Print
A review of the introduced herpetofauna of Mexico and Central America, with comments on the effects of invasive species and biosecurity methodology
expand article infoVíctor Hugo González-Sánchez, Jerry D. Johnson§, David González-Solís, Lydia Allison Fucsko|, Larry David Wilson
‡ El Colegio de la Frontera Sur, Chetumal, Mexico
§ The University of Texas at El Paso, El Paso, United States of America
| Swinburne University of Technology, Melbourne, Australia
¶ Centro Zamorano de Biodiversidad, Escuela Agrícola Panamericana Zamorano, Homestead, United States of America
Open Access

Abstract

Among the principal causes producing detrimental effects on global biodiversity are introductions of alien species. Very few attempts to control introduced amphibians and reptiles in Middle America (Mexico and Central America) can be identified, so listings are provided for 24 exotic species, 16 translocated species, and 11 species that were removed from the introduced species listing because of lack of substantiating evidence that they are from established populations. Biosecurity methods are also identified that can be applied for preventing, controlling, and managing introduced and especially invasive species.

Resumen

Entre las principales causas que producen efectos perjudiciales sobre la biodiversidad mundial se encuentran la introducción de especies exóticas. Se pueden identificar muy pocos intentos de controlar anfibios y reptiles introducidos en América Central (México y América Central), por lo que proporcionamos listas de 24 especies exóticas, 16 especies translocadas y 11 especies que eliminamos de la lista de especies introducidas debido a la falta de evidencia que corrobore que provienen de poblaciones establecidas. También identificamos métodos de bioseguridad que se pueden aplicar para prevenir, controlar y gestionar especies introducidas y especialmente invasoras.

Keywords

Biological invasion, exotic species, herpetofauna, introduced species, Middle America, translocated species

Palabras claves

Especies exóticas, especies introducidas, especies translocadas, herpetofauna, invasion biológica, mesoamérica

Introduction

Among the most important drivers for biotic extinctions are introduced invasive species (Mack et al. 2000; Simberloff 2005). This phenomenon is not new, since human-mediated biological invasions or translocations of non-native species (e.g., goats, pigs, cats, dogs, and rats, among various mammals) by humans have been occurring for at least 20,000 years (Hofman and Rick 2018). Geographic scope, frequency, and number of species involved, however, have increased as a direct consequence of growths in transportation and commercial activities, so few habitats on Earth remain devoid of species introduced by humans (Mack et al. 2000). Among others, the main pathways for introducing amphibians and reptiles are accidental transport in cargo shipments on land and over water; intentional and accidental release via pet trade; as biocontrol agents; those associated with human food consumption, as well as for aesthetic purposes (Kraus 2003). Among many other effects, invasive non-native species can have negative impacts at all biological levels, including genetic pollution and hybridization (Cedeño-Vázquez et al. 2008), competition and depredation (Sakai et al. 2001), introducing parasites (Williams et al. 2013), epizootics (Garner et al. 2006), zoonoses (Hulme 2014), habitat modification, by altering nutrient and energy cycles and biomass structure (Beard et al. 2002; Crooks 2002), shifting water cycles (Gallardo et al. 2015), and triggering trophic cascades (Zavaleta et al. 2001). The impact of invasive species, such as the anurans Lithobates catesbeianus and Rhinella horribilis, as two well-known examples, on human economics and human well-being is considered collateral damage, because of the loss and alterations to goods (agricultural crops, animal husbandry, forest products, fisheries) and services (clean water, climate stabilization, pollination, human culture, recreation), as identified by Pejchar and Mooney (2009) and Walsh et al. (2016).

A common misbelief is that the ecology of invasive species and the ecosystem alterations they produce are extensively documented (Cadotte 2006). This is only partially true, since most research on ecology of alien species involves significant taxonomic and geographical biases (Pyšek et al. 2008; MacIsaac et al. 2010). This situation is relevant since historically introduced amphibians and reptiles have received less attention in Mesoamerica than have other groups, such as mammals, vascular plants, and insects (Reed and Kraus 2010). This deficiency has led to an omission of the status of introduced and invasive species in several herpetofaunal inventories. Therefore, listings of exotic and invasive herpetofaunal species in Middle America (Mexico and Central America) have been comprised of only a few species, including members of the genus Boa, Morelet’s Crocodile (Crocodylus moreletii), the Pond Slider (Trachemys scripta), and the American Bullfrog (Lithobates catesbeianus). Gekkonid lizards are found frequently in listings of introduced species, but their ecology and potential impacts have received little attention. Presently, 78 amphibian and 198 recognized reptile species have become established outside their native ranges around the world (Capinha et al. 2017). The goal of this paper is to review and assess the current knowledge and status of members of the introduced herpetofauna in Middle America, their history of colonization, their impacts on ecosystems, and their current geographic distributions.

Materials and methods

We compiled a list of the introduced reptiles and amphibians in Middle America by examining relevant literature for the region, complemented with records obtained from GBIF (GBIF.org 2018), iNaturalist (inaturalist.org 2018), Amphibian Species of the World (Frost 2020), Reptile Database (Uetz et al. 2020), International Union for Conservation of Nature (2020), and VertNet (vertnet.org, 2018) platforms. In addition to these sources, for Mexico we considered only records with acceptable confirmation, properly georeferenced, and found in the literature and databases of the Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO 2018) and Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT 2017). Scientific names are based on Wilson et al. (2013a, b) and Johnson et al. (2015b), along with the most recent lists at Uetz et al. (2020) and Frost (2020), with full understanding that nomenclatural changes will occur regularly during future taxonomic revisions. Common names, when appropriate, follow Liner and Casas-Andreu (2008) for Mexico and adjacent Central America when species are shared between the two regions. Common names of species occurring only outside of Mexico and adjacent areas are those found in the literature and websites listed above. For convenience, we use the term “reptiles” to name taxonomic groups traditionally considered orders of the class Reptilia as listed by Uetz et al. (2020) (i.e.., Testudines – turtles; Crocodylia – crocodiles; Squamata – snakes and lizards), so we use the term herpetofauna when generally referring to the amphibians, turtles, crocodiles, snakes, and lizards occurring within Middle America. For more on our taxonomic positions, see the section below and especially Johnson et al. (2015b).

Our study area comprises Mexico and all Central American countries (Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, and Panama), ordered by latitude. Our use of the term “Middle America” refers to the Central American countries plus Mexico. We do not use the term “Mesoamerica,” since this label is generally considered more relevant in an anthropological and historical context (Romero-Contreras and Ávila-Ramos 1999), even though it is a commonly used synonym for Middle America.

In this paper, we consider introduced species to be populations whose presence in an area is attributed to human activities that enabled them to overcome biogeographical barriers that they otherwise could not cross and become established. We prefer introduced as the universal term over some others, such as exotic, non-native, alien, or non-indigenous, since the word “introduced” is more easily associated with human intervention (Young and Larson 2011). We will use the terms exotic and translocated most often, however, to describe the two major groupings of introduced species, and use other synonymous terms occasionally to lessen redundancy. This idea is clearly opposite to the concept of “native species,” to designate those that have evolved in a given area without human involvement, or that have arrived there by natural means without intentional or unintentional intervention of humans, from areas in which they are also native (Pyšek and Richardson 2010).

Introduced species are not all-encompassing on temporal and geographical bases, because not all introduced organisms manage to become “established;” i.e., surviving long enough to produce descendant lineages (Davis 2009). We also understand that some species occasionally can be “naturalized,” which are those introduced species sustaining self-replacing populations for several life cycles without, or despite, direct intervention by humans (Richardson et al. 2011). In this sense, “population persistence” is the ongoing accumulation of establishment successes by individuals arising within an area (Davis 2009). Non-native species do not have to be introduced necessarily within the totality of a region, but transported to areas within the same area outside their native range; these will be referred to as “translocated species” (Shine et al. 2000). Countries are good examples of useful geographic units for qualifying which type of introduction a population exemplifies (exotic versus translocated), although sometimes adjustments need to be made to meet criteria, such as between islands and the mainland belonging to the same country, or a slightly disconnected mainland from a large peninsula belonging to the same country (e.g., Baja California peninsula and mainland Mexico). A closely related concept is “invasive species,” here defined as those well-established introduced species having deleterious effects on native ecosystems as the result of increasing their population numbers. In addition, invasive species can have deleterious effects on native populations, even if their populations are not increasing, e.g., through the introduction of new diseases, which can then kill native species.

The “impact” of an introduced invasive species, either exotic or translocated, refers to how an introduced species distresses the physical, chemical, or biological environment, the effect of which might be evident at the genetic, individual, population, community, ecosystem, landscape, regional, or global levels (Parker et al. 1999; Mack et al. 2000; Richardson et al. 2011). Of course, we do not overlook the fact that impacts and the reasons for them might be controversial (Parker et al. 1999; Young and Larson 2011), and that many invasive species can have negative influences on cultural, economic, and social issues relating to alleged human welfare (McNeely 2001; Perrings et al. 2005; Pejchar and Mooney 2009). Those topics remain unexplored in the field of invasive species in the herpetological literature, so some of the invasive species covered herein will focus our discussions on known impacts affecting ecological components. Finally, we adopt the term “biosecurity,” as defined by Pyšek and Richardson (2010), as the management of risks posed by organisms to the economy, environment, and human health through exclusion (prevention of initial introduction), mitigation, adaptation, control, and eradication.

Our taxonomic positions follow those discussed in Johnson et al. (2010), Porras et al. (2013), and expanded upon by Johnson et al. (2015b), which are predicated on modern phylogenetic principles. We regard species as separate evolutionary lineages and consider them the lowest evolutionary lineage segment placed on a phylogeny in a formal phylogenetically based taxonomy. We also consider the subspecies category to be a taxonomic anachronism that should not be used in a formal classification hierarchy because subspecies merely represent geographic variations in populations of the same species that are connected by gene flow (intergradation) and not separate evolutionary lineages. We do, however, concede that officially classifying subspecies had much influence in the past, so we reference them in a historical perspective in a few taxa discussed below. In those cases, the taxonomy reverts to the binomial.

Results and discussion

Status of the exotic and translocated herpetofauna of Middle America

Currently, 40 species of herpetofaunal species are considered introduced to a region of Middle America, or indigenous to a region, but translocated to non-native areas. Of these, 24 are exotics (Tables 1, 2), whereas 16 are translocated (Tables 1, 3). Most are reptiles (30 [18 exotics and 12 translocations], and ten are anuran amphibians [six exotics and four translocations]). Three species are listed among the 100 worst invasive alien species of the world (Lowe et al. 2000): the American Bullfrog (Lithobates catesbeianus), the Puerto Rican Coqui (Eleutherodactylus coqui), and the Pond Turtle (Trachemys scripta). Twenty-six species are listed in the IUCN Red List of Threatened Species (IUCN 2014), with most of those, as expected, in the Least Concern category. Two, however, are under risk categories: The Spiny Chuckwalla (Sauromalus hispidus) as Near Threatened, and the Mexican Giant Musk Turtle (Staurotypus triporcatus) as Lower Risk/Near Threatened. Mexico has the largest number of introduced species (24; 13 exotics and 11 translocations), whereas El Salvador only has two species, both exotic. In addition to Mexico, Honduras also has translocated species, such as Ctenosaura similis on Isla Roatán (Table 1).

Table 1.

List of introduced and translocated herpetofauna in the countries from Middle America. Introduced herpetofauna in Middle America (Right total = number of countries in which the species has “exotic” or “translocated” status; Bottom total = number of exotic and translocated species in that country). Parentheses enclose the number of exotic and translocated species for that taxon.

Taxa Exotic (E) or translocated (T) Mexico Guatemala Belize Honduras El Salvador Nicaragua Costa Rica Panamá Total Main references
Amphibians (10)
Anurans (10)
Eleutherodactylidae (4)
Eleutherodactylus antillensis E + 1 de Sousa et al. (1989), Barker and Rodríguez-Robles (2017)
Eleutherodactylus coqui E + Barrantes-Madrigal (2017), Barrantes-Madrigal et al. (2019)
Eleutherodactylus johnstonei E + 1 Ibañez and Rand (1990), McCranie and Valdez-Orellana (2014)
Eleutherodactylus planirostris E + + + + + 5 Crawford et al. (2011), McCranie and Gutsche (2014), Barquero and Araya (2016), Cedeño-Vázquez et al. (2014), Alvarez-Romero et al. (2008)
Hylidae (2)
Osteopilus septentrionalis E + 1 Savage (2002)
Smilisca baudinii T + 1 Recuero et al. (2004)
Pipidae (1)
Xenopus laevis E + 1 Murphy (1983) Álvarez-Romero (2008), Peralta-García et al. (2014)
Ranidae (3)
Lithobates berlandieri T + 1 Rorabaugh and Servoss (2006)
Lithobates catesbeianus T + 1 Casas-Andreu et al. (2001), Lemos-Espinal and Smith (2016), Grismer (2002)
Lithobates forreri T + 1 Grismer (2002)
Reptiles (31)
Crocodylia (1)
Crocodylidae (1)
Crocodylus moreletii T + 1 Alvarez-Romero (2008)
Squamata (23)
Dactyloidae (4)
Anolis allisoni E + + + 3 Charruau et al. (2015), Schmidt (1941), McCranie and Köhler (2015), Glor et al. (2005)
Anolis carolinensis E + 1 Terán-Juárez et al. (2015)
Ctenonotus cristatellus E + 1 Savage (2002), Mayer (2010)
Norops sagrei E + + + + + + 6 Lee (1996), Sexton and Brown (1977), Stuart (1955), Savage and Bolaños-Vives (2005), Batista et al. (2019)
Gekkonidae (9)
Gehyra mutilata E + 1 Álvarez-Romero (2008) Cruz-Sáenz et al. (2017), Reynoso (1990a), Lemos-Espinal and Dixon (2013)
Gekko gecko E + 1 Meerman and Garel (2002)
Hemidactylus frenatus E + + + + + + + + 8 Weterings and Vetter (2017)
Hemidactylus garnotii E + + 2 Savage (2002), Morales et al. (2017)
Hemidactylus haitianus E + + 2 McCranie (2015), Auth (1994)
Hemidactylus mabouia E + + + + 4 Álvarez-Romero (2008), Gutsche and McCranie (2009), Abarca and Monge (2007), Auth (1994)
Hemidactylus turcicus E + + 2 Martínez-Hernández et al., (2017), McCoy (1970), Lee (1996)
Lepidodactylus lugubris E + + + + 4 Smith and Grant (1961), Savage (2002), Hoogmoed and Avila-Peres (2015)
Tarentola mauritanica E + Ortíz-Mena et al. (2019)
Iguanidae (5)
Ctenosaura conspicuosa T + 1 Grismer (2002)
Ctenosaura pectinata T + 1 Aguirre-Léon and Matías-Ferrer (2017)
Ctenosaura similis T + 1 McCranie and Valdés-Orellana (2014)
Sauromalus hispidus T + 1 Grismer (2002), Petren and Case (1997)
Sauromalus varius T + 1 Hollingsworth et al. (1997)
Leiocephalidae (1)
Leiocephalus varius E + 1 Schwartz and Thomas (1975), McCranie (2018)
Phrynosomatidae (1)
Uta stansburiana T + 1 Upton and Murphy (1997)
Sphaerodactylidae (1)
Sphaerodactylus argus E + + + 3 Harris and Kluge (1984), Lee (1996), Sunyer et al. (2013), Thomas (1975)
Boidae (1)
Boa imperator T + 1 López-González (1991)
Typhlopidae (1)
Indotyphlops braminus E + + + + + + 6 Wallach et al. (2014), Leets-Rodríguez et al. (2019)
Testudines (7)
Chelydridae (1)
Chelydra serpentina E + 1 Dixon (2013), Powell et al. (2016), Terán-Juárez et al. (2016)
Emydidae (1)
Trachemys scripta T + 1 Lavín et al. (2014)
Kinosternidae (1)
Kinosternon integrum T + 1 Iverson et al. (1998)
Staurotypidae (1)
Staurotypus triporcatus T + 1 Terán-Juárez et al. (2015)
Testudinidae (1)
Chelonoidis carbonarius E + Salazar-Saveedra et al. (2015), Villa et al. (1988)
Trionychidae (1)
Apalone spinifera T + 1 Grismer (2002), McGaugh and Janzen (2008)
Totals 24E/16T 29 4 5 9 2 6 9 10

Herpetofaunal introductions in Middle America can be traced back to colonial times and were associated with commercial routes between the Philippines and New Spain (mainly Acapulco, Mexico), or through the slave trade from Western Africa to the Caribbean and Antillean islands, and from there into Central America. More recent events involved the opening of the Panama Canal in 1914, the expansion of the irrigation infrastructure after the 1950’s in northern Mexico, and throughout Middle America due to the highly popular pet trade and agricultural practices. On the other hand, translocations have more obscure origins, and certainly some of those could have occurred in pre-Columbian times, like translocations of iguanid lizards onto several islands in the Sea of Cortes (also called the Gulf of California, or Mar de Cortés in Spanish) by the Seri society. Whereas it is often accepted in invasive species biology that 1492 is the cutoff date for delineation between native and non-native species, we are concerned at this point with translocated species, not non-native species. In addition, it is our opinion that the year 1492, as the time when Cristopher Columbus “discovered” the New World, is of disputable significance from a biological point of view. The matter of most significant concern, we think, is to what extent humans, whether from Spain or elsewhere, have had a hand in the movement of creatures around the world.

We recognize six major Middle American sites as “hotspots” for herpetofaunal invasions (four of which are depicted on Fig. 1): 1) Northwestern Baja California and nearby Río Colorado delta in the Mexicali Valley, where hydrological systems and agricultural channels most likely served as pathways for the invasion of several introduced species already established in California, Arizona, and New Mexico; 2) The Panama Canal, where concentrated traffic of cargo shipping has been an important source of invaders; 3) The Mexican Yucatan Peninsula, a tropical region with flat topography and low elevation, with its highest point around 300 m, containing extensive communication and tourism infrastructures; 4) The State of Veracruz, historically having the largest and most important Mexican sea port, as well as being a main logistic center for Mexico’s economy; 5) Major airports and seaports of each Middle American country; and 6) Insular systems within the Sea of Cortes and Pacific Ocean in northwestern Mexico, where distributional patterns of several species can be explained only by natural or translocated dispersal overwater and island hopping; some translocations could have occurred there during pre-Columbian or even prehistoric times.

Figure 1. 

The four main Middle American sites considered as “hotspots” for herpetofaunal invasions: Northwestern Baja California and nearby Río Colorado delta in the Mexicali Valley, the Panama Canal, the Mexican Yucatan Peninsula, and The Papaloapan and Panuco basins in the Mexican state of Veracruz. The airports and seaports are not depicted due their ubiquity.

Finally, we recognize the following four major causes of introductions: 1) accidentals, mainly small species transported inadvertently by cargo vehicles, most frequently gekkonid lizards and anurans; 2) intentional releases, principally associated with pet trade and as food resources; most significantly chelonians and iguanid lizards, respectively; 3) escapees from the farming industry; mainly crocodiles and anurans, such as Morelet’s Crocodiles and American Bullfrogs; and 4) expanding invasion fronts when introduced naturalized species with high reproduction potential are well adapted to altered habitats. This fourth mechanism is especially relevant in anurans, such as those within the genus Eleutherodactylus and other species of original dispersers along invasion fronts.

The exotic herpetofauna of Middle America

AmphibiaAnura – Frogs

Family Eleutherodactylidae

Eleutherodactylus antillensis (Reinhardt & Lütken, 1863)

The Antilles Robber Frog is native to Puerto Rico, the Virgin Islands, and several associated islands and cays in that region. Numerous individuals apparently were introduced in the late 1950’s or early 1960’s into Panama City, probably through ornamental plants or intentionally released by a family after returning from a trip to Puerto Rico (Barker and Rodríguez-Robles 2017). Since then, this frog has spread throughout suburban and rural gardens, abandoned parcels, and pastures in the Panama City metropolitan area (Barker and Rodríguez-Robles 2017) (Table 2, Map 1).

Table 2.

Distribution of introduced amphibians and reptiles in Mexico and Central American countries.

Map 1. Eleutherodactylus antillensis. Map 2. Eleutherodactylus coqui.
Map 3. Eleutherodactylus johnstonei. Map 4. Eleutherodactylus planirostris.
Map 5. Osteopilus septentrionalis. Map 6. Xenopus laevis.
Map 7. Chelonoidis carbonarius. Map 8. Anolis allisoni.
Map 9. Anolis carolinensis. Map 10. Ctenonotus cristatellus.
Map 11. Norops sagrei. Map 12. Gekko gecko.
Map 13. Gehyra mutilata. Map 14. Hemidactylus frenatus.
Map 15. Hemidactylus mabouia. Map 16. Hemidactylus garnotii.
Map 17. Hemidactylus turcicus Map 18. Lepidodactylus lugubris
Map 19. Tarentola mauritanica. Map 20. Leiocephalus varius.
Map 21. Sphaerodactylus argus. Map 22. Indotyphlops braminus.

Eleutherodactylus coqui (Thomas, 1966)

Fig. 2

The Puerto Rican Coqui was first reported in Costa Rica by García-Rodríguez et al. (2010). According to them, it was a recent introduction, probably around the end of the 1990’s by a pet trader who carried six individuals from Puerto Rico and released them in Turrialba, where their descendants invaded surrounding localities despite attempts to exterminate them with poison; afterwards, survivors were sold as pets in other parts of Costa Rica. Besides Turrialba, however, the only other known established Costa Rican population is in nearby Juan Viñas (Barrantes-Madrigal 2017; Barrantes-Madrigal et al. 2019) (Table 2, Map 2). This is especially troubling since it has had extensive documented negative impacts on the native biota of Hawaii (Beard et al. 2002; Beard et al. 2009), and is listed as one of the 100 world’s worst invasive introduced species (Lowe et al. 2000). The control or eradication of this species is still possible, since its distribution apparently is still restricted to a few localities in Costa Rica (Barrantes-Madrigal 2017).

Figure 2. 

Eleutherodactylus coqui. Southern Florida. Photograph by Louis Porras.

Eleutherodactylus johnstonei (Barbour, 1914)

Johnstone’s Whistling Frog is native to the Lesser Antilles and has been introduced into several Caribbean areas, South America, the United States, and Europe (Savage 2002). This frog is a particularly proficient invader since it easily establishes breeding sites and males vocalize almost immediately after being released into the wild, thereby out competing native frogs (Kaiser and Henderson 1994). In Central America, it currently occurs in residential suburbs of Panama City, where it was believed to have been introduced in the mid-1980’s, together with introduced plants (Ibañez and Rand 1990) (Table 2, Map 3). Savage (2002) reported a single specimen collected in a city park in San José, Costa Rica, but it apparently was alone and failed to establish a viable population (Savage and Bolaños-Vives 2005). Due to the lack of more records, Sasa et al. (2010) omitted this frog in their account of the Costa Rican herpetofauna. We agree with their opinion.

