalientes, and Guanajuato, two small
states in central Mexico, Ciudad de México, and Tlaxcala, and the Yucatan Pen-
insula states, Campeche, Yucatan, and Quintana Roo (Fig. 3). The next high-
est category (30-65) includes the northwestern and western states of Sonora,

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 293

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

110°0'O"W 100°0'0"W 90°0'O"W

Amphibians of Mexico

Gulf of Mexico

Pacific Ocean

Number of species

We 7-29
30-65

290) 145 O 290 580 870 1,160
WO cc-164 |
a ee Ki) ometers
Figure 3. Number of species of amphibians in Mexico by state. The color scale represents the species numbers from
low (green) to high (red).

Chihuahua, Durango, Sinaloa, and Nayarit, Jalisco, Colima, and Michoacan, and
the northeastern states, of Tamaulipas and San Luis Potosi, the east-central
states of Hidalgo and Querétaro, the central Mexican states including state of
México and Morelos, and southeast state of Tabasco. The highest category
(66-164) is comprised of a group of states in southern Mexico, including Guer-
rero, Puebla, Veracruz, Oaxaca, and Chiapas.

According to the number of species of reptiles by state the resulting cate-
gories are somewhat different than those for the amphibians, since the lowest
rank (32-120) is occupied by the states of Baja California Sur, a smaller central
Mexican swath that includes Zacatecas, Aguascalientes, Guanajuato, Queréta-
ro, Estado de México, Ciudad de México, Tlaxcala, and Morelos, the state of Coli-
ma, and the states of the Yucatan Peninsula (Campeche, Yucatan, and Quintana
Roo) (Fig. 4). The medium category (121-199) comprises the remaining states
in northern and central Mexico, and southeastern state of Tabasco. The high
category (200-328) consists of a single cluster of states that includes Guerrero,
Veracruz, Oaxaca, and Chiapas. The general pattern for the distribution of reptile
species in Mexico is a low level of species richness in Baja California Sur, the
central Mexican swath of states and Colima, and the states of Campeche, Yu-
catan, and Quintana Roo, a medium level in the remaining states in northern and
central Mexico, and the southeastern state of Tabasco, and lastly, a high level
in the southern and eastern states of Guerrero, Veracruz, Oaxaca, and Chiapas.

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 294

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

110°0'O"W 100°0'O"W 90°0'O"W

Reptiles of Mexico

Gulf of Mexico

Pacific Ocean

Number of species

ME 32-120
| 121-199
WO 200-328

Figure 4. Number of species of reptiles in Mexico by state. The color scale represents the species numbers from low
(green) to high (red).

Discussion

The richness of amphibian and reptile species in Mexico has been the subject
of study for several decades, generating various works listing the herpetofauna
at the country level. In this study, we update the list of species to 1,421, of which
1,405 are native species, and 16 are non-native. The number of native species we
record in this study is higher compared to that of recent works by Balderas-Valdiv-
ia and Gonzdalez-Hernandez (2021; 1,389 species), and Balderas-Valdivia et al.
(2022; 1,393 species) for the herpetofauna of Mexico, based on new discoveries.

The high species richness and endemism of both groups stands out com-
pared to that of other groups of vertebrates in Mexico (Smith and Lemos-Espinal
2022), since Johnson et al. (2017) noted that the country of Mexico contains
the fifth largest amphibian fauna and the second largest reptile fauna in the
world. At that time, the size of the Mexican amphibian fauna was smaller than
only those of Brazil, Colombia, Peru, and Ecuador, and the Mexican reptile fauna
was smaller than only that of Australia.

The brief historical review shown in the introduction of this work highlights
the continuous study of the herpetofauna to date, since it began with the ac-
knowledgement of the amphibian and reptile fauna by the Aztecs, which lat-
er increased with the Spanish conquest of the Americas (Martin del Campo
1979). Although Europeans made this conquest in the 17" century, the principal

