Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64536, mayo 2025 (Publicado May. 15, 2025)

Annotated list of amphibians and reptiles of Costa Rica: The role of the

Museum of Zoology in cataloging the country’s herpetological diversity

Mahmood Sasa1, 2, 3 *; https://orcid.org/0000-0003-0118-5142

Erick Arias1, 2; https://orcid.org/0000-0002-4449-1070

Gerardo Chaves2; https://orcid.org/0000-0002-4301-6569

1. Escuela de Biología, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica; msasamarin@gmail.com

(*Correspondence), eapiedra@gmail.com

2. Museo de Zoología, Centro de Investigaciones en Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa

Rica; cachi13@gmail.com

3. Instituto Clodomiro Picado, Facultad de Microbiología, Universidad de Costa Rica.

Received 02-IX-2024. Corrected 14-III-2025. Accepted 14-III-2025.

ABSTRACT

Introduction: The herpetological collection of the Museum of Zoology at the University of Costa Rica (MZUCR)

has been key to safeguarding a sample of the diversity of amphibians and reptiles of Costa Rica and maintaining

an up-to-date catalog of species present in the country. This unit develops phylogenetic and biogeographic stud-

ies on these animals and plays an important role in taxonomic decisions about them.

Objectives: In this work, we review and update the list of amphibians and reptiles of Costa Rica, commenting on

cases with some taxonomic controversy. In addition, we briefly account for the MZUCR herpetological collection

and explore its history.

Methods: We update the list of present species based on identifying specimens kept in our collection and review-

ing relevant scientific literature. In addition, we compile the information available in internal documents and

scientific publications to establish the history of the collection.

Results: We recognized 221 species of amphibians and 257 species of reptiles in Costa Rica, representing 17 and

36 families, respectively. More than 99% of amphibian species and 97% of reptiles are represented in our collec-

tion, which currently has more than 24 000 properly cataloged specimens. The collection houses 39 holotypes

and 190 paratypes of 47 species, demonstrating its key role in taxonomic studies of the local herpetofauna.

Although amphibian and reptile sampling cover almost the entire country, the northern lowlands and the

Caribbean foothills of the Talamanca Mountain range require more significant collecting efforts.

Conclusions: Despite its long and intense history of biological exploration, records of Costa Rica’s herpetologi-

cal diversity continue to increase. Much of this growth is a result of efforts made at the MZUCR. Since its origin

as a modest private collection in the early 1960s, it has grown into one of the most comprehensive collections

of amphibians and reptiles inhabiting Costa Rica. It also functions as a research unit that generates substantial

biological knowledge about both groups.

Keywords: Costa Rican herpetofauna; checklist; MZUCR; CIBET; Universidad de Costa Rica.

https://doi.org/10.15517/rev.biol.trop..v73iS2.64536

SUPPLEMENT

SECTION: MUSEUM

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64536, mayo 2025 (Publicado May. 15, 2025)

INTRODUCTION

Despite its reduced surface area, just over

51 000 km2, Costa Rica maintains an extraor-

dinary diversity of amphibians and reptiles that

allows it to surpass even megadiverse countries

(sensu Mittermeier et al., 1998) when estimates

are made per surface area unit (Savage, 2002;

Sasa et al., 2010). This small country has one

of the region’s oldest traditions in herpeto-

logical studies, spanning more than 150 years

of studies in this field (Cope, 1875; Picado,

1931; Savage, 2002). Because of this long his-

tory, the number of amphibian and reptile

species inhabiting Costa Rica has not remained

stable: new species are constantly being added

to the lists of recognized species, including new

descriptions, synonymized species resurrec-

tions, and new country distribution records.

Taylor (1952a), Taylor (1952b), Taylor

(1954a), Taylor (1954b), Taylor (1956) under-

took the first comprehensive work on the

herpetofauna of Costa Rica during the twen-

tieth century, laying the foundation for future

research. His report included a staggering 143

amphibians (113 Anura, 27 Caudata, and 3

Gymnophiona) and 227 reptiles (76 Saurian,

and 151 Serpentes). Savage & Villa (1986) iden-

tified 150 amphibians and 212 reptiles thirty

years later, building upon Taylor’s work. In his

magnum opus on Costa Rican herpetofauna,

Savage (2002) reported 172 amphibians and

211 reptiles inhabiting this country almost

two decades afterward. A few years later, the

numbers increased again: Savage & Bolaños

(2009) updated the list to 190 amphibians and

235 reptiles, while Sasa et al. (2010) recognized

189 amphibians and 231 reptiles. More recently,

Leenders (2016) Leenders (2019) recognized

207 amphibians and 245 reptiles in the country,

continuing the legacy of previous researchers

(Fig. 1).