Eleutherodactylus planirostris (Cope, 1862)

Fig. 3

The Greenhouse Frog is extremely small-sized (adults < 30 mm in length), is native to Cuba, Bahamas, and Cayman Islands (Crawford et al. 2011; Olson et al. 2012), and currently is found in Honduras in San Pedro Sula, La Paz, and Isla Guanaja, Islas de la Bahía (McCranie and Gutsche 2014). This frog has also been introduced into Puerto Limón, Costa Rica (Barquero and Araya 2016), the Miskito Cays of Nicaragua, in Panama, the Mexican Yucatan Peninsula (Cedeño-Vázquez et al. 2014; García-Balderas et al. 2016; Ortíz-Medina et al. 2017), and Veracruz, Mexico (Álvarez-Romero et al. 2008); the last authors noted that this species was first reported there in 1974; no other record existed from Veracruz until the phylogenetic analysis of Contreras-Calvario et al. (2018) concluded that the Greenhouse Frog population of the Gulf versant is related to Cuban populations, thus inferring a different colonization event from the E. planirostris on the Yucatan Peninsula, which is closer to Panama and Philippines populations (Cedeño-Vázquez et al. 2014). Recent reports, however, are found for Veracruz and Morelos in the iNaturalist platform (inaturalist.org 2018) (Table 2, Map 4). Environmental impacts produced by the diminutive Greenhouse Frogs need to be determined, since there has been no direct evidence for it being particularly harmful. Still, possible negative impacts include: predation on native invertebrates, competition for food with other insectivorous vertebrates, vulnerability to depredation that could limit its dispersal, but paradoxically it could become an abundant food source for other introduced species, thus facilitating their establishment and pending impacts (Olson et al. 2012).

Figure 3. 

Eleutherodactylus planirostris. Southern Florida. Photograph by Louis Porras.

Family Hylidae

Osteopilus septentrion,alis (Duméril & Bibron, 1841)

The Cuban Tree Frog is native to Cuba, Bahamas, and the Cayman Islands (Duellman 2001). In Central America, it is only known from Puerto Limón, Costa Rica, where it arrived in the mid 1980’s as an accidental introduction from cargo ships (Savage 2002) (Table 2, Map 5). Due to high adaptability to humid disturbed areas and its explosive breeding behavior, it has a high potential to spread quickly to other populated areas (Savage 2002).

Family Pipidae

Xenopus laevis (Daudin, 1802)

Fig. 4

The African Clawed Frog is native to “extreme southern Angola…south to Cape Region of Rep. South Africa thence east and north in savanna habitats through Zimbabwe and southeastern Zambia to Malawi” (Frost 2020). This frog is now a widespread invader in lentic waters, and its accidental or deliberate introductions are associated with uses as a laboratory animal or pet (Álvarez-Romero et al. 2008; Measey et al. 2012). Although we know now that Batrachochytrium is native to Asia (O’Hanlon et al. 2018), the earliest known presence of Batrachochytrium dendrobatidis fungal infection outside its native range came from X. laevis populations in Africa (Weldon et al. 2004). Thus, the association with global trade markets and frequent releasing of this frog into the wild was credited for initiating the chytridiomycosis epidemics (Kraus 2009). The frog’s presence in Mexico was documented first in the 1980’s from a single observation (Murphy 1983), probably a result of dispersing individuals from established populations in southern California (Peralta-García et al. 2014). Today, it is confined to the Tijuana and Ensenada region in northwestern Baja California (Lavín et al. 2014), with large populations reported at sites near Rosarito (Peralta-García et al. 2014) (Table 2, Map 6). Further range expansion of this species is highly probable, since the landscapes all along northern Baja California and northern Mexican Plateau contain suitable habitats (Measey et al. 2012). It is not clear, however, if by “Baja” Measey et al. (2012) meant the total Baja California peninsula or only the Mexican state of Baja California.

Figure 4. 

Xenopus laevis. Rosarito, Baja California, Mexico. Photograph by Anny Peralta.

Reptiles – Testudines – Turtles

Family Chelydridae

Chelydra serpentina (Linnaeus, 1758)

The Eastern Snapping Turtle is mentioned in the “Lista de las Especies Exóticas Invasoras para México” as present in the northern Mexican states of Coahuila and Chihuahua (SEMARNAT 2017), based on a distribution terminating at the Rio Grande as mapped by Stebbins (2003), who didn’t identify any verified records for Mexico. Several publications, including Lemos-Espinal and Smith (2007a, b), Lemos-Espinal and Smith (2016), Lemos-Espinal et al. (2017), and several chapters in Lemos-Espinal (2015) covering the US-Mexico border states alongside the Rio Grande, did not list C. serpentina as having substantiated records from Mexico. Terán-Juárez et al. (2016) discussed the probable presence of this species in the Río Grande basin across the border from Hidalgo County, Texas, and came to the conclusion that Eastern Snapping Turtles most likely inhabit the border region of south Texas and adjacent Mexico. Michael J. Forstner (personal communication) observed them in Tamaulipas, but didn’t take any voucher specimens. Dixon (2013) thought that the Hidalgo, Texas, and adjacent Tamaulipas turtles were introduced from allopatric native populations farther northeast in Texas, a pattern depicted on the map by Powell et al. (2016). Even though we do not have a map showing this species’ range in Mexico, we are under the impression that enough evidence now exists to consider Eastern Snapping Turtles in Tamaulipas as exotic in Mexico, being introduced originally from native Texas populations to the north.

Cupul-Magaña and Rubio-Delgado (2003) reported an Eastern Snapping Turtle from Puerto Vallarta, Jalisco, which they considered either a released individual or an escaped pet. Cruz-Sáenz et al. (2017) apparently did not think that the Puerto Vallarta individual was a member of an established population and did not list this species for Jalisco; we agree with that evaluation.

Family Testudinidae

Chelonoidis carbonarius (Spix, 1824)

The Red-footed Tortoise´s original distribution range extends from central Panama, through Colombia and the Atlantic versant of the Amazonas in Brazil, as far south as Paraguay and northern Argentina (Köhler 2008). This species was listed originally as part of the Nicaraguan herpetofauna by Villa et al. (1988), who indicated that J. Villa found a single individual on Big Corn Island in 1964. They also mentioned the testimony of a local inhabitant who claimed to have owned a specimen of this species in her childhood, 50 years prior to the interview. If this testimony is true, the red-footed tortoise would have been on Big Corn Island by the end of the 1930’s. Thus, this tortoise was known on Great Corn Island by the second half of the 20th century, but since the other known specimens were individuals held in captivity or kept as pets, the Red-footed Tortoise was omitted from the Nicaraguan herpetofauna by subsequent listings (e.g., Köhler 1999, 2001; Ruíz-Pérez 1996). Salazar-Saveedra et al. (2015), however, reported a breeding wild population on Great Corn Island, and pointed out that this population could be of exotic origin (Table 2, Map 7). They also stated that the known records of C. carbonarius on mainland Nicaragua (departments of Nueva Segovia and Masaya) might have originated from the Corn Islands. Sunyer and Martínez-Fonseca (2015) accepted the Red-footed tortoise as a member of the Nicaraguan herpetofauna and remarked that this species is alien in the country.

Reptiles – Squamata – Lizards

Family Dactyloidae

The anole family Dactyloidae contains the second highest number of introduced species (four species) in Middle America, and has exotic members distributed in all countries except El Salvador and Nicaragua. Norops sagrei is the most widespread anole in Middle America.

Anolis allisoni (Barbour, 1928)

Fig. 5

Allison’s Anole is listed by Álvarez-Romero et al. (2008) as being exotic in Mexico. Lee (1996) considered its occurrence on Cozumel Island doubtful, and González-Sánchez et al. (2017) reported that the only populations of this species in Mexico occur on the cays of Banco Chinchorro, Quintana Roo. Anolis allisoni also can be found on other Caribbean islands, such as Half Moon Cay, Belize (Schmidt 1941) and on Islas de Barbareta, Guanaja, Morat, Roatán, Utila, and Cayos Cochinos within the Islas de la Bahía complex in Honduras, as well as near the northern coastal regions, such as in La Ceiba, Atlántida (McCranie and Gutsche 2009), on the Honduran mainland; populations from Utila and La Ceiba might be recent introductions (McCranie and Köhler 2015) (Table 2, Map 8).

Figure 5. 

Anolis allisoni. Banco Chinchorro Atoll, Mexico. Photograph by Víctor H. González-Sánchez.

Recent evidence (Glor et al. 2005) indicated that A. allisoni is native to Cuba. The Mexican and Honduran populations have little variation in genetic distances when compared to Cuban populations, which suggests recent introductions onto the offshore islands of Mexico and Central America, as well as mainland areas of Honduras. In addition, McCranie and Gutsche (2009) indicated that the A. allisoni population (first mainland record for Honduras) in the Caribbean port of La Ceiba is a recent introduction from the Islas de la Bahía.

Anolis carolinensis (Voigt, 1832)

Presence of the Green Anole in Mexico was suggested by Conant and Collins (1998), because of a supposed record from Tamaulipas, but no evidence was provided. Álvarez-Romero et al. (2008) advocated that at least one population in Tamaulipas might be native, but they did not provide any specific locality or give any justification for that reasoning. Farr (2015) did not list the Green Anole from Tamaulipas. The occurrence of this species in Tamaulipas remained controversial until Terán-Juárez et al. (2015) reported photographic records from a courtyard in Ciudad Valle Hermoso where A. carolinensis had been observed for at least ten years; they also indicated its presence in the city of Matamoros. We consider A. carolinensis an exotic species in Mexico, based on Dixon’s (2013) assertion that records from Texas border counties were probable introductions and that its known allopatric native distribution lies to the north of the border region in Texas (Powell et al. 2016; map only) (Table 2, Map 9).

In Belize, the Green Anole is only known from a single specimen collected in 1966 on Half-Moon Cay (UF 23924); no other records were reported thereafter (Lee 1996), although some lizards sighted in 1996 resembled A. carolinensis (Platt et al. 1999). This last claim should be treated with caution, as the very similar-looking A. allisoni also occurs on that cay, and can be easily misidentified. Lee (1996) hinted that if the Green Anole ever occurred on Half-Moon Cay, it might be extirpated now due to displacement by A. allisoni. Stafford et al. (2010) did not list A. carolinensis for Belize, so we adopt this view by not recognizing this species as presently having an established population in that country.

Ctenonotus cristatellus (Duméril & Bibron, 1837)

Fig. 6

The Crested Anole is native to Puerto Rico and the Virgin Islands (Köhler 2008). In Central America, it has been established only in Costa Rica, where it has been observed in Puerto Limón (Savage 2002), Cahuita (Mayer 2010), Guayacán, and Valle de Rosas, Limón province, and in Turrialba, Cartago province (Savage 2002). This anole was first recorded in 1970 from Limón (Mayer 2010) and Fitch (1975) noted an explosive population increase of these anoles in a Limón city park and wondered if colonization by C. cristatellus might have caused the extirpation of Gonatodes albogularis in that park (Table 2, Map 10).

Figure 6. 

Ctenonotus cristatellus. Southern Florida. Photograph by Louis Porras.

In Mexico, C. cristatellus was reported from states on the Yucatan Peninsula, first by a single record from Cozumel, Quintana Roo, although Lee (1996) considered its presence there doubtful. Colston et al. (2015) reported Norops cristatellus from Calakmul, Campeche, but González-Sánchez et al. (2017) did not list it for the region because they thought the record needed verification, especially since Calakmul is a popular study site for working herpetologists and no other records are known from there. We concur that this species does not have an established population on Cozumel or Calakmul, so we remove it from the list of introduced species in Mexico.

Norops sagrei (Duméril & Bibron, 1837)

The Cuban Brown Anole is native to Cuba, the Bahamas, and Cayman Islands, but it is unclear if all populations in Jamaica were introduced, or if some were native (McCranie and Köhler 2015). It is important to note that the taxonomic status and identity of this species remain unclear for many populations in Mexico and northern Central America, since the validity of the “native” subspecies N. s. mayensis seriously has been questioned (González-Sánchez et al. 2017). Moreover, the description of this subspecies was made from a single population from Isla Polao, in the region of Laguna de Términos, Campeche, and supported only by morphological characters (Smith and Burger 1949). Anoles as a group can have much geographic variability in morphology, as exemplified by McCranie and Köhler’s (2015: 169) statement, “using dewlap color as a diagnostic character for any N. sagrei complex population (including isolated island populations) might not be informative.” In any case, considering the unlikelihood that N. s. mayensis is a valid taxonomic lineage (see our taxonomic positions above in the methods section), and because of the long history of N. sagrei colonizing Mexico and Central America, any defining characteristics of a separate evolutionary lineage (=species) were probably eliminated by genetic intergradation with other alien N. sagrei populations. Because of this and the close association of this species with human mediated disturbed habitats, we regard the N. sagrei complex, with one exception, as a single exotic species within the region. The exception was the recent resurrection by McCranie and Köhler (2015) of N. nelsoni, an allopatric N. sagrei complex species from the Islas del Cisne, Honduras.

The naturalized distributional range in Mexico of N. sagrei comprises all the inland regions and several insular systems off the Yucatan Peninsula (González-Sánchez et al. 2017; Lee 1996), Tabasco (Lee 1996), Ciudad Altamira, Tamaulipas (Terán-Juárez et al. 2015), and Minatitlán and Catemaco, Veracruz (Zamora-Abrego et al. 2006). Specimens labeled Anolis sagrei in the Zoological Collection of El Colegio de la Frontera Sur (ECOSUR), in San Cristóbal de las Casas, Chiapas (Muñoz-Alonso 2006), are from Ocozocoautla de Espinosa (Coita), Chiapas, which is located in the semi-arid Central Depression region. Johnson et al. (2010, 2015a) did not report that locality and no other records are known to exist in Chiapas.

In Central America, the Cuban Brown Anole occurs throughout Belize (Lee 1996) and adjacent Caribbean lowlands of Guatemala (Stuart 1955). In Honduras, this species is known to occur at Puerto Cortés and San Pedro Sula, Cortés, and at La Ceiba and Tela, Atlántida, on the northern mainland and on the islands of Roatán and Utila (McCranie and Köhler 2015). An introduced population also exists in the vicinity of Limón, Costa Rica (Savage and Bolaños-Vives 2005). Batista et al. (2019) reported established populations of N. sagrei at several sites within and around Panama City. They also mentioned that those populations might have become established approximately five years ago, and arrived there as released pets, or more probably, from shipments arriving at the port of Balboa (Table 2, Map 11).

Family Gekkonidae

Not surprisingly, the geckos contain the highest number of introduced species (10 species) for the region, distributed among two families (i.e., Gekkonidae and Sphaerodactylidae) in Middle America (Table 1). Because of their notorious colonization ability, geckos are frequently referred to as “weedy” species (Kluge 1969).

Hemidactylus is the most speciose genus, comprising five introduced species (H. frenatus, H. garnotii, H. haitianus, H. mabouia, and H. turcicus). At least one of these species occurs in each country of Middle America, but only H. frenatus is present in all seven (Table 2). Hemidactylus is a species-rich genus (167 species; Uetz et al. 2020), native to tropical areas of Asia and Africa and the Mediterranean region, most of which species have small native distributional ranges. A few species, however, can be found almost worldwide in tropical and subtropical areas due to either human intervention or possibly by having undertaken long transmarine journeys (Carranza and Arnold 2006).

Gekko gecko (Linnaeus, 1758)

Fig. 7

The Tokay Gecko is very well known due to its use in scientific research (Roesler et al. 2011), and popularity in the pet trade (Kraus 2009). It was originally native to eastern India, Nepal, Southeast Asia, China, and the Malayan Archipelago (Roesler et al. 2011). Stafford et al. (2010) listed this species as exotic in Belize. The population was reported from South Water Caye, a small sandy caye measuring 8.2 ha on the Belizean barrier reef (Meerman and Garel 2002) (Table 2, Map 12). Apparently, the introduction of the Tokay Gecko occurred around 1994, when a tourism industry worker brought several individuals to South Water Caye and intentionally released them. The introduction of G. gecko and the declining numbers of Aristelliger georgeenis and Phyllodactylus tuberculosus on that island might be related (Meerman and Garel 2002).

Figure 7. 

Gekko gecko. Southern Florida. Photograph by Louis Porras.

Gehyra mutilata (Wiegmann, 1834)

The Stump-toed Gecko is native to the Pacific basin region of Southeast Asia, where it has dispersed among Indian and Oceanic islands since the times of pre-Polynesian navigators (Fisher 1997). Rocha et al. (2009) suggested that G. mutilata is a complex of two cryptic lineages, one of them involving the resurrection of G. insulensis for several, if not all, Pacific islands, which includes the former Mexican, but now French Isla Clipperton (Isla de la Pasión). Lorvelec and Pascal (2006) hypothesized that the time and source of invasion on that atoll could have been during the 1950’s from Mexican Pacific ports in Nayarit or Sinaloa, but that allegation was merely speculation. If true, however, Mexican populations on the Pacific side would correspond to G. insulensis. Since Rocha et al. (2009) did not include Mexican samples in their study, we retain the name G. mutilata until more evidence is provided.

The date for the introduction of G. mutilata into Mexico is unknown, although Lemos-Espinal and Dixon (2013) remarked that before the advent of aviation, this species was restricted almost exclusively to the Philippines, although specimens collected in Nayarit by the end of the 19th century already existed (GBIF.org 2018). In addition, Ineich and Blanc (1987) hypothesized that it could have been present in Mexico during the 18th Century by way of maritime trade between New Spain and the Philippines. In Mexico, it is now known from the Pacific versant in Sinaloa, Nayarit, Guerrero, and Chiapas (Álvarez-Romero et al. 2008), Jalisco (Cruz-Sáenz et al. 2017), and Baja California Sur (Reynoso 1990a; Lovich et al. 2009), and some Pacific Islands, such as Isabel (Quijada-Mascareñas and Canseco-Márquez 2007). On the Atlantic versant, established populations have been reported only from Veracruz (Álvarez-Romero et al. 2008) and Ciudad Valles, San Luis Potosí (Lemos-Espinal and Dixon 2013) (Table 2, Map 13).

Hemidactylus frenatus (Duméril & Bibron, 1836)

Fig. 8

The Common House Gecko is a well-known successful colonizer of urban environments (Lee 1996). Even though its native range is uncertain, it probably can be restricted to southern India, Sri Lanka, Burma, southern China, Malayan Peninsula, and Philippines (Savage 2002). As an introduction in the Americas, it occurs on both versants from Florida and California through Mexico and Central America to Brazil (Weterings and Vetter 2017), from sea level to 1,545 m elevation (Mata-Silva et al. 2013). Introductions of this gecko possibly could lead to competitive exclusion of native gecko populations and to extinction of insular endemics (Cole et al. 2005).

Figure 8. 

Hemidactylus frenatus. León, Nicaragua. Photograph by Javier Sunyer.

It is believed that H. frenatus arrived on the North American continent around the 16th century by means of maritime commerce between the Philippines and Acapulco, Mexico (Álvarez-Romero et al. 2008). Farr (2011), however, argued that the first records of this species date to the end of the 19th century or even as late as the 1930’s. If so, colonization during the colonial period, by an introduced lizard common in hotels of Acapulco today, should have been recorded early on by the first collectors visiting Mexico, but since there are no records from then, introductions into Central American countries might have been even more recent. For example, in Costa Rica the first reports of H. frenatus were made after 1990, according to Savage (2002). This exotic species also has been reported from all other Central American countries and many states in Mexico (Uetz et al. 2020). Hemidactylus frenatus first was recorded from peninsular Baja California Sur, Mexico, by Reynoso (1990b) from the city of La Paz; Grismer (2002) also observed an established population in Loreto, located north of La Paz on the peninsula. Most recently, the species was reported for the first time from any island in the Sea of Cortes (Isla El Pardito) by Dayton et al. (2020) (Table 2, Map 14). Mata-Silva et al. (2013) found this species to be very common in Oaxaca City, Oaxaca at an elevation of 1,545 m.

Hemidactylus garnotii (Duméril & Bibron, 1836)

The Indo-Pacific or Garnot’s House Gecko, is native to the Indo-Pacific basin, and is widespread on several islands in Oceania, Asia, and the Pacific Ocean. In Middle America, it was introduced at several ports and urban centers in Costa Rica (Savage 2002; Köhler 2008) and in Guatemala, Guatemala (Morales et al. 2017). This species is a parthenogenetic all-female species and, therefore, it seems to have no social hierarchy (Frankenberg 1982); it is a territorial and aggressive species (Frankenberg 1984), and successful colonizer of urban and other anthropogenic habitats, with high potential for expansion (Savage 2002). The first record in Middle America was from San José, Costa Rica, in 1992 (Savage 2002), whereas in Guatemala its introduction seems to be more recent, but from an unknown source (Morales et al. 2017) (Table 2, Map 16).

Hemidactylus haitianus (Meerwarth, 1901)

The Haitian House Gecko has had a complex and unclear taxonomic history. Traditionally, it was considered two different taxa: the Middle American and West Indian populations of H. brooki haitianus, type locality marked as “Haití, Port-au-Prince” (Powell and Maxey 1990), and H. angulatus, type locality “West Coast of Africa” (Hallowell 1854). Powell et al. (1996) considered H. brooki haitianus a full species (H. haitianus) for lizards native to the West Indies, including Cuba and Puerto Rico. Further revisions revealed H. haitianus as nested inside a larger clade that includes the African populations of H. brooki along with H. angulatus (Carranza and Arnold 2006; Weiss and Hedges 2007; Bauer et al. 2010; Rösler and Glaw 2010). Weiss and Hedges (2007) and Gamble et al. (2011) reported virtually no genetic divergence among the populations of this complex in the Greater Antilles, and clustered them with African populations, thus corroborating the introductory origin of H. haitianus for the Antilles. The source of invasion to the New World might have been through the slave trade between western Africa and the West Indies (Weiss and Hedges 2007). Consequently, the populations of H. brooki in Honduras (McCranie 2015) and Panama (Auth 1994) and those of H. angulatus in South America might have had their origins by following colonial trade routes between the Antilles and Middle America, although a date has not been suggested. Even if the H. angulatus-brooki complex is not fully resolved, the name H. haitianus, as suggested by Bauer et al. (2010), is used herein for individuals occurring throughout the Western Hemisphere.