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 295

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

impulse that led to our current knowledge of the Mexican herpetofauna did not
occur until the second half of the 19" century and was made possible through
the contributions of Smith and Smith (1976) and Flores-Villela (1993b). By late
19" century the number of species in the country was ca. 219, but by the end
of 20" century it had increased to 995 species (Fig. 1). This dynamic increase
was driven primarily by the continuous work of Mexican (Alvarez-del Toro 1982;
Flores-Villela and Canseco-Marquez 2004; Lemos-Espinal and Smith 2009,
2016; Lemos-Espinal and Dixon 2013; Ramirez-Bautista et al. 2014; Mata-Silva
et al. 2015; Lemos-Espinal et al. 2016) and American herpetologists (Wilson et
al. 2010; 2013a, b; Johnson et al. 2017). At the beginning of the 21% century,
the increased herpetofaunal species richness resulted from new state records
(Badillo-Saldafia et al. 2019; Tepos-Ramirez et al. 2022) and the descriptions of
new species (Flores-Villela and Canseco-Marquez 2004; Lara-Tufifio et al. 2016;
Campbell et al. 2018; Garcia-Castillo et al. 2018; Garcia-Vazquez et al. 2018a,
b; Barragan-Reséndiz et al. 2022; Flores-Villela et al. 2022a, b; Nieto-Montes de
Oca et al. 2022; Griinwald et al. 2023). All these studies increased the herpeto-
fauna to a current total of 1,421 species.

Based on this number, the herpetofauna of Mexico has been analyzed by au-
thors in other related fields, including conservation (Wilson et al. 2010) and bio-
geography (Wilson et al. 2013a, b; Johnson et al. 2017). The species endemism
value of 63.1% is similar to that reported by Johnson et al. (2017; 61.2%) but is
higher than that reported for Central America by Mata-Silva et al. (2019; 56.9%).
By comparing the 2017 value and the present values for Mexican endemism,
evidently the species added to the herpetofauna for ca. the last five years have
been reported primarily as endemic to the country, as opposed to reported for
Mexico from outside the country.

The states in southeastern Mexico, including Oaxaca, Chiapas, and Veracruz,
have shown the highest amount of species richness (Figs 2—4; Suppl. material 3;
Balderas-Valdivia and Gonzdlez-Hernandez 2021; Balderas-Valdivia et al. 2022),
and these values are partly supported by such factors as environmental hetero-
geneity, climate, vegetation types, and their influence on biogeographic regions,
as documented by other studies (Ochoa-Ochoa et al. 2014; Ramirez-Bautista et
al. 2014; Cruz-Elizalde et al. 2016, among others). Note that states in the north-
ern part of the country, which are located within a longer territorial extension,
have a low number of species (Fig. 2; Suppl. material 3). These states are also
characterized by an environmental homogeneity in vegetation types and other
physiographic characteristics (Lemos-Espinal and Dixon 2013; Lemos-Espinal
et al. 2016). For states with a small territorial extension, such as Guerrero, Pueb-
la, Jalisco, Tamaulipas, San Luis Potosi, Michoacan, Nayarit, and Hidalgo, their
high species richness might be explained not only by their environmental hetero-
geneity, but also by the convergence of biogeographic zones (Ochoa-Ochoa et
al. 2014; Ramirez-Bautista et al. 2014; Palacios-Aguilar and Flores-Villela 2018;
Smith and Lemos-Espinal 2022). These results have been explained for other
studies on the Mexican herpetofauna (Smith and Lemos-Espinal 2022), as well
as in other biological groups such as mammals (Ceballos et al. 1998) or birds
(Ramirez-Albores et al. 2020). In these studies, the authors show a trend in which
the tropical zones, and mainly in the Mesoamerican region and low latitudes,
the richness of species increases. This trend is also supported by the complex
orography and diverse environments in these areas (Ochoa-Ochoa et al. 2014).

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 296

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

Future studies on systematics (Campillo-Garcia et al. 2021), regional check-
lists, and isolated geographic records will keep increasing the number of spe-
cies. In this sense, the continuous study of the herpetofauna conjugates on
plans regarding conservation, since a high percentage of herpetofaunal spe-
cies are endemic to the country, and many are regarded as threatened on ac-
count of human activities, e.g., overpopulation, habitat fragmentation, pollution,
agriculture, among other factors (Wilson et al. 2010; Johnson et al. 2017).

Acknowledgments

We thank Itzel Magno Benitez, César Adrian Diaz Marin, Luis Manuel Badillo Sal-
dana, Daniel Lara Tufifo, Ismael Reaho Hernandez, and Raquel Hernandez Austria
for logistical help during this study. We also wish to thank Lydia Allison Fucsko
for her perceptive review of this paper. We thank Julio Lemos Espinal, Luis Can-
seco Marquez, and Uri Garcia Vazquez for helpful comments on the manuscript.