Hemidactylus mabouia (Moreau De Jonnès, 1818)

Moreau’s Tropical House Gecko has no clear-cut geographic origin. The type locality is marked as “Antilles,” and restricted to St. Vincent Island by Stejneger (1904). Nonetheless, its actual origin was without doubt on the African continent, where it is widespread, ranging from southern Africa northward to Liberia and Ethiopia (Álvarez-Romero et al. 2008). An interesting hypothesis suggests arrival of this species in the New World by accidental transport on slave ships along routes from Africa to the West Indies and South America. There is no full concordance, however, between slave ship routes and the distribution of H. mabouia (Kluge 1969). Based on the long history of maritime trade between Africa and the Antilles and the 1818 description of the species, it can surely be said that the type specimen from the Antilles represents an introduced population. Moreover, the introductions in Mexico and Central America might be the result of maritime trade from the West Indies (Álvarez-Romero et al. 2008). Currently, introduced populations of H. mabouia are found in Veracruz (Ochoa-Ochoa et al. 2006; Álvarez-Romero et al. 2008), and Tamaulipas, Mexico (Sosa-Tovar, et al. 2019), Islas de la Bahía, Honduras (Gutsche and McCranie 2009), San José, Costa Rica (Abarca and Monge 2007), and Panama (Auth 1994) (Table 2, Map 15).

Hemidactylus turcicus (Linnaeus, 1758)

The Mediterranean House Gecko is native to coastal areas of the Mediterranean, where it is widespread across southern Europe, the Levant, and more sporadically in North Africa (Martínez-Hernández et al. 2017). In Mexico, it is known from many states and places, including: Yucatan Peninsula, Baja California, Sonora, Sinaloa, Chihuahua, Coahuila, Durango, San Luis Potosí, Aguascalientes, Ciudad de Mexico, Morelos, Nuevo León, Puebla, Tamaulipas, Oaxaca, Chiapas (Martínez-Hernández et al. 2017), and Querétaro (Tepos-Ramírez, 2019). Its first introduction probably occurred around Acapulco, from colonial-period trade with inhabitants of the Pacific islands (Álvarez-Romero et al. 2008), although another possible source was from Veracruz, where H. turcicus was recorded in 1895. Subsequently, H. exsul was described in 1906 from Progreso, Yucatán, which in fact was based on a specimen of H. turcicus (McCoy 1970). In Panama, it is known from the Canal Zone (McCoy 1970), where the introduction took place after the opening of the Panama Canal in 1914. Stafford et al. (2010) suggested the possible presence of this species in Belize, but we were unable to find records on GBIF.org (2018), or in the available literature, so it is not recognized herein for that country (Table 2, Map 17).

Lepidodactylus lugubris (Duméril & Bibron, 1836)

Fig. 9

The Mourning Gecko is native to southeast Asian and Indo-Australian regions, and currently is distributed worldwide in the tropics from sea level up to 700 m elevation (Köhler 2008). Probably much of its dispersal potential comes from being a complex of parthenogenetic lineages that includes diploid and triploid forms because of hybridization between L. moestus and an undescribed species (Fujita and Moritz 2009). In the Americas, it was first reported in the mid 1950’s at Ft. Clayton, in the Canal Zone, Panama (Smith and Grant 1961). Even though prior collected specimens exist from the 1910’s in that area, it is also known from Bocas del Toro, Panamá (Hoogmoed and Avila-Pires 2015), Golfo Dulce, and Peninsula de Osa and Punta Arenas departments, both from the Pacific versant of Costa Rica (Savage 2002). This species is known also in Costa Rica from the Gandoca-Manzanillo Wildlife Refuge, Talamanca, Limón, and from Tirimbina Biological Reserve and La Virgen of Sarapiquí, Heredia (Jiménez and Abarca 2014). This gecko is established in Nicaragua on the southeastern Caribbean coast at elevations lower than 10 m (Hoogmoed and Avila-Pires 2015), and on the Corn Islands (Sunyer et al. 2013), where individuals probably arrived on cargo ships sometime around 1975 at Bluefields and/or Great Corn Island (Henderson et al. 1976). Hoogmoed and Avila-Pires (2015) thought that the Mourning Gecko should not be listed for Mexico because of inconsistences in the literature and lack of vouchered museum specimens. A recent (November 2017) photographic record for the port of Mazatlán, Sinaloa, however, exists in the iNaturalist platform (GBIF.org 2018). In addition, Ahumada-Carrillo and Weatherman (2018) found a recent established population in Puerto Vallarta, Jalisco, Mexico (Table 2, Map 18).

Figure 9. 

Lepidodactylus lugubris. Big Corn Island, Nicaragua. Photograph by Javier Sunyer.

Tarentola mauritanica (Linnaeus, 1758)

Fig. 10

The Moorish Gecko is native to the European and North African Mediterranean basin (Rato et al. 2015). Ortíz-Medina et al. (2019) recently reported this species from cargo containers stored in a warehouse in Progreso, Yucatan, Mexico. They mentioned that six to eight “unusual-looking lizards” were sighted there in early 2017, but only two specimens were captured and identified as T. mauritanica. The population in Progresso, however, was supposedly established, which is most likely factual, considering the number of individuals observed at the site, in addition to the well-known success of gekkonid lizards for becoming established species. Another record is known from Guadalajara, Mexico, through a photographic entry in inaturalist.org (2018), but no other records of the Moorish Gecko exist for Mexico, and no validating museum vouchers could be found for Guadalajara (Table 2, Map 19). We have not overlooked the recent evidence suggesting that T. mauritanica constitutes a species complex (Rato et al. 2016), but choose to maintain the traditional nomenclature while awaiting full resolution of the group.

Figure 10. 

Tarentola mauritanica. Progreso, Yucatan, Mexico. Photograph by Javier Ortíz-Medina.

Family Leiocephalidae

Leiocephalus varius (Garman, 1887)

The Cayman Curly-tailed Lizard is native to the Grand Cayman Islands (Garman 1887; Echternacht et al. 2011) and was first reported on the Swan Islands, Honduras, in the mid 1970’s (Schwartz and Thomas 1975), where it was conspicuous on human buildings and other places on Big Swan Island (McCranie et al. 2017). While some authors listed it as native to Honduras (Townsend and Wilson 2010; Solís et al. 2014), McCranie et al. (2017) suggested that this lizard was introduced recently to the Swan Islands, although circumstances or the date of the introduction were not provided. We note that McCranie (2018) recently elevated L. varius to a full species from L. carinatus varius, therefore, L. varius replaces L. carinatus as the exotic species on the Swan Islands (Table 2, Map 20).

Family Sphaerodactylidae

Sphaerodactylus argus (Gosse, 1850)

The Ocellated Dwarf Gecko is native to Jamaica, Cuba, and adjacent islands in the Bahamas and Antilles, including the Colombian Isla de San Andrés (Harris and Kluge 1984). It was introduced into the northern Yucatan Peninsula of Mexico (Lee 1996), Isla del Maíz Grande (Corn islands), Nicaragua (Thomas 1975; Sunyer et al. 2013), and on several islands on the Bocas del Toro and San Blas Archipelagos in Panama (Harris and Kluge 1984; Savage 2002). Approximate dates of introductions on the islands of Central America are unknown, but before Thomas (1975) no previous record for Middle America was available.

Records of S. argus from the northern coast of Yucatan (Lee 1996) might be the result of active maritime commerce in the region. Furthermore, this gecko was mentioned as occurring on the insular systems of Costa Rica (Lee 1996), but no evidence or citation was provided. Sphaerodactylus argus has not been reported on mainland Costa Rica (Savage 2002; Savage and Bolaños-Vives 2005; Köhler 2008; Sasa et al. 2010), although its close proximity to the Bocas del Toro islands could lead to future colonization of mainland areas adjacent to those islands (Savage 2002). Sphaerodactylus argus should not be confused with what was formerly known as S. argus continentalis (now S. continentalis), which ranges from the Isthmus of Tehuantepec, Mexico, into central Honduras (McCranie and Hedges 2012) (Table 2, Map 21).

Reptiles – Squamata – Snakes

Family Typhlopidae

Indotyphlops braminus (Daudin, 1803)

Fig. 11

The Brahminy Blindsnake is the most widespread alien reptile in the world (Capinha et al. 2017). This snake is a small-sized (mean total length < 130 mm) brown to black-colored species (Wallach 2009), with a secretive fossorial lifestyle occupying soil and leaf litter (Álvarez-Romero et al. 2008), and can easily be confused with earthworms (Wallach 2009). The wide distributional range of this snake can be explained by the ease with which this snake is carried inadvertently within root masses of potted plants being shipped world-wide by the garden industry (Álvarez-Romero et al. 2008). The reproductive characteristics of this species (unisexual, triploid, and parthenogenetic) also allows a single individual to establish a new population (Vitt and Caldwell 2014).

Figure 11. 

Indotyphlops braminus. Ixil, Yucatan, Mexico. Photograph by Luis Díaz-Gamboa.

The type locality is reportedly the region of Coromandel, southeast India (Wallach et al. 2014). The center of origin of I. braminus is difficult to discern with precision, however, due to its almost cosmopolitan distribution (Wallach et al. 2014; Capinha et al. 2017), previous unclear taxonomy, frequent misidentifications (Wallach 2009), and a long history of exceptional dispersal ability. The only certain location for the center of origin is that it should be an undetermined site in the Old-World tropics (Álvarez-Romero et al. 2008). This snake also has a hybrid origin with the parental species still undetermined (Vitt and Caldwell 2014). Currently, it is widespread in Mexico, Guatemala, Belize, Honduras, and El Salvador (Wallach et al. 2014; Lee 1996). In Nicaragua, it is known from a recent record (Leets-Rodríguez et al. 2019) from a locality near Managua. Due the nature of the record (five specimens found accidentally in a suburban backyard), it is probable that the Brahminy Blindsnake is widespread in urban and suburban areas of Managua. Arrival in Mexico was most likely sometime during the 16th century via maritime trade between the Philippines and New Spain at the Acapulco port (Álvarez-Romero et al. 2008). Care should be taken not to confuse I. braminus with other local members of Typhlopidae inhabiting the American tropics (Table 2, Map 22).

The translocated herpetofauna of Middle America

Several species of amphibians and reptiles found in a region are translocated when introduced into areas of the same region outside their native ranges. We will consider only those with sound evidence of having been translocated by human activities and that have established populations (Table 1).

AmphibiaAnura – Frogs

Family Hylidae

Smilisca baudinii (Duméril & Bibron, 1841)

The Mexican Treefrog ranges from “Extreme southern Texas (USA) and southern Sonora and southwestern Chihuahua (Mexico) south (including the Balsas Depression of Mexico) in tropical lowlands to Costa Rica on the Pacific slope; including the Tres Marias Islands off the coast of Nayarit, Mexico” (Frost, 2020). This species also can be found on other Mexican Pacific Islands (Woolrich-Piña et al. 2016), but historically was unknown on the Baja California Peninsula (Grismer 2002; Lovich et al. 2009). Records of S. baudinii exist from near the village of Todos Santos, Baja California Sur, which could represent an unlikely relictual population, or a more reasonably explained accidental translocation from any of the mainland ferry ports where this frog naturally occurs (Recuero et al. 2004) (Table 3, Map 23).

Table 3.

Non-native distribution of translocated amphibians and reptiles in Mexico and Central America.

Map 23. Smilisca baudinii Map 24. Lithobates berlandieri
Map 25. Lithobates forreri Map 26. Apalone spinifera
Map 27. Ctenosaura conspicuosa Map 28. Ctenosaura pectinata
Map 29. Ctenosaura similis Map 30. Sauromalus hispidus
Map 31. Sauromalus varius Map 32. Uta stansburiana
Map 33. Boa imperator

Family Ranidae

Lithobates berlandieri (Baird, 1859)

The native range of the Rio Grande Leopard Frog extends from southeastern New Mexico and central Texas southward into Mexico (Stebbins 2003) along the Gulf lowlands into the northern half of Veracruz (Zaldı́var-Riverón et al. 2004). It is unclear what the status is for populations in the Mexican portion of the lower Colorado Basin, where Grismer (2002) reported a sighting of what he believed to be an individual L. berlandieri at the confluence of the Hardy and Colorado rivers in the Mexicali Valley, Baja California. Photographic evidence of this species near San Luis Río Colorado, Sonora, Mexico, was reported by Rorabaugh and Servoss (2006) from a concrete-lined ditch passing through an agricultural field. The invasion front originated somewhere in southwestern Arizona where the Colorado and Gila rivers meet, which is the same area where L. berlandieri was collected in a survey in 1981 (Platz et al. 1990). The most probable act of introduction into Mexico was not a single event, but rather a secondary effect of several fish transplant operations into the Yuma, Arizona region from New Mexico in the late 1960’s or early 1970’s (Platz et al. 1990). It is likely that the Mexican populations, if they are established, originated as an expansion of the invasion front, using the Colorado and Gila River systems, and adjacent agricultural canals as dispersal routes (Rorabaugh et al. 2002). It is unknown what the impact of L. berlandieri is on the biodiversity of Baja California, but it has been associated with historical declines of populations of other native leopard frogs, such L. yavapaiensis, in areas of the lower Rio Colorado basin in Arizona (Rorabaugh et al. 2002) (Table 3, Map 24).

Lithobates catesbeianus (Shaw, 1802)

Fig. 12

The American Bullfrog originally ranged from southeastern Canada and central and eastern United States into northeastern Mexico (Conant and Collins 1998). It has a long and extensive history of introductions into Mexico, with first reports made near Cadereyta, Nuevo León, in 1853 (Ramos-Guerra and Gatica 2014). Since then, feral populations have been established in many Mexican states and places, including Chihuahua, Durango, and San Luis Potosí (Lemos-Espinal and Smith 2016), Sinaloa, Sonora, Morelos, Ciudad de Mexico, Puebla, San Luis Potosí (Casas-Andreu et al. 2001), Aguascalientes (Avila-Villegas et al. 2007), Baja California (Grismer 2002) and Hidalgo (Ramírez-Bautista et al. 2014), where it is probably linked to the extirpation of L. yavapaiensis and Incilius alvarius, and declines in Hyliola cadaverina (as Hyla regilla) and Thamnophis hammondii in several oases on the Baja California Peninsula (Grismer 2002). Apparently, American Bullfrogs were introduced intentionally in Costa Rica (Savage and Bolaños-Vives 2005), although the previously known population in La Garita is now thought to be nonexistent (IUCN 2015) (Table 3, Map 25).

Figure 12. 

Lithobates catesbeianus. Near Chihuahua International Airport, Chihuahua, Mexico. Photograph by Rubén A. Carbajal-Márquez.

Due to a high reproductive rate and generally destructive behaviors, L. catesbeianus has great potential for being harmful to native species, and has already been associated with declining and disappearing populations of native amphibians around the world (Casas-Andreu et al. 2001). This frog is considered one of the 100 world’s worst invasive introduced species (Lowe et al. 2000). Also of special concern is the relationship of American Bullfrogs with deadly pathogens, such as Batrachochytrium dendrobatidis and Ranavirus (Schloegel et al. 2009; Kolby et al. 2014). Recently, an outbreak of Ranavirus was reported in captive American Bullfrogs from a farm in Guasave, Sinaloa, in northwestern Mexico (Saucedo et al. 2019). Presently, this pathogen seems not to have spread into wild amphibian populations in Sinaloa, but the risk of Ranavirus becoming widespread is high, since there are several susceptible frog species in that area (Saucedo et al. 2019).

Lithobates forreri (Boulenger, 1883)

Forrer’s Leopard Frog’s native distribution was considered to be on the mainland Pacific versant of Mexico from Sonora (Zaldı́var-Riverón et al. 2004) into Costa Rica (Savage 2002). Grismer (2002) reported an introduced population in the water systems near Rancho San Juanito within the La Presa region, 100 km north of La Paz, Baja California Sur, suggesting to us a probable intentional translocation from mainland ferry ports across the Sea of Cortes from 1991 to 1993. Those dates were based on personal correspondence between a local rancher and L. L. Grismer (Table 3, Map 25).

Reptiles – Crocodylia – Crocodiles

Family Crocodylidae

Crocodylus moreletii (Duméril & Bibron, 1851)

Fig. 13

Morelet’s Crocodile originally ranged only along the Atlantic lowlands of Middle America, from Tamaulipas, Mexico, to northern Guatemala and adjacent Belize (Cedeño-Vázquez et al. 2012). This crocodile formerly was considered an endangered species and subject to strict conservations measures. Fortunately, in the last few decades, significant recovery of populations has occurred within its native range. There has been an increase in the number of sites dedicated to its conservation by captive breeding, but also for exploiting the animals for food and hides; unfortunately, these farms are mainly found on the Pacific versant of Mexico outside its native range (Álvarez-Romero et al. 2008). As a negative side issue, in Mexico there have been several incidences of C. moreletii escaping from these farms into the wild, with populations being established primarily in the states of Oaxaca (Lagunas de Chacahua), Sinaloa, and Colima (Laguna de Alcuzahue) (Álvarez-Romero et al. 2008; Lavín et al. 2014). The first documented case took place during the 1970’s when several Morelet’s Crocodiles were taken from Tabasco to Lagunas de Chacahua, Oaxaca, in order to establish a hide factory there, but after a few years, the project was abandoned and several individuals escaped into the wild (Serrano-Gómez et al. 2016). In addition, the population near Villa Flores in the Central Depression of Chiapas could be an intentional translocation from areas to the north of there (Álvarez-Romero et al. 2008), although that locality is higher up on the Gulf versant with potential riverine access to the lowlands, at least in the past.

Figure 13. 

Crocodylus acutus. Hybrid pattern (see tail) with C. moreletii. APFF Yum Balam, Quintana Roo, Mexico. Photograph by Julio César Gutiérrez-Ramírez.

The invasion of aquatic habitats by C. moreletii might have serious consequences for native biotic communities, since it is a large top predator (Álvarez-Romero et al. 2008). Although C. moreletii and C. acutus (American Crocodile) are sympatric in some areas of their native ranges, in places were C. moreletii had been translocated it tended to out-compete and displace C. acutus (Lavín et al. 2014). Several cases of hybridization between the two species have been documented from the Mexican Yucatan Peninsula (Cedeño-Vázquez et al. 2008; Rodriguez et al. 2008), Oaxaca (Serrano-Gómez et al. 2016), and Belize (Ray et al. 2004), which is particularly critical for C. acutus (Serrano-Gómez et al. 2016), since the genome of C. moreletii seems to have a higher fitness value (Lavín et al. 2014). This apparently is true on the Yucatan Peninsula, given the rarity of C. acutus in areas where C. moreletii is much more abundant. Actions to prevent genomic pollution of American Crocodiles should be encouraged (Cedeño-Vázquez et al. 2008). Furthermore, undocumented crocodiles could become vectors for infectious diseases, such the West Nile Virus (González-Sánchez et al. 2017).

Lazcano-Barrero (1993) acknowledged several intentional releases of C. moreletii on Isla Contoy from 1981 to 1991. The individuals came from zoos and from seizures at regional fairs. Very likely, however, those crocodiles emigrated or failed to establish there, since Morelet’s crocodiles are not listed as part of the Contoy herpetofauna (Arriaga y Ramírez-Bautista 2008; González-Sánchez et al. 2017). Also, it is unknown if those released crocodiles contributed to genetic pollution of C. acutus populations on Contoy, since studies on hybridization between C. moreletii and C. acutus on the Mexican Yucatan Peninsula did not include samples from that island (Cedeño-Vázquez et al. 2008; Rodriguez et al. 2008; Machkour-M’Rabet et al. 2009) (Fig. 13).

Reptiles – Testudines – Turtles

Family Emydidae

Trachemys scripta (Thunberg in Schoepff, 1792)

Fig. 14

A common misbelief in Mexico is that the Pond Slider is native to Japan, thus the common name “Tortugas japonesas” (Japanese turtles) is used frequently. The species’ native geographic range, as presently understood, includes the southeastern United States and adjacent lowlands of northeastern Mexico (Rhodin et al. 2017), but due to their intensive husbandry and commercialization as pets, this turtle has become the most widespread chelonian in the world (Standfuss et al. 2016). It is almost impossible to determine exact invasion routes or dates of introduction, since these turtles have been subjected to extensive illegal trafficking, and can easily be purchased in pet stores and markets throughout Mexico (Yáñez-Arenas et al. 2016). Still, it seems highly probable that most introductions in Mexico occurred during the 1980’s through 1990’s, when “Japanese Turtles” gained immense popularity among pet owners.

Any review of literature covering the distribution of this introduced turtle should be made with special care, since the name Trachemys scripta, until recently, included almost every Pond Turtle population ranging throughout Middle America, unfortunately, listed as subspecies of T. scripta (Campbell 1998; Köhler 2008). Perhaps this is the reason why T. scripta appears in the Reptile Database as being introduced into all countries of Central America (Uetz et al. 2020). Johnson et al. (2015b) gave an account of the taxonomic and nomenclatural history of many taxa being considered up until that time, but some of that information has changed. Many subspecies of Pond Turtles recently have been rightly elevated to full species, thereby making them native populations to their inclusive ranges throughout Middle America (e.g., Seidel 2002; Fritz et al. 2012; McCranie et al. 2013; Parham et al. 2013, 2015). According to Parham et al. (2015), the native species in Middle America that occurs geographically closest to T. scripta is T. venusta in northeastern Mexico, which in itself has gone through taxonomic reorganization, so today its range is primarily restricted to the Atlantic versant of Middle America into northern South America.

Our review of introduced species will only cover what has been called the Red-eared Slider, T. scripta elegans (Wied, 1838), which is listed among the 100 most dangerous invasive species, according to the Invasive Species Specialist Group (Lowe et al. 2000), although we herein do not consider subspecies as a legitimate formal taxonomic category (see Johnson et al. 2015b), but only “pattern classes” of geographic variants exhibited within a single species (Grismer 2002). The Elegans pattern class individuals of T. scripta are of special concern due to their deleterious tendencies to outcompete other turtles for basking sites, and as possible vectors for spreading diseases (Lavín et al. 2014).

Established introduced populations of the Elegans pattern class of T. scripta are disconnectedly distributed in several parts of Mexico, such as in Baja California, within several northern states, along the southern Pacific slopes, and on the Yucatan Peninsula (Lavín et al. 2014). We must clarify that the latest revision of the herpetofauna of the Mexican Yucatan Peninsula (González-Sánchez et al. 2017) failed to mention T. scripta, since it was thought that distributional records at that time from the peninsula corresponded to its close relative, T. venusta. These authors did overlook the report published by Böehm (2013), however, for some turtles from populations living within cenotes on the Yucatan Peninsula that clearly resembled the Elegans pattern class of T. scripta.

In Honduras, T. scripta has been observed in Río Llanitos, Santa Bárbara, and Isla Guanaja in the Islas de la Bahía (= Bay Islands) (McCranie et al. 2005; Solís et al. 2014). McCranie and Valdés-Orellana (2014), however, did not know whether the few known specimens from Guanaja were part of an established population, or merely individual escaped pets. McCranie and Valdés-Orellana (2014) also specifically mentioned a female from Isla de Guanaja at Savannah Bight (FMNH 283584) that was suspected to be an escaped pet because both T. scripta and T. ornata (= T. venusta; Parham et al. 2013, 2015) are regularly kept as pets by local citizens. Another report was published by McCranie and Valdés-Orellana (2014) of a vouchered, but supposedly uncatalogued T. scripta in the collection of UNAH from 11.9 km north of Cofradía, Cortés, on the mainland, located 290 km southwest of Savanna Bight on Isla Guanaja.

Kraus (2009) listed the red-eared slider (Elegans pattern class of T. scripta) as introduced in Panama, citing Moll (1995), but that reference was an editorial letter in which the author only mentioned having collected specimens in Panama; thus, he didn’t provide any specific locality, date, or voucher specimen. Jaramillo et al. (2010) did not list T. scripta as an introduced species in Panama. Therefore, we have found no verified report for any established populations in Panama (Figure 14).

Figure 14. 

Trachemys scripta elegans (Elegans pattern class of T. scripta). Xcunya, Yucatan, Mexico. Photograph by Luis Díaz-Gamboa.

Trachemys venusta (Gray, 1855)

The Mesoamerican Slider, as discussed elsewhere, was involved in the taxonomic dispute associated with T. ornata (Fritz et al. 2012; Parham et al. 2015). Presently, T. ornata and T. venusta are considered as separate species, with T. ornata ranging along the Pacific lowlands of Mexico from Sinaloa to at least southeastern Guerrero, and T. venusta ranging from Tamaulipas, Mexico, on the Atlantic lowlands into South America (Parham et al. 2015; Legler and Vogt 2013). Much of the taxonomic controversy involved slider turtles sampled from the vicinity of Acapulco, Guerrero, which Parham et al. (2015) showed to be translocated T. venusta. We agree with their conclusion.

Family Kinosternidae

Kinosternon integrum (Le Conte, 1854)

A single individual of the Mexican Mud Turtle was captured 29 March 2007 in a perennial pool at the bottom of a rocky canyon in the Sierra La Laguna, Baja California Sur (Luja et al. 2007). Apparently, this was one of a pair mentioned by a local settler as being released into the pool by someone else, most likely in the second half of the 1980’s; the other turtle was found dead at the site approximately ten years after its original release. Luja et al. (2007) made no mention about where the turtles originated. We assume the turtles were translocated by someone from the mainland on the Pacific versant of northwestern Mexico. Ferryboats regularly carry people across the Sea of Cortes from the Port of Pichilinque, near La Paz, Baja California Sur, to ports in Sinaloa at Topolobampo (near Los Mochis) in the northwest and Mazatlán in southwestern portion of the state. For now, we assume an established translocated population is probable at this site.

Iverson et al. (1998) stated that K. integrum populations in the Valley of Mexico were introduced, but they did not provide arguments to support that idea. It was also suggested that the probable extinction of K. hirtipes hirtipes, a supposed endemic subspecies of mud turtle known only from three to five localities in the Valley of Mexico (Legler and Vogt 2013; Rhodin et al. 2017), “may or may not be associated with the introduction of K. integrum, which has become very common there” (van Dijk et al. 2007); no evidence or explanation was given to confirm that assessment. Ramírez-Bautista et al. (2009) indicated that K. integrum and K. hirtipes are sympatric in Lago de Xochimilco (Distrito Federal), but they did not indicate if K. integrum was introduced or that K. hirtipes had become extinct there. Kinosternon integrum also inhabits Canal de Chalco, in the state of Mexico, and K. hirtipes also occurs in Lago de Tlahuac. We assume for now that both species have established populations in the Valley of Mexico, and that K. integrum was introduced there.

Family Trionychidae

Apalone spinifera (Le Sueur, 1827)

The Spiny Softshell has a limited native range in Mexico, principally in drainage systems associated with the Rio Grande, which is the border with Texas and a very small segment of New Mexico near El Paso, with neighboring states in Mexico (Chihuahua, Coahuila, Nuevo León, and Tamaulipas), then continuing along the Gulf lowlands to approximately Soto la Marina, Tamaulipas (Legler and Vogt 2013). Farr (2015) apparently considered this turtle native throughout Tamaulipas, as did Lemos-Espinal and Dixon (2013) for the record of A. spinifera in San Luis Potosí. This turtle has been reported four times from translocated populations in northwestern Mexico (Rorabaugh and Lemos-Espinal 2016), once in the lower basin of Río Colorado and its drainages in the Mexicali Valley, Baja California (Grismer 2002), and three times in Sonora, twice from the Welton Canal area in the southeastern Río Colorado Valley (Rorabaugh et al. 2008) and once from the Río San Rafael in the Municipality of Cananea, 449 km to the east-southeast (Rorabaugh and King 2013). The introductions in Sonora are probably the result of expansion of an invasion front that might have originated by intentional introductions of Spiny Softshell turtles (together with fishes and frogs) by ranchers along the Gila River early in the 20th century. From there, the range expanded until reaching the lower Colorado Basin and Mexicali Valley (Miller 1946), where reportedly they were once plentiful, but now in decline due to hunting pressures (Mellink and Ferreira-Bartrina 2000); Spiny Softshell meat has been served frequently in Chinese restaurants in Mexicali (Grismer 2002) (Table 3, Map 26).

In the Cuatro Ciénegas area of Coahuila, Mexico, A. spinifera arrived at some local water sources when irrigation channels were opened from the northeast in the 1880’s (McGaugh and Janzen 2008). It has been linked to the impending extinction of the endemic Black Softshell (A. atra) due to hybridization, although this claim is yet to be fully confirmed, since “pure” individuals of A. atra could still be found during the 1970’s and 1980’s (Webb 1973; Legler and Vogt 2013), especially at the type locality. McGaugh and Janzen (2008) thought that there was insufficient molecular evidence to differentiate the two turtles and concluded they were conspecific, but Bonin et al. (2006) and Legler and Vogt (2013) regarded them as separate species. Wilson and Johnson (2010) reviewed the evidence and decided to continue recognizing A. atra until a study showing complete genetic introgression between all populations of the two species refutes the claim that they are separate evolutionary lineages. We agree with that conclusion because of historical genetic isolation and the reasons enunciated by Wilson and Johnson (2010). If it turns out to be true that a translocated invasive population of A. spinifera has hybridized to the point of full genetic introgression with A. atra, then, unfortunately, this situation will become a prime example of genetic extinction of a formerly endemic species by hybridization with a non-endemic species.

Outside of northern Mexico, a single record exists for A. spinifera from Jalisco in an artificial pond in Puerto Vallarta (Cupul-Magaña 2012), but nothing was said about it being from an established population. According to F. G. Cupul-Magaña (pers. comm.), the turtle was still alive as of August 2018 and living in an open-air aquarium in Puerto Vallarta, but he does not know the original date of translocation or where it originated. Cruz-Sáenz et al. (2017) did not list A. spinifera as a member of the Jalisco herpetofauna, presumably because they did not consider it an established population; we agree with that conclusion.

There are, however, documented localities in Guerrero for A. spinifera from along the Río Balsas drainage, one from the vicinity of Colonia Valerio Trujano, near the Mezcala Bridge, Municipality of Edwardo Neri, that seems to be from an established population. Local residents indicated that it had been present there since the 1950’s (Lemos-Espinal et al. 1999). Two turtles were taken from that area and according to the authors, were deposited in the Herpetology Collection, Unidad de Biología, Tecnología y Prototipos, UNAM, Campus Iztacala, Tlalnepantla, Estado de México. The other locality that allegedly has an established population is from the Municipality of Copalillo, 5 km northeast of Papalutla at the edge of Río Atoyac; a specimen from there was deposited in the Colección del Laboratorio de Herpetología Vivario, Facultad de Estudios Superiores Iztacala, UNAM (CLHV 4462-E) (Jiménez-Arcos et al. 2009).

Castro-Franco and Zagal (2004) reported an adult female A. spinifera captured 17 October 1999 in the Río Amacuzac on the Sierra de Huautla Natural Protected Area near Las Huertas Spa, Municipality of Tlaquiltenango, Morelos, which was deposited in the Herpetological Collection of the Universidad Autónoma del Estado de Morelos (EBUM 2898). Another turtle was observed previously in that same Protected Area in Cruz Pintado Pond, but was not captured. Those turtles indicate a probable established population, and were thought to have been released pets.

García-Vázquez et al. (2009) first reported A. spinifera as occurring in Puebla, Mexico, without commenting about it being introduced to the state or not. Most recently, Woolrich et al. (2017) considered the Puebla record as an introduction, a determination with which we agree.

Reptiles – Squamata – Lizards

Family Iguanidae

Ctenosaura conspicuosa (Dickerson, 1919)

The Isla San Esteban Spiny-tailed Iguana only exists on Cholludo and San Esteban islands, in the Sea of Cortes, located in close proximity to the coast of Sonora, Mexico. The cultural evidence suggests that C. conspicuosa populations on both islands could be due to a prehistoric introduction of C. nolascensis from Isla San Pedro Nolasco by the Seri culture (Grismer 2002; Nabhan 2002). The divergence time between those two lineages, however, is much older (~890, 000 years), which coincides with the detachment of Isla San Esteban from mainland Sonora (Edwards et al. 2005). Even so, there seems to be enough molecular and cultural evidence to indicate that the population of C. conspicuosa present on Isla Cholludo was introduced indeed from Isla San Esteban by the Seri culture (Buckley et al. 2016) (Table 3, Map 27).

Ctenosaura pectinata (Wiegmann, 1834)

The Western Spiny-tailed Iguana naturally occurs in low to intermediate elevations primarily on the Pacific versant of Mexico from Sinaloa into Chiapas, including subhumid interior basins and valleys and offshore islands (Uetz et al. 2020). This iguana was introduced on the remote Isla Clarion sometime in the mid-1990´s (Aguirre-Léon and Matías-Ferrer 2017; CONANP 2018), although the path of introduction is unknown. We can be sure that they were introduced, since spiny-tailed iguanas are conspicuous animals, and not shy around human presence; thus, they should be noted easily, but were not reported in previous herpetofaunistic listings for that island (Townsend 1890; Brattstrom 1955, 1990). Whether there is an impact of C. pectinata on native species, such as the endemic Urosaurus clarionensis is unknown (Table 3, Map 28).

Ctenosaura similis (Gray, 1831)

Fig. 15

The Black Iguana naturally occurs on the Atlantic and Pacific versants from Mexico, below the Isthmus of Tehuantepec, through all countries in Central America (McCranie et al. 2005; Köhler 2008; Buckley et al. 2016). This lizard was reported as introduced to Isla Roatán, Honduras, where it was detected in 2012 (Pasachnik 2013); apparently, the site of initial translocation was on a small island off the south coast of Roatán near Coxen Hole. The Black Iguana could be an immediate threat to native C. oedirhina, since it might compete for resources and/or hybridize (McCranie and Valdés-Orellana 2014). Probably other insular populations of C. similis in the Caribbean waters of Mexico and Central America are intentional introductions, as exemplified by Sunyer et al. (2013), who mentioned probable translocations of C. similis and Iguana iguana as food sources for residents of the Corn Islands, Nicaragua, although this opinion was not based on empirical fact. A Mexican Navy cadet told one of us (VHGS) that both the Black Iguana and Green Iguana were introduced several decades ago as ornamental species by navy cadets in Banco Chinchorro, Quintana Roo, Mexico, but there is no way to verify that claim. Thus, except for the Black Iguana population on Roatán, we do not consider the other locations as having confirmed translocations (Table 3, Map 29).

Figure 15. 

Ctenosaura similis. A juvenile specimen from Little Corn Island, Nicaragua. Photograph by Javier Sunyer.

Sauromalus hispidus (Stejneger, 1891)

The Spiny Chuckwalla inhabits several islands within the Gulf of California, including: Angel de La Guarda, Alcatraz, Cabeza de Caballo, Flecha, Granito, Mejía, Piojo, Pond, San Lorenzo Norte, San Lorenzo Sur, and numerous islands in Bahía de Los Ángeles (Grismer 2002; Buckley et al. 2016); a single individual also was reported from Isla Rasa (Velarde et al. 2008). The Spiny Chuckwalla was probably introduced from Ángel de la Guarda onto La Ventana and Smith islands, since individuals from both populations have “a nearly identical cytochrome b sequence as some Ángel de la Guarda S. hispidus.” The species also could have been translocated to San Lorenzo Sur, but the possibility exists that there was a previous land bridge involved (Petren and Case 1997). The Spiny Chuckwalla population on Isla Alcatraz might encompass introduced hybrids (Grismer 2002), since hybridization and genetic introgression have been suggested among S. hispidus, S. varius, and S. obesus (Robinson 1972). Kraus (2009) indicated a date of introduction to be around 1920, but he didn’t give concrete reasons for that estimation, and overlooked the much earlier cultural evidence of insular translocations by Seri navigators (Nabhan 2000, 2002). Shaw (1946) pointed out that some introductions probably occurred naturally due to Ospreys (Pandion haliaetus) inadvertently dropping live chuckwallas into their nests; he also was aware of translocations of individuals by the Seri culture as food sources (Table 3, Map 30).

Sauromalus varius (Dickerson, 1919)

The Piebald Chuckwalla is only known from the islands of San Esteban and Roca Lobos in the Sea of Cortes (Grismer 2002). Hollingsworth et al. (1997) reported the first record for Roca Lobos, which they considered an introduced population based on the lack of morphological divergence between that population and the one on San Esteban. They also thought that natural colonization seemed unlikely since Isla Salsipuedes, with no S. varius, lies between the two islands and would have represented a barrier to direct overwater dispersal. Hollingsworth et al. (1997) also pointed out that the S. varius population on Roca Lobos could have been an intentional introduction by researchers attempting an experiment on the effects of colonization and evolution of insular populations. Considering the decades of studies they quoted, the introduction might have occurred in the midpoint or end of the 1970’s. A decade after the initial report, the Isla San Esteban Chuckwalla population was reported as being “healthy and reproducing” (Lovich and Mahrdt 2007) (Table 3, Map 31).

Family Phrynosomatidae

Uta stansburiana (Baird & Girard, 1852)

The Side-blotched Lizard is a common widespread generalist, occurring in the western United States, northern Mexico, and along the Baja California Peninsula and many of its associated islands (Grismer 2002). Interestingly, U. stansburiana is present on islands in the Sea of Cortes and Pacific Ocean, and on those with both continental and oceanic origins (Murphy and Aguirre-León 2002), some of which also are associated with endemic species of Uta (Grismer 2002). Certainly, many of those populations are natural overwater dispersal colonizers and, taking into account the anthropological evidence of translocations, some insular populations, assuredly, could be introduced (Murphy and Aguirre-León 2002). This assertion seems to be supported at least by the Uta populations on Isla La Raza, a tiny volcanic island in the Sea of Cortes, where molecular evidence suggests colonization from the north, from Ángel de la Guarda and/or Isla Mejía populations, something that Murphy and Aguirre-León (2002) considered unlikely by natural over water current dispersal, since the severe upwellings in that area should impede that movement. Additionally, the anthropological evidence of human occupancy on Isla La Raza and the lack of genetic differentiation points to accidental introduction, probably during prehistoric times (Upton and Murphy 1997) (Table 3, Map 32).

Reptiles – Squamata – Snakes

Family Boidae

Boa imperator (Daudin, 1803)

The Central American Boa Constrictor, formerly a subspecies of Boa constrictor (Reynolds et al. 2014), is widespread in Middle America, ranging on the Atlantic versant of Mexico to the Isthmus of Tehuantepec and on both Atlantic and Pacific sides below the Isthmus to northwestern Colombia (Card et al. 2016; Suárez-Atilano et al. 2017). Despite being native on the mainland Caribbean versant of Mexico, including the Yucatan Peninsula and several offshore islands, it was considered an introduced invasive species on Cozumel Island, Quintana Roo (Martínez-Morales and Cuarón 1999; González-Sánchez et al. 2017), although reasons for its distributional status were controversial as late as 2008 (Álvarez-Romero et al. 2008). It is clear now that B. imperator was unknown on Cozumel until 1971, when according to local independent informants, cinematographers filming the movie “El Jardín de la Tía Isabel” released several boas of various sizes in order to create a more “exotic” atmosphere (González-Sánchez et al. 2017). This conclusion was reinforced by the fact boas had not appeared in any biological listing until the inventory made by López-González (1991). A molecular study by Vázquez-Domínguez et al. (2012) documented the existence of a founder effect in the genome of Cozumel’s B. imperator and close phylogenetic ties with populations on the adjacent mainland.

Martínez-Morales and Cuarón (1999) suggested a possible link between the arrival of B. imperator on Cozumel with the decline of several native species, such as the Cozumel Thrasher (Toxostoma guttatum), Central American Agouti (Dasyprocta punctata), Cozumel Raccoon (Procyon pygmaeus), Cozumel Coati (Nasua narica), and the Cozumel Curassow (Crax rubra), among others. Of those, the Cozumel Raccoon is under a highly critical extinction risk (de Villa-Meza et al. 2011). Thus, it is imperative that a plan should begin now to eliminate or tightly manage this invasive snake species that is threatening Cozumel’s wildlife. We are unaware of any attempted B. imperator control program on Cozumel Island, however, beyond surveys or ecological studies.

Charruau et al. (2015) suggested that B. imperator might be alien to Cayo Centro (Banco Chinchorro, Quintana Roo), since it is a large reptile and not seen previously by local anglers, although the management program for that reserve already listed this snake as occurring there. A molecular analysis, however, should be done in order to clarify the origin of the Cayo Centro population (Table 3, Map 33).

Reported introduced species not on our list of established populations in Middle America

Taricha torosa (Rathke, 1833)

Murphy and Méndez de la Cruz (2010) listed the California Newt (Salamandridae) as introduced into Baja California, but they did not provide any details or explanations. Grismer (2002) considered it a species that probably occurs in northern Baja California. Kuchta (2005) listed it based on supposed records for northwestern Baja California by Slevin (1928), Smith and Taylor (1948), and others, but all of those records need verification. Taricha torosa definitely occurs naturally in San Diego County, California, a few kilometers from the Mexican border (Kuchta 2005), therefore, the most probable scenario of potential extant Mexican populations reflects natural range expansions from the San Diego populations. Thus, at this time, we do not consider it a native or an introduced species in Mexico.

Gopherus agassizii (Cooper, 1861)

The taxonomic history of the G. agassizii species complex of Desert Tortoises (Testudinidae) generally had been unresolved (Murphy et al. 2011). Until recently, only G. agassizii was officially recognized and naturally resided in the Mojave and Sonoran Deserts in the USA and Mexico, southward in Sonora (including Tiburón Island) into areas of Sonoran desertscrub, Sinaloan thornscrub, and tropical deciduous forest to around Alamos, Sinaloa. Reported populations in Baja California and Baja California Sur were mostly associated with introductions (Bury et al. 2002; Murphy et al. 2011; Legler and Vogt 2013). The species complex has undergone recent taxonomic revisions (Murphy et al. 2011; Edwards et al. 2016) and future studies might well lead to additional taxonomic modifications.

Murphy et al. (2011) removed G. agassizii (Agassiz’s Desert Tortoise) from the herpetofauna of Mexico by formally describing G. morafkai (Morafka’s Desert Tortoise). Gopherus morafkai also ranges into the USA, primarily in Sonoran Desert areas in Arizona south and east of the Colorado River. Gopherus agassizii is now considered native to the Mojave Desert of California, Nevada, Utah, small areas in northwestern and southwestern Arizona, and a small section of the Sonoran Desert in southeastern California. Edwards et al. (2016) further divided the Mexican populations into G. morafkai and a new species, G. evgoodei (Goode’s Desert Tortoise), which is endemic to Mexico and native to primarily Sinaloan thornscrub and tropical deciduous forests in east-central and southern Sonora and northern Sinaloa. The two species occasionally hybridize in the ecotone between Sonoran desertscrub (G. morafkai preferred habitat) and Sinaloan thornscrub (G. evgoodei preferred habitat); no hybrids were observed from tropical deciduous forest habitats.

Ottley and Velazquez-Solís (1989) described a new species of tortoise (as Xerobates lepidocephalus), based on one live individual and shell remains of another, from the Cape region of Baja California Sur, specifically from the Sierra San Vicente, 1.5 km north of the Buena Mujer Dam, 20 km south of La Paz. Crumly and Grismer (1994) noted that its morphological variation fell within that expected for individuals from Sonoran populations, thus they considered X. lepidocephalus as a junior synonym of G. agassizii; they further opined that the population was a probable introduction. Grismer (2002), however, considered the possibility of a native relict population existing in the Cape region that later was reduced considerably when goat grazing was allowed. To us, it also seems plausible that the construction of Buena Mujer Dam in the 1980’s would have destroyed a large area of suitable tortoise habitat, leaving only a few survivors. Murphy et al. (2011) inconclusively pointed out that the holotype of X. lepidocephalus might correspond to a hybrid, so if this is the case, it supports the hypothesis of an introduced population, since hybrid lineages are common in non-native species (Edwards et al. 2010). The hybrids most likely would be between G. morafkai and G. evgoodei that originated as pets taken to the La Paz area. On the other hand, if a population of native tortoises exists in the La Paz region, then they certainly should be considered a separate evolutionary species (Murphy et al. 2011), because of its allopatric distribution compared to that of G. agassizii, G. morafkai, and G. evgoodei. In that case, measures to guarantee the population’s protection should be made available as soon as possible (Murphy and Mendez de la Cruz 2010).

At this point, we do not think that the population around La Paz has been identified definitively as being G. morafkai, G. evgoodei, G. agassizii, or one of hybrid origin. We also consider any inhabitants that will be found in northeastern Baja California will probably fall within the native range of G. morafkai, unless they can be shown positively to be part of translocated G. agassizii, G. evgoodei, or hybrid populations. Thus, it will not be appropriate at this time to list any population of the G. agassizii species group of Desert Tortoises as being introductions within northwestern Mexico.