Additional information

Conflict of interest

No conflict of interest was declared.

Ethical statement

No ethical statement was reported.

Funding

Funding from CONABIO JMO001 was provided to ARB, and PAPIME UNAM PE205618
to CUBV.

Author contributions

Conceptualization: LWP RCE, CUBV, LDW, JDJ, UHS, AJXGH, ARB, CBI, VMS. Data cu-
ration: CUBV, LWP JDJ, LATH, RCE, AJXGH, ARB, VMS, UHS, LDW, CBI. Formal analy-
sis: LATH, AJXGH, LDW, RCE, VMS, CJUBV, ARB. Funding acquisition: RCE, AJXGH, CJBV,
ARB. Investigation: LATH, LDW, UHS, ARB, RCE, AJXGH, CUBV. Methodology: JDJ, CBI,
UHS, LDW, CJBV, LWP, LATH, ARB, VMS, RCE. Project administration: RCE, LDW, ARB. Re-
sources: CUBV, RCE, AJXGH, ARB, VMS, LATH, LDDW. Software: RCE. Supervision: RCE,
ARB, VMS, LDW. Validation: LDW, RCE, AJXGH, LWP, CUBV, JDJ, VMS, ARB. Visualization:
ARB, LDW, RCE. Writing - original draft: ARB, UHS, RCE. Writing - review and editing: LWP
VMS, CJBV, ARB, JDJ, LDW, AUXGH, RCE, CBI, LATH.

Author ORCIDs

Aurelio Ramirez-Bautista © https://orcid.org/0000-0003-2948-0178

Raciel Cruz-Elizalde © https://orcid.org/0000-0001-9630-7063

Christian Berriozabal-Islas © https://orcid.org/0000-0002-3739-3990

Uriel Hernandez-Salinas © https://orcid.org/0000-0001-9630-7063

Larry David Wilson © https://orcid.org/0000-0003-4969-0789

Jerry D. Johnson © https://orcid.org/0000-0002-2135-4866

Carlos Jesus Balderas-Valdivia © https://orcid.org/0000-0002-1378-7610
Adriana J. X. Gonzalez-Hernandez © https://orcid.org/0000-0002-6445-3338
Vicente Mata-Silva © https://orcid.org/0000-0001-8123-1844

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 297

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

Data availability

All of the data that support the findings of this study are available in the main text or
Supplementary Information.

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Supplementary material 1

List of 430 taxa of native (and 2 non-native) amphibians of Mexico

Authors: Aurelio Ramirez-Bautista, Lizzeth A. Torres-Hernandez, Raciel Cruz-Elizalde
Christian Berriozabal-Islas, Uriel Hernandez-Salinas, Larry David Wilson, Jerry D. John-
son, Louis W. Porras, Carlos Jesus Balderas-Valdivia, Adriana J. X. Gonzalez-Hernan-
dez, Vicente Mata-Silva

Data type: Species list (xlsx. file)

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.1166.86986.suppl1

Supplementary material 2

List of 975 taxa of native (and 14 non-native) reptiles of Mexico

Authors: Aurelio Ramirez-Bautista, Lizzeth A. Torres-Hernandez, Raciel Cruz-Elizalde
Christian Berriozabal-Islas, Uriel Hernandez-Salinas, Larry David Wilson, Jerry D. John-
son, Louis W. Porras, Carlos Jesus Balderas-Valdivia, Adriana J. X. Gonzalez-Hernan-
dez, Vicente Mata-Silva

Data type: Species list (xlsx. file)

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.1166.86986.suppl2

ZooKeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 305

Aurelio Ramirez-Bautista et al.: Herpetofauna of Mexico

Supplementary material 3

Territorial area, percentage territorial of county, and total number of species
of amphibians, reptiles, and herpetofauna of each state in Mexico

Authors: Aurelio Ramirez-Bautista, Lizzeth A. Torres-Hernandez, Raciel Cruz-Elizalde
Christian Berriozabal-Islas, Uriel Hernandez-Salinas, Larry David Wilson, Jerry D. John-
son, Louis W. Porras, Carlos Jesus Balderas-Valdivia, Adriana J. X. Gonzalez-Hernan-
dez, Vicente Mata-Silva

Data type: table (docx. file)

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.1166.86986.suppl3

Zookeys 1166: 287-306 (2023), DOI: 10.3897/zookeys.1166.86986 306