Gopherus berlandieri (Agassiz, 1857)

A single specimen of the Texas Tortoise (Testudinidae) was found in a city park at Puerto Vallarta, Jalisco, Mexico (Cupul-Magaña 2012). Cruz-Sáenz et al. (2017) did not consider it as part of the Jalisco herpetofauna, nor is there any other record for this species outside its native range in northeastern Mexico, so we concur that it was not part of an established population.

Staurotypus triporcatus (Wiegmann, 1828)

Until recently, the Mexican Giant Musk Turtle (Staurotypidae) was considered native to the Atlantic lowlands from central Veracruz, Mexico, through the southern Yucatan Peninsula, and on to the western Caribbean lowlands of Honduras (Legler and Vogt 2013; Rhodin et al. 2017). Terán-Juárez et al. (2015) reported this turtle near Ocampo, Tamaulipas, Mexico, ca. 524 km to the north of the closest known locality in central Veracruz. They considered this record the result of an introduction due to the large hiatus between those localities. Terán-Juárez et al. (2016), however, later regarded the population as native because there was no empirical evidence to support its translocation by human activities from farther down the Gulf Coastal Plain. Our experience with S. triporcatus indicates that individuals rarely leave water sources and cross roads like other kinosternid turtles, especially those in the genus Kinosternon and even Claudius. Legler and Vogt (2013) also mentioned that S. triporcatus in Belize never were observed on land during its activity season. Crossing roads could be a good source for translocating turtles along roadways, but if that rarely happens with S. triporcatus, the capability of being translocated is diminished. Until additional information indicates otherwise, we agree with Terán-Juárez et al. (2015) that the records came from a marginal area of its native range in northeastern Mexico.

Trachemys ornata (Gray, 1831)

The Trachemys scripta species group (Emydidae) has had a confusing taxonomic history in Middle America, especially those populations occurring in tropical latitudes (Johnson et al. 2015a), so more work is required to properly determine native species boundaries (Parham et al. 2013, 2015). The Ornate Slider is presently considered a Mexican endemic ranging on the Pacific lowlands of Mexico below 300 m elevation from Sinaloa to 4 km northwest of Ixtapa, Guerrero, which is ca. 220 km northwest of the Acapulco, Guerrero area (Mertz et al. 2015). This turtle also was reported to occur in several lagoons around Acapulco and sold for food in local markets (Legler and Vogt 2013). According to Parham et al. (2015), however, all samples they evaluated near Acapulco were T. venusta, which they considered translocations from the Atlantic lowlands of Mexico. Rhodin et al. (2017) alleged that an isolated record of T. scripta (= T. ornata?) in Michoacán and other remote records on the Pacific lowlands in Guerrero might have been introduced as well. A connection between the coastal lagoon systems on the Pacific lowlands, however, might have existed in the past (Legler and Vogt 2013), and possibly still does, so it seems likely to us that the range of T. ornata might extend even farther down the Pacific lowlands past the Acapulco region. We will not include T. ornata as anything other than native until the origins and taxonomic status of those Pacific lowland populations are fully resolved.

Cnemidophorus ruatanus (Barbour, 1928)

The Ruatan Whiptail (Teiidae) was reported (as Cnemidophorus lemniscatus) by Stafford and Meyer (2000) from Monkey River town, Toledo District, Belize; Stafford et al. (2010), without comment, considered it as an introduced species. McCranie and Hedges (2013) elevated C. ruatanus to species level from its previous status as a subspecies of C. lemniscatus. Due to the proximity of this record to other known localities, however, and for maintaining consistency in a coastal scheme of distribution (McCranie and Hedges 2013), we consider this population as being a marginal natural expansion of its total distributional range.

Gonatodes albogularis (Duméril & Bibron, 1836)

Fig. 16

Identifying the native distribution of the Yellow-headed Gecko (Sphaerodactylidae) is problematical because of its wide range in parts of Middle America, northern South America, and on many islands in the West Indies (Uetz et al. 2020). A pertinent question is whether its native range is restricted to Middle and South America and it is introduced in the West Indies, or vice versa. Stuart (1963) was under the impression that all West Indian records were introductions. Villa et al. (1988) and Köhler (2008) did not mention it ranging into the West Indies, which we infer to mean that those individuals were not considered native to that area. Others have apparently included the West Indies as part of the natural range (e.g., Lee 2000; Savage 2002; McCranie et al. 2006; Johnson et al. 2010); most of those sources also reported G. albogularis as being introduced into Florida, without referring to its possible origin.

Figure 16. 

Gonatodes albodularis. Ometepe island. Nicaragua. Photograph by Javier Sunyer.

On the mainland, G. albogularis occurs from the Pacific slopes of the Soconusco and Sierra Madre regions in Chiapas, Mexico, through Central America into northern Colombia and Venezuela (Johnson et al. 2015a), at elevations lower than 1,000 m. Its northernmost locality on the Atlantic versant of Mexico recently was reported as Minatitlán, Veracruz, by Guzmán-Guzmán and Palma-Martínez (2016). It was discovered in Belize City in 1996 (Lee 2000) and Stafford et al. (2010) thought it had been introduced recently into that country, but with no substantiated evidence. Until new information demonstrates otherwise, we regard G. albogularis as being native to Middle and South America and exotic in the West Indies and Florida.

Phyllodactylus nocticolus (Dixon, 1964)

The occurrence of Peninsular Leaf-toed Gecko (Phyllodactylidae) on Isla Tiburón could be due to natural over-water island hopping, accidental introduction (Murphy and Aguirre-León 2002), ancient historical translocation from other islands in the Sea of Cortes (Nabhan 2000), or even by paleotectonic activity (Blair et al. 2009). Without specific evidence to support any of the above-mentioned scenarios, we elect not to list this species as introduced at this time, but it is certainly a candidate worth further investigation.

Phyllodactylus xanti (Cope, 1863)

Mellink (2002), citing Nabhan (2002), indicated that the Cabo Peninsular Leaf-toed Gecko (Phyllodactylidae) was introduced involuntarily from mainland Baja California onto San Lorenzo, San Esteban, Tiburón, and Alcatraz islands, probably by boats. We could not find any corroborating reference or study suggesting that scenario. Thus, because we could not specifically determine if the P. xanti population on Tiburón Island originated there by natural dispersal or by some sort of human-mediated introduction process (see P. nocticolus account above), we decided not to include this species on our list of introductions at this time.

Sauromalus obesus (Baird, 1859)

A chance exists that the Western Chuckwalla (Iguanidae) might have been translocated to several islands in the Sea of Cortes, based on the same reasons as those for S. hispidus and S. varius (Nabhan 2002), but not enough evidence was presented to support that contention. Therefore, we are awaiting more verification before deciding to place it on our list of species introduced into Mexico.

Varanus exanthematicus (Bosc, 1792)

The Savanna Monitor (Varanidae) is known in Mexico from a single record in Puerto Vallarta, Jalisco (Cupul-Magaña 2012). This record probably corresponds to an escaped pet or intentional release. We know of no other record from anywhere else in Middle America, so we do not consider the lizard as coming from an established population.

Considerations for management and control measures for introduced species

The main difficulty for management and control of the introduced invasive herpetofauna of Middle America is the lack of documentation. Most conservation managers or agencies have little to no tradition of publishing in the refereed scientific literature, so, for the most part, successes and failures of invasive species are found in grey literature sources (Simberloff 2009). Even more daunting is the fact that among known eradication/management programs cited in the literature, few concern reptiles (Campbell III et al. 1999; Toda et al. 2010) or amphibians (Miller 2006; Greenlees et al. 2018), and hardly any evaluate effectiveness of their control techniques (Rodda et al. 1999; Davis et al. 2015; Haramura et al. 2017; Muller and Schwarzkopf 2017). The main consequence of this “nothing can be done” approach is the lack of evidence for successful programs (Simberloff 2009). Therefore, the first challenge must be to encourage managers to publish their results in the scientific literature.

A key component of any successful control or eradication program is early detection of an invasion and a quick response (Simberloff 2009). Unfortunately, only in limited cases are introductions detected early, so usually dates are merely approximations or suppositions. The true origins of many species are often uncertain, especially if they are common, and times and places of the introduction are unknown; many are commonly considered native even though their true place of origin is unresolved. Those instances refer to the “cryptogenic species” of Carlton (1996), a good example of which is the status of A. allisoni being listed as native to the Yucatan Peninsula of Mexico (González-Sánchez et al. 2017), Belize (Stafford et al. 2010), and Honduras (Solís et al. 2014; McCranie 2015), even though others consider them introductions, including Glor et al. (2005) and Álvarez-Romero et al. (2008). Additionally, in some situations, some apparent “endemic” species might be, in fact, the result of an ancient anthropogenic introduction of an alien species; this situation can occur particularly in insular ecosystems after thousands of years of isolation, and are referred to as “ethnospecies” (Hofman and Rick 2018). This explanation could apply to some endemic iguanids (such as Sauromalus and Ctenosaura) on islands in the Sea of Cortes; there has been speculation about the origin of the population of C. conspicuosa on San Esteban island, which is believed to have originated from translocated individuals of C. nolascensis from San Pedro Nolasco Islands by the Seri people (Grismer 2002). Davy et al. (2011), however, refuted this notion and specified that C. nolascensis and C. conspicuosa diverged thousands of years before human colonization of the Sea of Cortes. Still, the idea should not be discarded, as some iguanid endemic lineages on those islands might be ethnospecies.

Another key component of control and eradication programs involves the correct identification of the presumed introduced species by qualified specialists and their training of non-expert volunteers so to avoid misidentification of native species as introduced ones. These specialists also need to participate in the removal programs. These lessons have come from the work of Rick Shine and his colleagues on the Cane Toad or Marine Toad (Rhinella marina) in Australia, as reported in numerous publications, including the book Cane Toad Wars (Shine 2018). Nonetheless, the work of Shine and his colleagues has demonstrated that simple, physical removal of adult toads is ineffective in reducing their numbers or in curtailing their spread through Australia and that what is more promising is an approach using integrated pest control (Shine 2015). Shine (2015: 312) indicated that “Cane toads are formidable invasion machines, and it is unlikely that any single method will ever eradicate them. Even with a combination of methods, landscape-scale extirpation is vanishingly unlikely. However, the new weapons developed out of recent ecological research on this high-profile invasive anuran provide great encouragement.” Shine (2015: 314) further noted that the release of “juvenile (and thus, non-lethal) toads at the current invasion front [can be employed] to train native predators to avoid toads as prey” and that “funnel-traps baited with toad toxins can eradicate toad tadpoles from natural water bodies.” In general, he concluded that “cane toads in Australia provide a clear example that [one needs] to understand an invasive species if [one wants] to control it.” This “whole-biology” approach also is stressed by Tyler (2006) in his discussion of the potential use of pheromones of both tadpoles and adults in the control of invasive populations of this toad.

A common challenging situation emerges in cases where some translocated species are under legal protection, but are exotic or even invasive in other parts of the country. This situation keeps conservation managers from performing effective eradication or control measures on those invasive populations (Lazcano et al. 2010). For example, Morelet’s Crocodile and Boa imperator (as Boa constrictor) appear under the categories of “subject to special protection” and “threatened,” respectively, by Mexican legislation (D.O.F. 2015). Obviously, when describing an invasive species, its location of controlling action should be strictly identified (Richardson et al. 2011), or an accurate delineation of both the native areas where the species resides and where it is invasive be given (Ochoa-Ochoa et al. 2017).

The first obvious method for controlling introduced species of amphibians and reptiles is direct capture/sacrifice and trapping methodology. Some biologists, however, are opposed to the sacrificing such introduced creatures, for ethical reasons. This position is entirely understandable and is complex enough to require adequate discussion elsewhere. Several capture techniques include using nooses, pitfall traps, funnel traps, sticky traps, rubber bands, firearms, blowguns, and road cruising, among others, whose effectiveness are well known to herpetologists, thus we will not detail them here. Less common, but potentially successful procedures are discussed below.

Chemical control has a long tradition for managing invasive mammals, but its use has been employed infrequently for herpetofaunal control. The most publicized instances involved Brown Tree Snakes (Boiga irregularis) on Guam, where Brooks et al. (1998) tested lethality of dermal and oral drugs. Clark and Savarie (2012) and Clark et al. (2012) used and evaluated bait poisoning techniques. We are unaware of similar techniques being applied on invasive snakes in Middle America, but evaluating drug toxicity and establishing bait-poisoning plots for controlling B. imperator on Cozumel Island should be evaluated seriously for a long-term control or eradication program.

Invasive anuran control programs in Australia included spraying lethal chemicals in water sources (Kelehear et al. 2012). In the USA, Witmer et al. (2015) evaluated toxicity of different compounds on American Bullfrogs (Lithobates catesbeianus). Several species of anurans and other amphibians have evolved species-specific embryogenic suppression pheromones to reduce intraspecific competition, whereby older larvae produce substances that inhibit development of younger conspecifics (Tingley et al. 2017). The feasibility of using these embryogenic inhibitors for controlling invasive anurans in water sources has received recent attention, with some promising results (Clarke et al. 2016).

A key aspect for the success of some invasive anurans are potent chemical defenses that most vertebrate predators cannot tolerate. Nonetheless, several invertebrate groups are immune to those toxins, and many of them, such as dragonfly nymphs, fishing spiders, water beetles, ants, crabs, and crayfish, voraciously consume tadpoles or early stage metamorphs. Thus, introducing native invertebrate predators can be a biological control option when the only species in the water source is an alien anuran (Cabrera-Guzmán et al. 2012; Cabrera-Guzmán et al. 2015).

A major advantage for invasive species when reaching a new area is access to a parasite-free space (Torchin et al. 2003). With this notion in mind, it makes sense to introduce the relevant parasite into the area in order to diminish the fitness of the invasive population (Torchin et al. 2003; Finnerty et al. 2018). Even if the invasive species does not develop a parasitic disease, accumulation of sub-lethal effects can have significant consequences on overall performance, growth rates, and reproduction potential of infected toads, as reported by Finnerty et al. (2018). Another example of biological control used in Hawaii on the invasive Eleutherodactylus coqui was infecting them with the lungworm Rhabdias elegans (Marr et al. 2010).

A successful molecular tool for early detection and monitoring aquatic and semiaquatic invader species is testing water sources for their waterborne environmental DNA (eDNA), as described by Jerde et al. (2011) and Bohmann et al. (2014). So far, however, this technique has been used mainly for detecting rare or elusive species, although it was used to detect invasive amphibians (Dejean et al. 2012) and reptiles (Piaggio et al. 2014; Hunter et al. 2015). The advantage of eDNA is appealing because it can improve chances for detecting hard to observe species or to identify species displaying crypsis (Jerde et al. 2011; Piaggio et al. 2014). This method also allows conservation monitors to identify key amphibian breeding sites, and could be a valuable tool for locating strategic invasion places, such as those near airports, maritime ports, and plant nurseries (Tingley et al. 2017). In addition, it can be a complementary tool for post-eradication confirmation surveys.

A key feature for managing introduced species, especially in large areas, is to identify important sites to focus control and eradication efforts, as well as to prevent invasions and/or reinvasions before they occur. Environmental niche modelling (ENM) has been a helpful tool for identifying potential corridors among the sources and areas vulnerable to invasions (Peterson 2003; van Wilgen et al. 2009). In fact, the use of ENM for managing and controlling introduced herpetofaunal species recently has been increasing, with several examples originating in Middle America (Rödder et al. 2008; van Wilgen et al. 2009; Lira-Noriega and Ramírez 2016; Yáñez-Arenas et al. 2016).

Decision makers often require methods that help them justify and decide where, when, and on which species to target conservation and/or control programs. For these reasons, there exist protocols that can be used to determine if a species is potentially at risk and deserves attention. Generally, these protocols consist of a questionnaire that must be answered by a specialist or by a panel of experts; examples are revealed in “Método de Evaluación Rápida de Invasividad (MERI) para Especies Exóticas en México” (González-Martínez et al. 2017), and in the Harmonia+ and Pandora+ protocols for invasive species and invasive pathogens, respectively (D’hondt et al. 2015).

Finally, and most important of all, is the human component, which is pivotal for the success of any management program governing actions associated with invasive species (McNeely 2001; García-Llorente et al. 2008). The most obvious reason is that humans are often the determining factor when they transport species into new areas voluntarily or involuntarily, thereby enabling individual animals to cross biogeographical barriers (Vitousek et al. 1997; Mack et al. 2000). Personal attitudes of people relating to introduced species also strongly influence management preferences by giving more support to non-intervention approaches when they think animals have equal rights of existence versus control program when they consider human intervention acceptable for maintaining ecosystem integrity (Sharp et al. 2011). In addition, how urgent or harmful people perceive the risk of invasive species to be can directly reflect a potential willingness to help provide funding or other support for developing strategic management or control programs (García-Llorente et al. 2011). Therefore, any management policy should include serious sociological decisions, together with effective proclamation and educational campaigns incorporating biosecurity as a real value for local citizens.

Conclusions

As noted previously, interest in introduced invasive species is taxonomically skewed toward other vertebrates, such as mammals, birds, or fishes. Thus, with a few exceptions, the ecological influences and damages caused by introduced invasive amphibians and reptiles are unknown. This lack of knowledge can have ominous consequences, such as taxonomic uncertainty, causing voids in legislation, and omissions of reptiles and amphibians in many biosecurity protocols or practices. Additionally, a great proportion of the literature on monitoring or control programs corresponds to technical reports not easily accessible to other researchers and/or managers. A first barrier to overcome is to encourage managers and researchers to identify results of monitoring and control programs on the invasive herpetofauna and to have that information published in accredited journals.

For conservationists to influence protocols, it is imperative that they promote stricter legislation on damaging practices, such as the pet trade, which should be discouraged, especially in those species with high potential of being harmful to Middle American ecosystems, even if they are already present, like Trachemys spp., or those that are common in the pet trade of a region, but not yet reported in the wild, such as Varanus exanthematicus and Python molurus. Stricter legislation should not be limited to vertebrate species, but also extended to pathogens associated with amphibians (or reptiles), such Batrachochytrium dendrobatidis, Saprolegnia parasitica, and Ranavirus spp. We are aware that such listing of potential harmful pathogens exists for Mexico (D.O.F. 2016), but it is unknown to us if any equivalent legislation exists in other Middle American countries.

Finally, the ever-growing trade of goods on a global scale, the increasing interest by people for keeping exotic pets, and the human persistence for environmental degradation will continue to favor arrival and settlement of invasive species. Regrettably, the frequency, scope, and intensity of biological invasions are expected to increase during the ensuing decades. Thus, the study and management of introduced amphibians and reptiles in Middle America is a topic that offers a wide spectrum of opportunities for career development associated with young researchers, conservationists, and other professionals dealing in ecological restoration.

We consider biological conservation as a human value, which includes a series of moral codes and behaviors that transcend time and culture and define us as a species. In this way, and like other values, it contributes to building prosperity and free coexistence among societies. It is also obvious that invasive species control and management is controversial, since many people put a high value on any single living organism, independent of its origin. If they accept invasive species to be a legitimate part of our ecological footprint, however, it is clear that we have a moral and ethical responsibility to educate them on the negative impacts invasive species have on the overall well-being of our biosphere. At the same time, it is also our responsibility to help maintain biosecurity and ecological restoration measures as advocates to prevent, mitigate, and remediate damages caused by invasive species until the majority of humanity accepts the fact that being good stewards of our living spaces is the right thing to do. Conservation professionals also must understand what motivates people’s different attitudes towards invasive species (e.g., why they transport them, whether they perceive them as harmful or not, or whether they are willing to accept control methods or not) in order to develop meaningful programs that discourage harmful behaviors and promote more responsible attitudes. Therefore, integration of the human and technical component is fundamental for accepting biosecurity as one more principle guiding societal behavior.

Acknowledgements

We thank Lizbeth E. Lara-Sánchez for her review and comments on the control methods section. We are also thankful to Rubén Alonso Carbajal-Márquez, Luis Díaz-Gamboa, Julio Gutiérrez-Ramírez, Javier Ortiz-Medina, Louis W. Porras, Javier Sunyer, and Anny Peralta for allowing us to use their photographic material to illustrate this paper. We also are grateful to Vicente Mata-Silva for assistance with corrections to one of the distribution maps and to Louis W. Porras for supplying us with information concerning the references.

References

  • Abarca J, Monge AM (2007) Geographic distribution: Costa Rica, San José: Hemidactylus mabouia. Herpetological Review 38(3): 351–352.
  • Aguirre-Léon G, Matías-Ferrer N (2017) Islas Revillagigedo: sus reptiles y el redescubrimiento de una especie. La Jornada Ecológica 209: 16–18.
  • Ahumada-Carrillo IT, Weatherman GN (2018) A confirmed established population of Lepidodactylus lugubris (Duméril & Bibron, 1836) (Squamata: Gekkonidae) in Mexico. Mesoamerican Herpetology 5(1): 179–180.
  • Álvarez-Romero J, Medellín RA, Gómez de Silva H, Oliveras de Ita A (2005) Boa constrictor. Vertebrados Superiores Exóticos en México: Diversidad, Distribución y Efectos Potenciales. Instituto de Ecología, Universidad Nacional Autónoma de México. Bases de datos SNIB-CONABIO. Proyecto U020. México. D.F.
  • Álvarez-Romero JG, Medellín RA, Oliveras de Ita A, Gómez de Silva H, Sánchez O (2008) Animales Exóticos en México: Una Amenaza para la Biodiversidad, México D.F. , Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, Instituto de Ecología, UNAM, Secretaría del Medio Ambiente y Recursos Naturales, 502 pp.
  • Arriaga A, Ramírez-Bautista A (2008) A Checklist of Reptiles from the Parque Nacional Isla Contoy, México. Boletín de la Sociedad Herpetológica Mexicana 16(2): 36–40.
  • Avila-Villegas H, Rodríguez-Olmos LP, Lozano-Román LF (2007) Rana toro (Lithobates catesbeianus): anfibio introducido en Aguascalientes, México. Boletín de la Sociedad Herpetológica Mexicana 15(1): 16–17.
  • Barker BS, Rodríguez-Robles JA (2017) Origins and genetic diversity of introduced populations of the Puerto Rican Red-Eyed Coqui, Eleutherodactylus antillensis, in Saint Croix (U.S. Virgin Islands) and Panamá. Copeia 105(2): 220–228. https://doi.org/10.1643/CG-16-501
  • Barquero MD, Araya MF (2016) First record of the Greenhouse Frog, Eleutherodactylus planirostris (Anura: Eleutherodactylidae), in Costa Rica. Herpetology Notes (9): 145–147.
  • Barrantes-Madrigal J (2017) El Coquí Común (Eleutherodactylus coqui) en Costa Rica: Selección de Hábitat, Etapa de Invasiòn y Aspectos Sociales Antrópicos que Podrían Influir en su Dispernsión y Manejo. M. Sc. Thesis. Universidad Nacional Costa Rica, 81 pp.
  • Barrantes-Madrigal J, Parallada MS, Alvarado G, Chaves-Acosta VJ (2019) Distribution and invasion progress of Eleutherodactylus coqui (Anura: Eleutherodactylidae) introduced in Costa Rica. Phyllomedusa: Journal of Herpetology 18(1): 101–107. https://doi.org/10.11606/issn.2316-9079.v18i1p101-107
  • Batista A, Ponce M, Garcés O, Lassiter E, Miranda M (2019) Silent pirates: Anolis sagrei Duméril & Bibron, 1837 (Squamata, Dactyloidae) taking over Panama City, Panama. Check List 15(3): 455–459. https://doi.org/10.15560/15.3.455
  • Bauer AM, Jackman TR, Greenbaum E, Giri VB, de Silva A (2010) South Asia supports a major endemic radiation of Hemidactylus geckos. Molecular Phylogenetics and Evolution 57(1): 343–352. https://doi.org/10.1016/j.ympev.2010.06.014
  • Beard KH, Price EA, Pitt WC (2009) Biology and Impacts of Pacific Island invasive species. 5. Eleutherodactylus coqui, the Coqui Frog (Anura: Leptodactylidae). Pacific Science 63(3): 297–316. https://doi.org/10.2984/049.063.0301
  • Blair C, Méndez de La Cruz F, Ngo A, Lindell J, Lathrop A (2009) Molecular phylogenetics and taxonomy of leaf-toed geckos (Phyllodactylidae: Phyllodactylus) inhabiting the peninsula of Baja California. Zootaxa 2027(1): 28–42. https://doi.org/10.11646/zootaxa.2027.1.2
  • Böehm S (2013) Records of invasive Trachemys scripta elegans (Wied-Neuwied, 1839), in cenotes of the Yucatan Peninsula, Mexico. Herpetozoa 26: 98–100.
  • Bohmann K, Evans A, Gilbert MTP, Carvalho GR, Creer S, Knapp M, Yu DW, de Bruyn M (2014) Environmental DNA for wildlife biology and biodiversity monitoring. Trends in Ecology & Evolution 29(6): 358–367. https://doi.org/10.1016/j.tree.2014.04.003
  • Bonin F, Devaux B, Dupré A (2006) Turtles of the World. The Johns Hopkins University Press, Baltimore, 416 pp.
  • Brattstrom BH (1955) Notes on the Herpetology of the Revillagigedo Islands, Mexico. The American Midland Naturalist 54(1): 219–229. https://doi.org/10.2307/2422189
  • Brooks JE, Savarie PJ, Johnston JJ (1998) The oral and dermal toxicity of selected chemicals to brown tree snakes (Boiga irregularis). Wildlife Research 25(4): 427–435. https://doi.org/10.1071/WR97035
  • Buckley LJ, de Queiroz K, Grant TD, Hollingsworth BD, Iverson JB, Pasachnik SA, Stephen CL (2016) A checklist of the iguanas of the world (Iguanidae; Iguaninae). Herpetological Conservation and Biology 11(Monograph 6): 4–46.
  • Bury RB, Germano DJ, Van Devender TR, Martin BE (2002) The desert tortoise in Mexico. Distribution, ecology, and conservation. In: Van Devender TR (Ed.) The Sonoran Desert Tortoise. Natural History, Biology and Conservation. University of Arizona Press, Tucson, 86–108. https://doi.org/10.2307/j.ctvfjcx1x.8
  • Cabrera-Guzmán E, Crossland MR, Shine R (2012) Predation on the eggs and larvae of invasive cane toads (Rhinella marina) by native aquatic invertebrates in tropical Australia. Biological Conservation 153: 1–9. https://doi.org/10.1016/j.biocon.2012.04.012
  • Cadotte MW (2006) Darwin to Elton: early ecology and the problem of invasive species. In: Cadotte MW, McMahon SM, Fukami T (Eds) Conceptual Ecology and Invasion Biology: Reciprocal Approaches to Nature (Vol. 1). Springer, New York, 15–33. https://doi.org/10.1007/1-4020-4925-0_2
  • Campbell III E, Rodda GH, Fritts T, Bruggers R (1999) An integrated management plan for the Brown Treesnake (Boiga irregularis) on Pacific islands. In: Rodda GH, Sawai Y, Chiszar D, Tanaka H (Eds) Problem Snake Management: the Habu and the Brown Treesnake. Cornell University Press, Ithaca, New York, 423–435.
  • Campbell JA (1998) Amphibians and Reptiles of Northern Guatemala, the Yucatán, and Belize. University of Oklahoma Press, Norman, 380 pp.
  • Capinha C, Seebens H, Cassey P, García-Díaz P, Lenzner B (2017) Diversity, biogeography and the global flows of alien amphibians and reptiles. Diversity and Distributions 23(11): 1313–1322. https://doi.org/10.1111/ddi.12617
  • Card DC, Schield DR, Adams RH, Corbin AB, Perry BW, Andrew AL, Pasquesi GIM, Smith EN, Jezkova T, Boback SM, Booth W, Castoe TA (2016) Phylogeographic and population genetic analysis reveal multiple species of Boa and independent origins of insular dwarfism. Molecular Phylogenetics and Evolution 102: 104–116. https://doi.org/10.1016/j.ympev.2016.05.034
  • Carranza S, Arnold EN (2006) Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia:Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 38(2): 531–545. https://doi.org/10.1016/j.ympev.2005.07.012
  • Casas-Andreu G, Aguilar X, Cruz RA (2001) La introducción y el cultivo de la rana toro (Rana catesbeiana) ¿Un atentado a la biodiversidad de México? Ciencia Eergo-sum 8(1): 62–67.
  • Castro-Franco R, Bustos-Zagal MG (2004) Additional records and range extensions of reptiles from Morelos, México. Herpetological Review 35(2): 196–197.
  • Cedeño-Vázquez JR, González-Vázquez J, Martínez-Arce A, Canseco-Márquez L (2014) First record of the invasive greenhouse frog (Eleutherodactylus planirostris) in the Mexican Caribbean. Revista Mexicana de Biodiversidad 85(2): 650–653. https://doi.org/10.7550/rmb.43234
  • Cedeño-Vázquez JR, Platt SG, Thorbjarnarson J (2012) Crocodylus moreletii. The IUCN Red List of Threatened Species 2012. [Accessed 6 February 2021]
  • Cedeño-Vázquez JR, Rodriguez D, Calmé S, Ross JP, Densmore LD, Thorbjarnarson JB (2008) Hybridization between Crocodylus acutus and Crocodylus moreletii in the Yucatan Peninsula: I. evidence from mitochondrial DNA and morphology. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology 309(10): 661–673. https://doi.org/10.1002/jez.473
  • Charruau P, Díaz de la Vega Pérez AH, Méndez de la Cruz FR (2015) Reptiles of Banco Chinchorro: updated list, life history data, and conservation. The Southwestern Naturalist 60(4): 299–312. https://doi.org/10.1894/0038-4909-60.4.299
  • Clarke GS, Crossland MR, Shine R (2016) Can we control the invasive cane toad using chemicals that have evolved under intraspecific competition? Ecological Applications 26(2): 463–474. https://doi.org/10.1890/14-2365
  • Clark L, Savarie PJ (2012) Efficacy of aerial broadcast baiting in reducing brown treesnake numbers. Human-Wildlife Interaction 6(2): 212–221.
  • Clark L, Savarie PJ, Shivik JA, Breck SW, Dorr BS (2012) Efficacy, effort, and cost comparisons of trapping and acetaminophen-baiting for control of brown treesnakes on Guam. Human-Wildlife Interactions 6(2): 222–236. https://www.jstor.org/stable/24874096
  • Colston TJ, Barão-Nóbrega JAL, Manders R, Lett A, Wilmott J, Cameron G, Hunter S, Radage A, Littlefair E, Williams RJ, Lopez Cen A, Slater K (2015) Amphibians and reptiles of the Calakmul Biosphere Reserve, México, with new records. Check List 11(5): 1–7. https://doi.org/10.15560/11.5.1759
  • CONANP (2018) Programa de Manejo Parque Nacional Revillagigedo, Mexico. Comisión Nacional de Areas Naturales Protegidas, 328 pp.
  • Conant R, Collins JT (1998) A Field Guide to Reptiles and Amphibians of Eastern and Central North America, Third edition. Houghton Mifflin, Boston, Massachusetts, 450 pp.
  • Contreras-Calvario AI, Mora-Reyes A, Parra-Olea G, Mendoza AM (2018) New records of the introduced Eleutherodactylus planirostris (Anura: Eleutherodactylidae), in the state of Veracruz, México. Herpetological Journal 28(2): 96–99.
  • Crawford AJ, Alonso R, Jaramillo CA (2011) DNA barcoding identifies a third invasive species of Eleutherodactylus (Anura: Eleutherodactylidae) in Panama City, Panama. Zootaxa 2890(1): 65–67. https://doi.org/10.11646/zootaxa.2890.1.6
  • Crumly CR, Grismer LL (1994) Validity of the tortoise Xerobates lepidocephalus Ottley and Velazques in Baja California. In: Bury RB, Germano DJ (Eds) Biology of North American Tortoises. Fish and Wildlife Research 13: 33–37.
  • Cruz-Sáenz D, Muñoz-Nolasco FJ, Mata-Silva V, Johnson JD, García-Padilla E, Wilson LD (2017) The herpetofauna of Jalisco, Mexico: composition, distribution, and conservation status. Mesoamerican Herpetology 4(1): 22–118.
  • Cupul-Magaña FG (2012) Varanus exanthematicus (Bosc, 1792), Apalone spinifera emoryi (Le Sueur, 1827) y Gopherus berlandieri (Agassiz, 1857): reptiles exóticos en el área urbana de Puerto Vallarta, Jalisco, México. Cuadernos de Herpetología 26(1): 59–60. http://hdl.handle.net/10915/18207
  • Cupul-Magaña FB, Rubio-Delgado A (2003) Distribución de dos especies de tortugas dulceacuícolas, Kinosternon chimalhuaca y Chelydra serpentina (Testudines), en Puerto Vallarta, Jalisco, México. Boletín de la Sociedad Herpetológica Mexicana 11: 49–50.
  • Davis JL, Alford RA, Schwarzkopf L (2015) Some lights repel amphibians: implications for improving trap lures for invasive species. International Journal of Pest Management 61(4): 305–311. https://doi.org/10.1080/09670874.2015.1058991
  • Davis MA (2009) Invasion Biology. Oxford University Press on Demand, Oxford, 244 pp.
  • Davy CM, Méndez-de la Cruz FR, Lathrop A, Murphy RW (2011) Seri Indian traditional knowledge and molecular biology agree: no express train for island-hopping spiny-tailed iguanas in the Sea of Cortés. Journal of Biogeography 38(2): 272–284. https://doi.org/10.1111/j.1365-2699.2010.02422.x
  • Dayton GH, Croll DA, Fuentes Cuevas S (2020) Geographic distribution: Mexico, Baja California Sur: Hemidactylus frenatus. Herpetological Review 51(2): e271.
  • Dejean T, Valentini A, Miquel C, Taberlet P, Bellemain E, Miaud C (2012) Improved detection of an alien invasive species through environmental DNA barcoding: the example of the American bullfrog, Lithobates catesbeianus. Journal of Applied Ecology 49(4): 953–959. https://doi.org/10.1111/j.1365-2664.2012.02171.x
  • de Sousa F, Arosemena F, Castillo JA, Mallorga HM (1989) Una nueva distribución geográfica de Eleutherodactylus antillensis (Reinhardt y Lütken, 1863) (Aamphibia: Anura: Leptodactylidae), identificación y hábitos ecológicos en la ciudad de Panamá. Sciencia (Panamá) 1989: 87–102.
  • de Villa-Meza A, Avila-Flores R, Cuarón AD, Valenzuela-Galván D (2011) Procyon pygmaeus (Carnivora: Procyonidae). Mammalian Species 43(877): 87–93. https://doi.org/10.1644/877.1
  • D’hondt B, Vanderhoeven S, Roelandt S, Mayer F, Versteirt V, Adriaens T, Ducheyne E, San Martin G, Grégoire J-C, Stiers I, Quoilin S, Cigar J, Heughebaert A, Branquart E (2015) Harmonia+ and Pandora+: risk screening tools for potentially invasive plants, animals and their pathogens. Biological Invasions 17(6): 1869–1883. https://doi.org/10.1007/s10530-015-0843-1
  • Dixon JR (2013) Amphibians and Reptiles of Texas: With Keys, Taxonomic Synopses, Bibliography, and Distribution Maps. Texas A&M University Press, College Station, 443 pp.
  • D.O.F. (2015) Proyecto de Modificación del Anexo Normativo III, Lista de especies en riesgo de la Norma Oficial Mexicana NOM-059-SEMARNAT-2010, Protección ambiental-Especies nativas de México de flora y fauna silvestresCategorías de riesgo y especificaciones para su inclusión, exclusión o cambio-Lista de especies en riesgo. Diario Oficial de la Federación, 78 pp.
  • D.O.F. (2016) Acuerdo mediante el cual se dan a conocer en los Estados Unidos Mexicanos las enfermedades y plagas exóticas y endémicas de notificación obligatoria de los animales terrestres y acuáticos. Diario Oficial de la Federación, 61–76.
  • Duellman WE (2001) The Hylid frogs of Middle America (Vol. 2). Society for the Study of Amphibians and Reptiles, Contibutions in Herpetology 18, Society for the Study of Amphibians and Reptiles, Ithaca, New York, 1158 pp.
  • Echternacht A, Burton F, Blumenthal J (2011) The amphibians and reptiles of the Cayman Islands: Conservation issues in the face of invasions. Conservation of Caribbean Island Herpetofaunas 1: 129–147. https://doi.org/10.1163/ej.9789004194083.i-439.49
  • Edwards T, Bonine KE, Ivanyi C, Prescott R (2005) The molecular origins of spiny-tailed iguanas (Ctenosaura) on the grounds of the Arizona-Sonora Desert Museum. Sonoran Herpetologist 18(11): 122–125.
  • Edwards T, Jarchow CJ, Jones CA, Bonine KE (2010) Tracing genetic lineages of captive desert tortoises in Arizona. The Journal of Wildlife Management 7(4): 801–807. https://doi.org/10.2193/2009-199
  • Edwards T, Karl A, Vaughn M, Rosen P, Meléndez Torres C, Murphy RW (2016) The desert tortoise trichotomy: Mexico hosts a third, new sister-species of tortoise in the Gopherus morafkaiG. agassizii group. ZooKeys 562: 131–158. https://doi.org/10.3897/zookeys.562.6124
  • Farr WL (2015) Herpetofauna of Tamaulipas. Chapter 8. In: Lemos-Espinal JA, Hollingsworth B, Mahrdt C, Brennan T, Babb R. and Painter C (Eds) Amphibians and Reptiles of the US-Mexico Border States/Anfibios y reptiles de los estados de la frontera México-Estados Unidos. Texas A&M University Press, 101–121.
  • Finnerty PB, Shine R, Brown GP (2018) The costs of parasite infection: effects of removing lungworms on performance, growth and survival of free-ranging cane toads. Functional Ecology 32(2): 402–415. https://doi.org/10.1111/1365-2435.12992
  • Fitch HS (1975) Sympatry and interrelationships in Costa Rican anoles. Occasional Papers of the Museum of Natural History of the University of Kansas, University of Kansas 40: 1–60. https://doi.org/10.5962/bhl.part.17265
  • Frankenberg E (1984) Interactions between two Species of colonizing house geckos, Hemidactylus turcicus and Hemidactylus garnotii. Journal of Herpetology 18(1): 1–7. https://doi.org/10.2307/1563665
  • Fritz U, Stuckas H, Vargas-Ramírez M, Hundsdörfer AK, Maran J, Päckert M (2012) Molecular phylogeny of Central and South American slider turtles: implications for biogeography and systematics (Testudines: Emydidae: Trachemys. Journal of Zoological Systematics and Evolutionary Research 50(2): 125–136. https://doi.org/10.1111/j.1439-0469.2011.00647.x
  • Frost DR (2020) Amphibian Species of the World: an online database. Version 6.1. American Museum of Natural History, New York, New York. Electronic Database accessible at https://amphibiansoftheworld.amnh.org/index.php [Accessed 6 February 2021]
  • Fujita MK, Moritz C (2009) Origin and evolution of parthenogenetic genomes in lizards: current state and future directions. Cytogenetic and Genome Research 127(2–4): 261–272. https://doi.org/10.1159/000295177
  • Gallardo B, Clavero M, Sánchez Marta I, Vilà M (2015) Global ecological impacts of invasive species in aquatic ecosystems. Global Change Biology 22(1): 151–163. https://doi.org/10.1111/gcb.13004
  • García-Balderas CM, Cedeño-Vázquez JR, Mineros-Ramírez R (2016) Distribution Notes. Eleutherodactylus planirostris. Mesoamerican Herpetology 3(1): e173.
  • García-Llorente M, Martín-López B, González JA, Alcorlo P, Montes C (2008) Social perceptions of the impacts and benefits of invasive alien species: Implications for management. Biological Conservation 141(12): 2969–2983. https://doi.org/10.1016/j.biocon.2008.09.003
  • García-Llorente M, Martín-López B, Nunes PALD, González JA, Alcorlo P, Montes C (2011) Analyzing the social factors that influence willingness to pay for invasive alien species management under two different strategies: eradication and prevention. Environmental Management 48(3): 418–435. https://doi.org/10.1007/s00267-011-9646-z
  • García-Rodríguez A, Chaves G, Wainwright M, Villegas A (2010) Geographic distribution: Costa Rica, Cartago, Turrialba District: Eleutherodactylus coqui. Herpetological Review 41(3): e375.
  • García-Vázquez UO, Canseco-Márquez L, Gutiérrez-Mayén G, Trujano-Ortega M (2009) Actualización del conocimiento de la fauna herpetológica en el estado de Puebla, México. Boletín de la Sociedad Herpetológica Mexicana 17(1): 12–36.
  • Garner TW, Perkins MW, Govindarajulu P, Seglie D, Walker S (2006) The emerging amphibian pathogen Batrachochytrium dendrobatidis globally infects introduced populations of the North American bullfrog, Rana catesbeiana. Biology Letters 2(3): 455–459. https://doi.org/10.1098/rsbl.2006.0494
  • GBIF.org (2018) Global Biodiversity Information Facility. https://www.gbif.org [Accessed 6 February 2021]
  • González-Martínez AI, Barrios-Caballero Y, de Jesús S (2017) Análisis de riesgo para especies invasoras en México. El impacto de las especies exóticas invasoras en México. Revista Legislativa de Estudios Sociales y de Opinión Pública 103: 24–29.
  • González-Sánchez VH, Johnson JD, García-Padilla E, Mata-Silva V, DeSantis DL, Wilson LD (2017) The herpetofauna of the Mexican Yucatan Peninsula: composition, distribution, and conservation. Mesoamerican Herpetology 4(2): 264–380.
  • Greenlees MJ, Harris S, White AW, Shine R (2018) The establishment and eradication of an extra-limital population of invasive cane toads. Biological Invasions 20(8): 2077–2089. https://doi.org/10.1007/s10530-018-1681-8
  • Grismer LL (2002) Amphibians and Reptiles of Baja California, Including its Pacific Islands and the Islands in the Sea of Cortés. University of California Press, Berkeley, California, 409 pp. https://doi.org/10.1525/9780520925205
  • Gutsche A, McCranie JR (2009) Geographic distribution: Honduras, Islas de la Bahía: Hemidactylus mabouia. Herpetological Review 4(1): 112–113.
  • Guzmán-Guzmán S, Palma-Martínez M (2016) Geographic distribution: México, Veracruz, Municipality of Minatitlán: Gonatodes albogularis. Herpetological Review 47(3): e423.
  • Hallowell E (1854) Description of new species of reptilia from western Africa. Proceedings of the Academy of Natural Sciences of Philadelphia 64: 62–65.
  • Harris DM, Kluge AG (1984) The Sphaerodactylus (Sauria: Gekkonidae) of Middle America. Occasional Papers of the Museum of Zoology, University of Michigan (706): 1–59. http://hdl.handle.net/2027.42/57142
  • Henderson R, Villa J, Dixon JR (1976) Lepidodactylus lugubris (Reptilia: Gekkonidae): a recent addition to the herpetofauna of Nicaragua. Herpetological Review 7(4): e173.
  • Hollingsworth B, Mahrdt CR, Grismer LL, Banta BH, Sylber CK (1997) The occurrence of Sauromalus varius on a satellite islet of Isla Salsipuedes, Gulf of California, México. Herpetological Review 28(1): 26–28.
  • Hofman CA, Rick TC (2018) Ancient Biological Invasions and Island Ecosystems: Tracking Translocations of Wild Plants and Animals. Journal of Archaeological Research 26(1): 65–115. https://doi.org/10.1007/s10814-017-9105-3
  • Hoogmoed MS, Avila-Pires TC (2015) Lepidodactylus lugubris (Duméril & Bibron 1836) (Reptilia: Gekkonidae), an introduced lizard new for Brazil, with remarks on and correction of its distribution in the New World. Zootaxa 4000(1): 90–110. https://doi.org/10.11646/zootaxa.4000.1.4
  • Hunter ME, Oyler-McCance SJ, Dorazio RM, Fike JA, Smith BJ (2015) Environmental DNA (eDNA) sampling Improves occurrence and detection estimates of Invasive Burmese Pythons. PLoS ONE 10(4): 1–17. https://doi.org/10.1371/journal.pone.0121655
  • Ibañez R, Rand SA (1990) Geographic distribution: Panama, Province of Panama: Eleutherodactylus johnstonei. Herpetological Review 21(1): e37.
  • Ineich I, Blanc CP (1987) Le peuplement herpétologique de Polynésie française. Adaptations et aléas. Bulletin de la Societe Zoologique de France 112(3–4): 381–400.
  • IUCN (2014) IUCN Red List of Threatened Species. Version 2014.2. www.iucnredlist.org [Accessed 6 February 2021]
  • IUCN (2015) Lithobates catesbeianus [Online]. [Accessed 6 February 2021]
  • Iverson JB, Young CA, Berry JF (1998) Kinosternon integrum LeConte (Mexican Mud Turtle). Society for the Study of Amphibians and Reptiles, Catalogue of American Amphibians and Reptiles (CAAR): 652.1–652.6.
  • Jaramillo C, Wilson LD, Ibáñez R, Jaramillo F (2010) The herpetofauna of Panama: distribution and conservation status. In: Wilson LD, Townsend JH, Johnson JD (Eds) Conservation of Mesoamerican Amphibians and Reptiles. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 604–671.
  • Jiménez-Arcos VH, Cruz-Padilla SS, Escalona-López A, Casas-Andreu G, Centenero-Alcalá E (2009) Geographic distribution: México, Guerrero, Municipality of Copalillo: Apalone spinifera. Herpetological Review 40(4): e448.
  • Jiménez RR, Abarca JG (2014) Extensión de rango del gecko Lepidodactylus lugubris en la vertiente Caribe de Costa Rica. Herpetotropicos 10(1–2): 31–32.
  • Johnson JD, Mata-Silva V, Ramírez-Bautista A (2010) Geographic distribution and conservation of the herpetofauna of southeastern Mexico. In: Wilson LD, Townsend JH, Johnson JD (Eds) Conservation of Mesoamerican Amphibians and Reptiles. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 322–369.
  • Johnson JD, Mata-Silva V, Padilla EG, Wilson LD (2015a) The Herpetofauna of Chiapas, Mexico: composition, distribution, and conservation. Mesoamerican Herpetology 2(3): 272–329.
  • Johnson JD, Mata-Silva V, Wilson LD (2015b) A conservation reassessment of the Central American herpetofauna based on the EVS measure. Amphibian & Reptile Conservation 9(2): 1–94.
  • Kaiser H, Henderson RW (1994) The conservation status of Lesser Antillean frogs. Herpetological Natural History 2(2): 41–56.
  • Kluge AG (1969) The evolution and geographical origin of the New World Hemidactylus mabouia-brooki complex (Gekkonidae, Sauria). Miscellaneous Publications Museum of Zoology. University of Michigan (138): 1–78.
  • Kôhler G (1999) The Amphibians and Reptiles of Nicaragua: A distributional checklist with keys. Courier Forschungsinstitut Senckenberg, Frankfurt a. M. 213: 1–121.
  • Kôhler G (2001) Anfibios y Reptiles de Nicaragua. Herpeton, Verlag Elke Kôhler, Rohrstr. 22, 63075 Offenbach, Germany, 208 pp.
  • Köhler G (2008) Reptiles of Central America (2nd Edn.). Herpeton Verlag Elke Köhler, Offenbach, Germany, 400 pp.
  • Köhler G (2011) Amphibians of Central America. Herpeton, Verlag Elke Köhler, Offenbach, Germany, 379 pp.
  • Kolby JE, Smith KM, Berger L, Karesh WB, Preston A, Pessier AP, Skerratt LF (2014) First evidence of amphibian chytrid fungus (Batrachochytrium dendrobatidis) and Ranavirus in Hong Kong amphibian trade. PLoS ONE 9(3): e90750. https://doi.org/10.1371/journal.pone.0090750
  • Kraus F (2003) Invasion pathways for terrestrial vertebrates. In: Ruiz GM, Carlton JT (Eds) Bioinvasions: Pathways, Vectors, and Management Strategies. Island Press, New York, New York, 68–92.
  • Kuchta SR (2005) Taricha torosa. In: Lannoo MJ (Ed.) Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkley, 904–908.
  • Lavín PA, Lazcano D, Gadsden H (2014) Anfibios y reptiles exóticos y traslocados invasores. In: Mendoza R, Koleff P (Eds) Especies Acuáticas Invasoras en México, Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), Mexico City, Mexico, 435–444.
  • Lazcano-Barrero M (1993) Plan de acción para la conservación del cocodrilo americano (Crocodylus acutus) en Isla Contoy, Quintana Roo. Technical Report, Reserva Ecológica El Edén A. C. , Cancún, Quintana Roo, México, 9 pp.
  • Lazcano D, Mendoza-Alfaro R, Campos-Múzquiz L, Lavin-Murcio PA, Quiñónez-Martínez M (2010) Notes on Mexican herpetofauna 15: the risk of invasive species in northeastern Mexico. Bulletin of the Chicago Herpetological Society 45(7): 113–117.
  • Lee JC (1996) The Amphibians and Reptiles of the Yucatán Peninsula. Comstock Publishing Asociates, Cornell University Press, Ithaca, New York, 500 pp.
  • Leets-Rodríguez L, López-Guevara H, Sunyer J (2019) First country records of the invasive Brahminy Blindsnake Indotyphlops braminus (Daudin, 1803) (Squamata, Typhlopidae) from Nicaragua. Revista Nicaragüense de Biodiversidad (45): 3–11.
  • Lemos-Espinal JA (2015) Amphibians and Reptiles of the US–Mexico Border States. Texas A&M University Press. College Station, 614 pp.
  • Lemos-Espinal JA, Dixon JR (2013) Amphibians and Reptiles of San Luis Potosí. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 300 pp.
  • Lemos-Espinal JA, Smith GR, Woolrich-Peña GA, Cruz A (2017) Amphibians and Reptiles of Chihuahua, Mexico, with comparisons with adjoining states. ZooKeys 658: 105–130. https://doi.org/10.3897/zookeys.658.10665
  • Lemos-Espinal JA, Smith HM (2007a) Anfibios y Reptiles del Estado de Chihuahua, México. UNAM, CONABIO, Mexico, 613 pp.
  • Lemos-Espinal JA, Smith HM (2007b) Anfibios y Reptiles del Estado de Coahuila, México. UNAM, CONABIO, México, 550 pp.
  • Lemos-Espinal JA, Dixon JR (2013) Amphibians and Reptiles of San Luis Potosí. Eagle Mountain Publishing, Eagle Mountain, Utah, 300 pp.
  • Lemos-Espinal JA, Webb RG, Smith HM (1999) Emory’s Softshell Turtle, Apalone spinifera emoryi, in Mexico. Bulletin of the Maryland Herpetological Society 35: 40–42.
  • Liner EA, Casas-Andreu G (2008) Standard Spanish, English and Scientific names of the Amphibians and Reptiles of Mexico. Herpetological Circular 38, Society for the Study of Amphibians and Reptiles, Lawrence, Kansas, 162 pp.
  • Lira-Noriega A, Ramírez H (2016) Análisis de riesgo detallado para cinco especies de reptiles con potencial invasor de alto riesgo para México. In: CONABIO (Ed.) Proyecto GEF Aumentar las Capacidades Nacionales para el Manejo de las Especies Exóticas Invasoras (EEI) a través de la Implementación de la Estrategia Nacional de EEI. Ciudad de México, México, 129 pp.
  • López-González C (1991) Estudio Prospectivo de los Vertebrados Terrestres del Corredor Turístico Cancún-Tulum, Quintana Roo, Mexico. Tesis de Licenciatura, Universidad Nacional Autónoma de Mexico, 127 pp.
  • Lorvelec O, Pascal M (2006) Les vertébrés de clipperton soumis à un siècle et demi de bouleversements écologiques. Revue D’écologie 61(2): 135–158. http://hdl.handle.net/2042/55675
  • Lovich RE, Grismer LL, Danemann G (2009) Conservation status of the herpetofauna of Baja California, México and associated islands in the Sea of Cortez and Pacific Ocean. Herpetological Conservation and Biology 4(3): 358–378.
  • Lovich RE, Mahrdt CR (2007) Herpetofauna terrestre. In: Danemann G, Ezcurra E (Eds) Bahía de Los Angeles: Recursos Naturales y Comunidad. Línea Base 2007. Pronatura Noroeste AC, Secretaría de Recursos Naturales y Medio Ambiente, Instituto Nacional de Ecología y Museo de Historia Natural de San Diego, 495–521.
  • Lowe S, Browne M, Boudjelas S, De Poorter M (2000) 100 of the World’s Worst Invasive Alien Species: A Selection from the Global Invasive Species Database. Invasive Species Specialist Group, Auckland, New Zealand, 12 pp.
  • Luja VH, Blázquez MC, Rodríguez-Estrella R (2007) Tortuga introducida: reporte de Kinosternon integrum (Leconte, 1854) en Baja California Sur. Boletín de la Sociedad Herpetológica Mexicana 15(2): 40–41.
  • Machkour-M’Rabet S, Hénaut Y, Charruau P, Gevrey M, Winterton P, Legal L (2009) Between introgression events and fragmentation, islands are the last refuge for the American crocodile in Caribbean Mexico. Marine Biology 156(6): 1321–1333. https://doi.org/10.1007/s00227-009-1174-5
  • MacIsaac H, Tedla R, Ricciardi A (2010) Patterns and Rate of Growth of Studies. In: Ricardson DM (Ed.) Fifty Years of Invasion Ecology: The Legacy of Charles Elton. Blackwell Publishing Ltd. , Hoboken, New Jersey, 456 pp. https://doi.org/10.1002/9781444329988.ch5
  • Marr SR, Johnson SA, Hara AH, McGarrity ME (2010) Preliminary evaluation of the potential of the helminth parasite Rhabdias elegans as a biological control agent for invasive Puerto Rican Coquís (Eleutherodactylus coqui) in Hawaii. Biological Control 54(2): 69–74. https://doi.org/10.1016/j.biocontrol.2010.04.012
  • Martínez-Hernández A, Pineda-Manzano U, Bauer AM, Vázquez-Villa BM (2017) First record of the Mediterranean House Gecko, Hemidactylus turcicus Linnaeus 1758 in the metropolitan area of San Luis Potosí, Mexico. IRCF Reptiles & Amphibians 24(2): 143–147.
  • Martínez-Morales MA, Cuarón AD (1999) Boa constrictor, an introduced predator threatening the endemic fauna on Cozumel Island, Mexico. Biodiversity & Conservation 8(7): 957–963. https://doi.org/10.1023/A:1008815004072
  • Mata-Silva V, Johnson JD, Wilson LD, Rocha A, García-Grajales J (2013) Geographic distribution: Mexico, Oaxaca: Hemidactylus frenatus. Herpetological Review 44(4): 625–626.
  • Mayer GC (2010) Ecological distribution of the Puerto Rican crested anole, Anolis cristatellus cristatellus, in Cahuita, Costa Rica. Anolis Newsletter VI: 138–144.
  • McCoy CJ (1970) Hemidactylus turcicus. Catalogue of American Amphibians and Reptiles (CAAR), 87.1–87.2.
  • McCranie JR (2015) A checklist of the amphibians and reptiles of Honduras, with additions, comments on taxonomy, some recent taxonomic decisions, and areas of further studies needed. Zootaxa 3931: 352–386. https://doi.org/10.11646/zootaxa.3931.3.2
  • McCranie JR (2018) The lizards, crocodiles, and turtles of Honduras: systematics, distribution, and conservation. Bulletin of the Museum of Comparative Zoology 15(1): 1–129. https://doi.org/10.5962/bhl.title.155244
  • McCranie JR, Gutsche A (2009) Geographic distribution: Honduras, Atlántida: Anolis allisoni. Herpetological Review 40(1): e112.
  • McCranie JR, Gutsche A (2014) Geographic distribution: Honduras, Department of La Paz: Eleutherodactylus (Euhyas) planirostris. Herpetological Review 45(4): e653.
  • McCranie JR, Harrison A, Valdés-Orellana L (2017) Updated Population and Habitat Comments about the reptiles of the Swan Islands, Honduras. Bulletin of the Museum of Comparative Zoology 161(7): 265–284. https://doi.org/10.3099/MCZ34.1
  • McCranie JR, Hedges SB (2012) Two new species of geckos from Honduras and resurrection of Sphaerodactylus continentalis Werner from the synonymy of Sphaerodactylus millepunctatus Hallowell (Reptilia, Squamata, Gekkonoidea, Sphaerodactylidae). Zootaxa 3492: 65–76.
  • McCranie JR, Hedges SB (2013) A review of the Cnemidophorus lemniscatus group in Central America (Squamata: Teiidae), with comments on other species in the group. Zootaxa 3722(3): 301–316. https://doi.org/10.11646/zootaxa.3722.3.1
  • McCranie JR, Köhler G (2015) The anoles (Reptilia: Squamata: Dactyloidae: Anolis: Norops) of Honduras. Systematics, Distribution, and Conservation. Bulletin of the Museum of Comparative Zoology 161(11): 1–280. https://doi.org/10.3099/0027-4100-14.1.1
  • McCranie JR, Köhler F, Gutsche A, Valdés-Orellana L (2013) Trachemys grayi emolli (Testudines, Emydidae) in Honduras and its systematic relationships based on mitochondrial DNA. Zoosystematics and Evolution, Museum für Naturkunde 89(1): 21–29. https://doi.org/10.1002/zoos.201300003
  • McCranie JR, Wilson LD, Köhler G (2005) Amphibians and Reptiles of the Bay Islands and Cayos Cochinos, Honduras. Bibliomania! Salt Lake City, Utah, 210 pp.
  • McCranie JR, Townsend JH, Wilson LD (2006) The Amphibians and Reptiles of the Honduran Mosquitia. Krieger Publishing Company, Malabar, 291 pp.
  • McCranie JR, Valdés-Orellana L (2014) New island records and updated nomenclature of amphibians and reptiles from the Islas de la Bahía, Honduras. Herpetology Notes 7: 41–49.
  • McGaugh SE, Janzen FJ (2008) The status of Apalone atra populations in Cuatro Ciénegas, Coahuila, México: preliminary data. Chelonian Conservation and Biology 7(1): 88–95. https://doi.org/10.2744/CCB-0651.1
  • McNeely JA (2001) An introduction to human dimensions of invasive alien species. In: McNeely JA (Eds) The Great Reshuffling. Human Dimensions of Invasive Alien Species. IUCN. Gland, Switzerland and Cambridge, United Kingdom, 5–22.
  • Measey GJ, Rödder D, Green SL, Kobayashi R, Lillo F (2012) Ongoing invasions of the African clawed frog, Xenopus laevis: a global review. Biological Invasions 14(11): 2255–2270. https://doi.org/10.1007/s10530-012-0227-8
  • Mellink E (2002) Invasive vertebrates on islands of the Sea of Cortés. Invasive exotic species in the Sonoran region. University of Arizona Press, Tucson, 112–125.
  • Mellink E, Ferreira-Bartrina V (2000) On the wildlife of wetlands of the Mexican portion of the Río Colorado Delta. Bulletin of the Southern California Academy of Sciences 99(3): 115–115. https://scholar.oxy.edu/scas/vol99/iss3/2
  • Mertz WH, Costa-Campos CE, Valentim DSS, Furtado MFM (2015) Geographic distribution: México: Guerrero, Municipality of Zihuatanejo: Trachemys ornata. Herpetological Review 46(1): e58.
  • Moll EO (1995) The turtle Trachemys scripta and the pet trade. Aliens 1: 1–3.
  • Morales JR, Chang MJ, Acevedo ME (2017) First records of Hemidactylus garnotii Duméril & Bibron, 1836 (Squamata: Gekkonidae) in Guatemala. Mesoamerican Herpetology 4(4): 953–955.
  • Muller BJ, Schwarzkopf L (2017) Relative effectiveness of trapping and hand-capture for controlling invasive cane toads (Rhinella marina). International Journal of Pest Management 64(2): 185–192. https://doi.org/10.1080/09670874.2017.1363443
  • Muñoz-Alonso LA (2006) Actualización y enriquecimiento de las bases de datos del proyecto de evaluación y análisis geográfico de la diversidad faunística de Chiapas. El Colegio de la Frontera Sur. Informe final SNIB-CONABIO proyecto No. U014. México. http://www.conabio.gob.mx/institucion/proyectos/resultados/InfU014.pdf [Accessed 6 February 2021]
  • Murphy RW (1983) Paleobiogeography and genetic differentiation of the Baja California herpetofauna. Occasional Papers of the California Academy of Sciences 137: 1–48.
  • Murphy RW, Aguirre-León G (2002) The nonavian reptiles: origins and evolution. In: Case TJ, Cody ML, Ezcurra E (Eds) A New Island Biogeography of the Sea of Cortés. Oxford University Press, New York, 181–270. https://doi.org/10.1093/oso/9780195133462.003.0014
  • Murphy RW, Berry KH, Edwards T, Leviton AE, Lathrop A (2011) The dazed and confused identity of Agassiz’s land tortoise, Gopherus agassizii (Testudines, Testudinidae) with the description of a new species, and its consequences for conservation. ZooKeys 113: 39–71. https://doi.org/10.3897/zookeys.113.1353
  • Murphy RW, Méndez de la Cruz FR (2010) The Herpetofauna of Baja California and its associated islands; a conservation assessment and priorities. In: Wilson LD, Townsend JH, Johnson JD (Eds) Conservation of Mesoamerican Amphibians and Reptiles. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 238–273.
  • Nabhan GP (2000) Cultural dispersal of plants and reptiles to the Midriff Islands of the Sea of Cortés: integrating indigenous human dispersal agents into island biogeography. Journal of the Southwest 42(3): 545–558. https://www.jstor.org/stable/40170213
  • Ochoa-Ochoa L, Flores-Villela O, Ríos-Muñoz CA, Arroyo-Cabrales J, Martínez-Gordillo M (2017) Mexico’s ambiguous invasive species plan. Science 355(6329): 1033–1033. https://doi.org/10.1126/science.aam9400
  • O’Hanlon SJ, Rieux A, Farrer RA, Rosa GM, Waldman B (2018) Recent Asian origin of chytrid fungi causing global amphibian declines. Science 360(6389): 621–627. https://doi.org/10.1126/science.aar1965
  • Olson CA, Beard KH, Pitt WC (2012) Biology and impacts of Pacific island invasive species 8. Eleutherodactylus planirostris, the Greenhouse Frog (Anura: Eleutherodactylidae). Science 66(3): 255–270. https://doi.org/10.2984/66.3.1
  • Ortiz-Medina JA, Nahuat-Cervera PE, Cabrera-Cen D, Vega-Marcín A, Cedeño-Vázquez JR (2017) Distribution Notes. First records of the invasive Greenhouse Frog, Eleutherodactylus planirostris (Cope, 1862) (Anura: Eleutherodactylidae) in the state of Yucatán, Mexico. Mesoamerican Herpetology 4(1): 191–193.
  • Ortiz-Medina JA, Cabrera-Cen DI, Chan-Noh MM, Cedeño-Vázquez JR (2019) First record of the Moorish Gecko, Tarentola mauritanica (Linnaeus, 1758) (Squamata: Phyllodactylidae), in Mexico. Herpetology Notes (12): 971–974. https://www.biotaxa.org/hn/article/view/46810/56653
  • Ottley JR, Velázquez-Solis VM (1989) An extant, indigenous tortoise population in Baja California Sur, Mexico, with the description of a new species of Xerobates (Testudines: Testudinidae). The Great Basin Naturalist 49(4): 496–502. https://doi.org/10.5962/bhl.part.22649
  • Parham JF, Papenfuss TJ, van Dijk PP, Wilson BS, Marte C, Schettino LR, Simson WB (2013) Genetic introgression and hybridization in Antillean freshwater Turtles (Trachemys) revealed by coalescent analyses of mitochondrial and cloned nuclear markers. Molecular Phylogenetics and Evolution 67(1): 176–187. https://doi.org/10.1016/j.ympev.2013.01.004
  • Parham JF, Papenfuss TJ, Buskirk JR, Parra-Olea G, Chen J-Y, Simison WB (2015) Trachemys ornata or not ornata: reassessment of a taxonomic revision for Mexican Trachemys. Proceedings of the California Academy of Science (Series 4) 62(4): 359–367.
  • Parker IM, Simberloff D, Lonsdale WM, Goodell K, Wonham M, Kareiva PM, Williamson MH, Von Holle B, Moyle PB, Byers JE, Goldwasser L (1999) Impact: toward a framework for understanding the ecological effects of invaders. Biological Invasions 1(1): 3–19. https://doi.org/10.1023/A:1010034312781
  • Pasachnik SA (2013) Growth, reproduction, and diet of Roatán Spiny-tailed Iguanas, Ctenosaura oedirhina, with notes on the status of the species. Herpetological Conservation and Biology 8(1): 191–198.
  • Peralta-García A, Valdez-Villavicencio JH, Galina-Tessaro P (2014) African clawed frog (Xenopus laevis) in Baja California: a confirmed population and possible ongoing invasion in Mexican watershEds The Southwestern Naturalist 59(3): 431–434. https://doi.org/10.1894/NBF-12.1
  • Perrings C, Dalmazzone S, Williamson M (2005) The economics of biological invasions. In: Mooney HA, Mack R, McNeely JA, Neville LE, Schei PJ, Waage JK (Eds) Invasive Alien Species: A New Synthesis. Island Press, Washington, 16–36.
  • Peterson TA (2003) Predicting the geography of species invasions via ecological niche modeling. The Quarterly Review of Biology 78(4): 419–433. https://doi.org/10.1086/378926
  • Piaggio AJ, Engeman RM, Hopken MW, Humphrey JS, Keacher KL (2014) Detecting an elusive invasive species: a diagnostic PCR to detect Burmese Python in Florida waters and an assessment of persistence of environmental DNA. Molecular Ecology Resources 14(2): 374–380. https://doi.org/10.1111/1755-0998.12180
  • Platt SG, Meerman JC, Rainwater TR (1999) Diversity, observations and conservation of the herpetofauna of Turneffe, Lighthouse and Glovers Atolls, Belize. Bulletin British Herpetological Society 66(1): 1–13.
  • Platz JE, Clarkson RW, Rorabaugh JC, Hillis DM (1990) Rana berlandieri: recently introduced populations in Arizona and Southeastern California. Copeia 1990(2): 324–333. https://doi.org/10.2307/1446338
  • Porras LW, Wilson LD, Schuett GW, Reiserer RS (2013) A taxonomic reevaluation and conservation assessment of the Common Cantil, Agkistrodon bilineatus (Squamata: Viperidae): a race against time. Amphibian & Reptile Conservation 7(1): 48–73.
  • Powell R, Conant R, Collins JT (2016) Peterson’s Field Guide to Reptiles and Amphibians of Eastern and Central North America, Fourth Edition. Houghton Mifflin Harcourt Publishing Company, New York, 494 pp.
  • Powell R, Maxey SA (1990) Hemidactylus brookii. Catalogue of American Amphibians and Reptiles (CAAR), Society for the Study of Amphibians and Reptiles 493.1–493.3.
  • Powell R, Henderson RH, Adler K, Dundee HA (1996) An annotated Check list of West Indian amphibians and reptiles. In: Powell R, Henderson RW (Eds) Contributions to West Indian Herpetology: A Tribute to Albert Schwartz. Society for the Study of Amphibians and Reptiles, Ithaca, 51–93.
  • Pyšek P, Richardson DM, Pergl J, Jarošík V, Sixtová Z (2008) Geographical and taxonomic biases in invasion ecology. Trends in Ecology & Evolution 23(5): 237–244. https://doi.org/10.1016/j.tree.2008.02.002
  • Quijada-Mascareñas A, Canseco-Márquez L (2007) Geographic distribution: Mexico, Nayarit: Gehyra mutilata. Herpetological Review 38(4): e483.
  • Ramírez-Bautista A, Hernández-Salinas U, García-Vázquez UO, Leyte-Manrique A, Canseco-Márquez L (2009) Herpetofauna del Valle de México: Diversidad y Conservación. Universidad Autónoma del Estado de Hidalgo (ICBI), CONABIO, México, D. F., 213 pp.
  • Ramírez-Bautista A, Hernández-Salinas U, Cruz-Elizalde R, Berriozabal-Islas C, Lara-Tufiño D, Goyenechea Mayer-Goyeneche I, Castillo-Cerón JM (2014) Los Anfibios y Reptiles de Hidalgo, México: Diversidad, Biogeografía y Conservación. Sociedad Herpetológica Mexicana, A.C. Pachuca de Soto, Hidalgo, 387 pp.
  • Ramos-Guerra S, Gatica A (2014) Ecología alimentaria de la rana toro Lithobates catesbeianus (Shaw, 1802) en el noroeste de Chihuahua, México. In: Mendoza R, Koleff P (Eds) Especies acuáticas invasoras en México. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, México, 411–429.
  • Rato C, Harris DJ, Carranza S, Machado L, Perera A (2016) The taxonomy of the Tarentola mauritanica species complex (Gekkota: Phyllodactylidae): Bayesian species delimitation supports six candidate species. Molecular Phylogenetics and Evolution 94(A): 271–278. https://doi.org/10.1016/j.ympev.2015.09.008
  • Rato C, Harris DJ, Perera A, Carvalho SB, Carretero MA (2015) A combination of divergence and conservatism in the niche evolution of the Moorish Gecko, Tarentola mauritanica (Gekkota: Phyllodactylidae). PLoS ONE 10(5): e0127980. https://doi.org/10.1371/journal.pone.0127980
  • Ray DA, Dever JA, Platt SG, Rainwater TR, Finger AG (2004) Low levels of nucleotide diversity in Crocodylus moreletii and evidence of hybridization with C. acutus. Conservation Genetics 5(4): 449–462. https://doi.org/10.1023/B:COGE.0000041024.96928.fe
  • Recuero E, Martínez-Solano I, Parra-Olea G, García-París M (2004) Smilisca baudinii (Anura: Hylidae) in Baja California Sur, México. Herpetological Review 35(3): e296.
  • Reynolds RG, Niemiller ML, Revell LJ (2014) Toward a tree-of-life for the boas and pythons: multilocus species-level phylogeny with unprecedented taxon sampling. Molecular Phylogenetics and Evolution 71(1): 201–213. https://doi.org/10.1016/j.ympev.2013.11.011
  • Reynoso F (1990a) Geographic distribution: Mexico, Baja Californias Sur: Gehyra mutilata. Herpetological Review 21(1): e22.
  • Reynoso F (1990b) Geographic distriution: Mexico, Baja California Sur: Hemidactylus frenatus. Herpetological Review 21(1): e22.
  • Rhodin AGJ, Iverson JB, Bour R, Fritz U, Georges A, Shaffer HB (2017) Turtles of the world: annotated checklist and atlas of taxonomy, synonymy, distribution, and conservation status. In: Rhodin AGJ, Iverson JB, Van Dijk PP, Saumure RA, Buhlmann KA, Prichard PCH, Mittermeier RA (Eds) Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group, Chelonian Research Monograph 7: 1–292 https://doi.org/10.3854/crm.7.checklist.atlas.v8.2017
  • Richardson DM, Pyšek P, Carlton JT (2011) A compendium of essential concepts and terminology in invasion ecology. In: Ricardson DM (Ed.) Fifty Years of Invasion Ecology: The Legacy of Charles Elton. Blackwell Publishing Ltd., Hoboken, 409–421. https://doi.org/10.1002/9781444329988.ch30
  • Robinson MD (1972) Chromosomes, Protein Polymorphism, and Systematics of Insular Chuckwalla Lizards (Genus Sauromalus) in the Gulf of California, Mexico. Ph.D. Dissertation, University of Arizona, Tucson, 71 pp.
  • Rocha S, Ineich I, Harris DJ (2009) Cryptic variation and recent bipolar range expansion within the Stumped-Toed Gecko Gehyra mutilata across Indian and Pacific Ocean islands. Contributions to Zoology 78(1): 1–8. https://doi.org/10.1163/18759866-07801001
  • Rodda GH, Fritts TH, Clark CS, Gotte SW, Chiszar D (1999) A state of the art trap for the Brown Tree Snake. In: Rodda GH, Sawai Y, Chiszar D, Tanaka H (Eds) Problem Snake Management: the Habu and the Brown Tree Snake. Cornell University Press, Ithaca, New York, 268–305. https://doi.org/10.7591/9781501737688
  • Rödder D, Solé M, Böhme W (2008) Predicting the potential distributions of two alien invasive housegeckos (Gekkonidae: Hemidactylus frenatus, Hemidactylus mabouia). North-Western Journal of Zoology 4(2): 236–246.
  • Rodriguez D, Cedeño‐Vázquez JR, Forstner MRJ, Densmore LD (2008) Hybridization between Crocodylus acutus and Crocodylus moreletii in the Yucatan Peninsula: II. Evidence from microsatellites. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology 309: 674–686. https://doi.org/10.1002/jez.499
  • Roesler H, Bauer AM, Heinicke MP, Greenbaum E, Jackman T (2011) Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae). Zootaxa 2989: 1–50. https://doi.org/10.11646/zootaxa.2989.1.1
  • Romero-Contreras AT, Ávila-Ramos L (1999) Mesoamérica: historia y reconsideración del concepto. Ciencia Ergo-sum 6: 233–242.
  • Rorabaugh JC, King AD (2013) Geographic distribution: Mexico, Sonora, Municipality of Cananea: Apalone spinifera. Herpetological Review 44(1): 104–105.
  • Rorabaugh JC, Lemos-Espinal JA (2016) A Field Guide to the Amphibians and Reptiles of Sonora, Mexico. ECO Herpetological Publishing and Distribution, Rodeo, New Mexico, 687 pp.
  • Rorabaugh JC, Montoya ES, Gomez-Sapiens MM (2008) Geographic distribution: Mexico, Sonora: Apalone spinifera. Herpetological Review 39(3): e365.
  • Rorabaugh JC, Servoss JM (2006) Geographic distribution: Mexico, Sonora. Rana berlandieri. Herpetological Review 37(1): e102.
  • Rorabaugh JC, Sredl MJ, Miera V, Drost CA (2002) Continued invasion by an introduced frog (Rana berlandieri): southwestern Arizona, southeastern California, and Río Colorado, México. The Southwestern Naturalist 47(1): 12–20. https://doi.org/10.2307/3672797
  • Rösler H, Glaw F (2010) Morphologische variation und taxonomie von Hemidactylus brookii Gray, 1845 und Hemidactylus angulatus Hallowell, 1854 sowie phänotypisch ähnlicher taxa. Spixiana 33(1): 139–160.
  • Ruíz-Pérez GA (1996) Claves Preliminares para Reconocer a los Reptiles de Nicaragua. Centro de Derecho Ambiental y Promoción para al Desarrollo, Managua, Nicaragua, 88 pp.
  • Sakai AK, Allendorf FW, Holt JS, Lodge DM, Molofsky J, With KA, Baughman S, Cabin RJ, Cohen JE, Ellstrand NC, McCauley DE, O’Neil P, Parker IM, Thompson JN, Weller SG (2001) The population biology of invasive species. Annual Review of Ecology and Systematics 32(1): 305–332. https://doi.org/10.1146/annurev.ecolsys.32.081501.114037
  • Salazar-Saavedra M, Loza JC, Fernández M, Martínez-Fonseca J, Dwyer Q, Sunyer J (2015) Chelonoidis carbonarius (Spix, 1824): a member of the Nicaraguan herpetofauna. Mesoamerican Herpetology 2(4): 571–573.
  • Sasa M, Chaves G, Porras LW (2010) The Costa Rican Herpetofauna: Conservation status and future perspectives. In: Wilson LD, Townsend JW, Johnson JD (Eds) Conservation of Mesoamerican Amphibians and Reptiles. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 510–603.
  • Saucedo B, Serrano JM, Jacinto-Maldonado M, Leuven RSEW, Rocha García AA, Méndez Bernal A, Gröne A, Van Beurden SJ, Escobedo-Bonilla CM (2019) Pathogen risk analysis for wild amphibian populations following the first report of a Ranavirus outbreak in farmed American Bullfrogs (Lithobates catesbeianus) from northern Mexico. Viruses 11(1): 11–26. https://doi.org/10.1016/j.ympev.2010.06.014
  • Savage JM (2002) The Amphibians and Reptiles of Costa Rica: a Herpetofauna Between Two Continents, Between Two Seas. University of Chicago Press, Chicago, 934 pp.
  • Schloegel LM, Picco AM, Kilpatrick AM, Davies AJ, Hyatt AD, Daszak P (2009) Magnitude of the US trade in amphibians and presence of Batrachochytrium dendrobatidis and Ranavirus infection in imported North American bullfrogs (Rana catesbeiana). Biological Conservation 142(7): 1420–1426. https://doi.org/10.1016/j.biocon.2009.02.007
  • Schwartz A, Thomas R (1975) A Check-list of West Indian Amphibians and Reptiles. Carnegie Museum of Natural History Special Publication No. 1. Pittsburgh, Pennsylvania, 216 pp. https://doi.org/10.5962/bhl.title.123681
  • Seidel ME (2002) Taxonomic observations on extant species and subspecies of slider turtles, genus Trachemys. Journal of Herpetology 36(2): 285–292. https://doi.org/10.2307/1566003
  • Serrano-Gómez SS, Guevara-Chumacero LM, Barriga-Sosa IDLA, Ullóa-Arvízu R, González-Guzmán S, Vázquez-Peláez CG (2016) Low levels of genetic diversity in Crocodylus acutus in Oaxaca and Guerrero, Mexico, and molecular-morphological evidence of the presence of C. moreletii. Biochemical Systematics and Ecology 69: 51–59. https://doi.org/10.1016/j.bse.2016.08.005
  • Shine C, Williams N, Gündling L (2000) A guide to designing legal and institutional frameworks on alien invasive species. IUCN, Gland, Switzerland, Cambridge, United Kingdom, and Bonn, Germany, 138 pp.
  • Shine R (2015) Reducing the Ecological Impact of Invasive Cane Toads. In: Canning-Clode J (Ed.) Biological Invasions in Changing Ecosystems: Vectors, Ecological Impacts, Management and Predictions. Warsaw: Walter de Gruyter. Warsaw, 301–317. https://doi.org/10.1515/9783110438666-019
  • Simberloff D (2009) We can eliminate invasions or live with them: successful management projects. In: Langor DW, Sweeney J (Eds) Ecological Impacts of Non-Native Invertebrates and Fungi on Terrestrial Ecosystems. Springer, Dordrecht, 149–157. https://doi.org/10.1007/978-1-4020-9680-8_11
  • Slevin JR (1928) The amphibians of western North America. Occasional Papers of the California Academy of Sciences 16: 1–152.
  • Smith HM, Burger L (1949) A new subspecies of Anolis sagrei from the Atlantic coast of tropical América. Anales del Instituto de Biología 20: 407–410.
  • Smith HM, Taylor EH (1948) An annotated checklist and key to the amphibians of Mexico. Smithsonian Institution, United States National Museum Bulletin 194, United States Government Printing Office, Washington, 118 pp. https://doi.org/10.5479/si.03629236.194
  • Solís J, Wilson L, Townsend J (2014) An updated list of the amphibians and reptiles of Honduras, with comments on their nomenclature. Mesoamerican Herpetology 1(1): 123–144.
  • Sosa-Tovar G, García-Padilla E, Wilson LD, Mata-Silva V (2019) Geographic distribution: Mexico, Tamaulipas, Municipality of Tampico: Hemidactylus mabouia. Herpetological Review 50(4): e746.
  • Stafford PJ, Meyer JR (2000) A Guide to the Reptiles of Belize. The Natural History Museum, London, Academic Press, London, United Kingdom, 356 pp.
  • Stafford PJ, Walker P, Edgar P, Pennj MG (2010) Distribution and conservation of the Herpetofauna of Belize. In: Wilson LD, Townsend JH, Johnson JD (Eds) Conservation of Mesoamerican Amphibians and Reptiles. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 370–404.
  • Standfuss B, Lipovšek G, Fritz U, Vamberger M (2016) Threat or fiction: is the pond slider (Trachemys scripta) really invasive in central Europe? A case study from Slovenia. Conservation Genetics 17(3): 557–563. https://doi.org/10.1007/s10592-015-0805-2
  • Stebbins RC (2003) A Field Guide to Western Reptiles and Amphibians. The Peterson Field Guide Series. Houghton Mifflin Company. New York, New York, 533 pp.
  • Stejneger L (1904) Herpetology of Porto Rico. Annual Report of the U.S. National Museum. U.S. Government Printing Office, Washington, 551–724.
  • Stuart LC (1955) A brief review of the Guatemalan lizards of the genus Anolis. Miscellaneous Publications of the Museum of Zoology University of Michigan (91): 1–31. http://hdl.handle.net/2027.42/56336
  • Stuart LC (1963) A checklist of the herpetofauna of Guatemala. Miscellaneous Publications of the Museum of Zoology, University of Michigan (122): 1–148. http://hdl.handle.net/11154/99592
  • Suárez-Atilano M, Rojas-Soto O, Parra J, Vázquez-Domínguez E (2017) The role of the environment on the genetic divergence between two Boa imperator lineages. Journal of Biogeography 44(9): 2045–2056. https://doi.org/10.1111/jbi.13006
  • Sunyer J, Nicholson KE, Phillips JG, Gubler JA, Obando LA (2013) Lizards (Reptilia: Squamata) of the Corn Islands, Caribbean Nicaragua. Check List 9(6): 1383–1390. https://doi.org/10.15560/9.6.1383
  • Sunyer J, Martínez-Fonseca J (2019) Fichas de las especies de anfibios y reptiles que han sido registradas para Nicaragua posterior a HerpetoNica (2015). Revista de Temas Nicaraguenses (139): 36–61.
  • Terán-Juárez SA, García-Padilla E, García-Morales LJ (2015) New records and distributional range extensions for amphibians and reptiles from Tamaulipas, México. Mesoamerican Herpetology 2(2): 208–213.
  • Terán-Juárez SA, García-Padilla E, Mata-Silva V, Johnson JD, Wilson LD (2016) The herpetofauna of Tamaulipas, Mexico: composition, distribution, and conservation. Mesoamerican Herpetology 3(1): 42–113.
  • Tepos-Ramírez M, Acosta-Ramírez A, Jiménez-Velázquez G, Álvarez-Alvarado L (2019) Geographic distribution: Mexico, Querétaro, Municipality of Querétaro: Hemidactylus turcicus. Herpetological Review 50(3): e526.
  • Tingley R, Ward-Fear G, Schwarzkopf L, Greenlees MJ, Phillips BL, Brown G, Clulow S, Webb J, Capon R, Sheppard A, Strive T, Tizard M, Shine R (2017) New weapons in the toad toolkit: a review of methods to control and mitigate the biodiversity impacts of invasive cane toads (Rhinella marina). The Quarterly Review of Biology 92(2): 123–149. https://doi.org/10.1086/692167
  • Toda M, Takahashi H, Nakagawa N, Sukigara N (2010) Ecology and control of the green anole (Anolis carolinensis), an invasive alien species on the Ogasawara Islands. In: Kawakami K, Okochi I (Eds) Restoring the Oceanic Island Ecosystem. Springer, Tokyo, 145–152. https://doi.org/10.1007/978-4-431-53859-2_22
  • Townsend CH (1890) Scientific results of explorations by the US Fish Commission steamer Albatross. No. XV. Reptiles from Clarion and Socorro Islands and the Gulf of California, with description of a new species. Proceedings of the United States National Museum 13: 143–144. https://doi.org/10.5479/si.00963801.800.143
  • Townsend JH, Wilson LD (2010) Conservation of the Honduran herpetofauna: issues and imperatives. In: Wilson LD, Townsend JH, Johnson JD (Eds) Conservation of Mesoamerican Amphibians and Reptiles. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 186–705.
  • Tyler MJ (2006) Cane toad pheromones. In: Molloy K, Henderson W (Eds) Science of Cane Toad Invasion and Control. Proceedings of the Invasive Animals CRC/CSIRO/Qld NRM&W Cane Toad Workshop. Brisbane, Queensland, 163–164.
  • Upton DE, Murphy RW (1997) Phylogeny of the Side-Blotched Lizards (Phrynosomatidae: Uta) based on mtDNA sequences: support for a midpeninsular seaway in Baja California. Molecular Phylogenetics and Evolution 8(1): 104–113. https://doi.org/10.1006/mpev.1996.0392
  • Vázquez-Domínguez E, Suárez-Atilano M, Booth W, González-Baca C, Cuarón AD (2012) Genetic evidence of a recent successful colonization of introduced species on islands: Boa constrictor imperator on Cozumel Island. Biological Invasions 14(10): 2101–2116. https://doi.org/10.1007/s10530-012-0217-x
  • Velarde E, Hollingsworth B, Rebman J (2008) Geographic distribution: Mexico, Baja California: Sauromalus hispidus. Herpetological Review 39(3): e368.
  • van Dijk PP, Hammerson G, Vazquez Diaz J, Quintero Diaz GE, Santos G, Flores-Villela O (2007) Kinosternon hirtipes (errata version published in 2016). The IUCN Red List of Threatened Species 2007: e.T63670A97381507, [Online]. [Accessed 6 February 2021]
  • van Wilgen NJ, Roura-Pascual N, Richardson DM (2009) A quantitative climate-match score for risk-assessment screening of reptile and amphibian introductions. Environmental Management 44(3): 590–607. https://doi.org/10.1007/s00267-009-9311-y
  • Villa J, Wilson LD, Johnson JD (1988) Middle American Herpetology: A Bibliographic Checklist. University of Missouri Press, Columbia, 131 pp.
  • Vitousek PM, D’Antonio CM, Loope LL, Rejmanek M, Westbrooks R (1997) Introduced species: a significant component of human-caused global change. New Zealand Journal of Ecology 21(1): 1–16. https://www.jstor.org/stable/24054520
  • Wallach V (2009) Ramphotyphlops braminus (Daudin): a synopsis of morphology, taxonomy, nomenclature and distribution (Serpentes: Typhlopidae). Hamadryad 34(1): 34–61.
  • Wallach V, Williams KL, Boundy J (2014) Snakes of the World: A Catalogue of Living and Extinct Species, CRC Press, Boca Raton, Florida, 1237 pp. https://doi.org/10.1201/b16901
  • Walsh JR, Carpenter SR, Vander Zanden MJ (2016) Invasive species triggers a massive loss of ecosystem services through a trophic cascade. Proceedings of the National Academy of Sciences 113(15): 4081–4085. https://doi.org/10.1073/pnas.1600366113
  • Webb RG (1973) Trionyx atra Webb and Legler: Black Softshell Turtle. Catalogue of American Amphibians and Reptiles (CAAR): 137.1.
  • Weiss AJ, Hedges SB (2007) Molecular phylogeny and biogeography of the Antillean geckos Phyllodactylus wirshingi, Tarentola americana, and Hemidactylus haitianus (Reptilia, Squamata). Molecular Phylogenetics and Evolution 45(1): 409–416. https://doi.org/10.1016/j.ympev.2007.01.006
  • Weterings R, Vetter KC (2017) Invasive house geckos (Hemidactylus spp.): their current, potential and future distribution. Current Zoology 64(5): 559–573. https://doi.org/10.1093/cz/zox052
  • Wilson LD, Johnson JD (2010) Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot. In: Conservation of Mesoamerican Amphibians and Reptiles. Wilson LD, Townsend JH, Johnson JD (Eds) Eagle Mountain Publishing, LC, Eagle Mountain, Utah, 30–235.
  • Wilson LD, Mata-Silva V, Johnson JD (2013a) A conservation reassessment of the Reptiles of Mexico based on the EVS measure. Amphibian & Reptile Conservation 7(1): 2–46.
  • Wilson LD, Johnson JD, Mata-Silva V (2013b) A conservation reassessment of the amphibians of Mexico based on the EVS measure. Amphibian & Reptile Conservation 7(1): 98–127.
  • Witmer GW, Snow NP, Moulton RS (2015) Efficacy of potential chemical control compounds for removing invasive American bullfrogs (Rana catesbeiana). SpringerPlus 4(1): 1–5. https://doi.org/10.1186/s40064-015-1319-6
  • Woolrich-Peña GA, García-Padilla E, DeSantis DL, Johnson DJ, Mata-Silva V, Wilson LD (2017) The herpetofauna of Puebla, Mexico: composition, distribution, and conservation status. Mesoamerican Herpetology 4(4): 2–95.
  • Woolrich-Piña GA, Ponce-Campos P, Loc-Barragán J, Ramírez-Silva JP, Mata-Silva V, Johnson JD, García-Padilla E, Wilson LD (2016) The herpetofauna of Nayarit, Mexico: composition, distribution, and conservation. Mesoamerican Herpetology 3(2): 376–448.
  • Yáñez-Arenas C, Díaz-Gamboa L, Rodríguez-Pérez A, Salmerón-Flores A, Patrón-Rivero C (2016) Análisis de riesgo de tortugas con potencial invasor en México. In: CONABIO and PNUD (Eds) GEF 0089333 “Aumentar las Capacidades Nacionales para el Manejo de las Especies Exóticas Invasoras (EEI) a través de la Implementación de la Estrategia Nacional de EEI, ” Ciudad de México: Unidad Académica de Yucatan-UNAM, 84 pp.
  • Zaldı́var-Riverón A, León-Regagnon V, Nieto-Montes de Oca A (2004) Phylogeny of the Mexican coastal leopard frogs of the Rana berlandieri group based on mtDNA sequences. Molecular Phylogenetics and Evolution 30(1): 38–49. https://doi.org/10.1016/S1055-7903(03)00141-6
  • Zamora-Abrego JG, García-Vázquez UO, Canseco-Márquez L, Nieto-Montes de Oca A (2006) Geographic Distribution: Mexico, Veracruz, Municipality of Minatitlán: Anolis sagrei. Herpetological Review 37(4): e493.