At the institutional level, the herpetofau-

na catalog is compiled in the Herpetological

RESUMEN

Lista anotada de anfibios y reptiles de Costa Rica: el papel del Museo de Zoología

en la catalogación de la diversidad herpetológica del país

Introducción: La colección herpetológica del Museo de Zoología de la Universidad de Costa Rica (MZUCR) ha

sido clave para resguardar una muestra de la diversidad de anfibios y reptiles de Costa Rica y mantener actuali-

zado el catálogo de las especies presentes en el país. Esta unidad desarrolla estudios filogenéticos y biogeográficos

sobre estos animales y juega un papel importante en las decisiones taxonómicas sobre ellos.

Objetivos: En este trabajo revisamos y actualizamos el listado de anfibios y reptiles de Costa Rica, comentando

casos con cierta controversia taxonómica. Además, hacemos un breve recuento de la colección herpetológica del

MZUCR y exploramos su historia.

Métodos: A partir de la identificación de especímenes mantenidos en nuestra colección y la revisión de literatura

científica relevante, actualizamos la lista de especies presentes. Además, recopilamos la información disponible en

documentos internos y publicaciones científicas para establecer la historia de la colección.

Resultados: Reconocemos 221 especies de anfibios y 257 de reptiles en Costa Rica, representando 17 y 36 fami-

lias, respectivamente. Más del 99% de las especies de anfibios y el 97% de reptiles están representadas en nuestra

colección, que actualmente cuenta con más de 24 000 ejemplares debidamente catalogados. La colección alberga

39 holotipos y 190 paratipos de 47 especies, lo que demuestra su papel clave en los estudios taxonómicos de la

herpetofauna local. A pesar de que el muestreo de anfibios y reptiles cubre casi todo el país, aun las tierras bajas

del norte y las estribaciones caribeñas de la Cordillera de Talamanca requieren de mayores esfuerzos de colecta.

Conclusiones: A pesar de su intensa y larga historia de exploración biológica, los registros de la diversidad her-

petológica de Costa Rica continúan aumentando. Gran parte de este crecimiento es resultado de los esfuerzos

realizados en el MZUCR. Desde su origen como una modesta colección privada a principios de la década de 1960,

se ha convertido en una de las colecciones más completas de anfibios y reptiles que habitan Costa Rica y funciona

como una unidad de investigación que genera un conocimiento biológico sustancial sobre ambos grupos.

Palabras clave: MZUCR; CIBET; lista de especies; herpetofauna costarricense; Universidad de Costa Rica.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64536, mayo 2025 (Publicado May. 15, 2025)

Collection of the Museum of Zoology at the

University of Costa Rica (MZUCR). This unit,

now part of the Center for Research in Biodiver-

sity and Tropical Ecology (CIBET) of the same

university (https://cibet.ucr.ac.cr/), evaluates

the taxonomic changes proposed by the spe-

cialized scientific community. It is also a center

for promoting the analysis of local diversity for

both classes of vertebrates, mainly through the

taxonomic review of the specimens preserved

there, their biogeography, and the exploration

of little-studied regions of the country.

In recent years, the advance of molecular

techniques to analyze the evolutionary rela-

tionships of taxonomically complex groups

(e.g., Terrarana anurans, Plethodontid sala-

manders) has generated a series of changes in

the nomenclature of many amphibians and

reptiles and in our understanding of the species

that inhabit the country. Added to this factor is

the description of new species, many of which

were derived from exploring remote regions

in the Talamanca Mountain range by MZUCR

staff and associates (Arias et al., 2018; Arias et

al., 2023; García-Rodríguez et al., 2016). For

these reasons, we present here an updated list

of amphibian and reptile species recognized

in Costa Rica. We also briefly describe the

MZUCR herpetological collection and discuss

its origin and development.

MATERIALS AND METHODS

To account for the amphibian and reptile

species in Costa Rica, we reviewed the material

available in our collection and the descriptions

of species not represented there. In addition,

the data on admissions and the number and

identity of specimens were obtained from the

collection’s physical and digital catalogs. We

examined the specialized literature for new

taxonomic proposals for the species known to

the country. To do so, we relied on two recog-

nized databases that are periodically reviewed:

Amphibian Species of the World: An Online

Reference. (version 6.2 https://amphibiansoft-

heworld.amnh.org/index.php) and The Reptile

Database (http://www.reptile-database.org).

This review of the literature and our taxonomic

criteria allowed us to endorse the classifica-

tion presented here. To prepare the historical

synopsis of the herpetological collection at

MZUCR, we used internal museum reports

as well as information recently published by

other curators of this unit (Angulo et al., 2016;

Springer, 2019).

RESULTS

List of amphibians and reptiles of Costa

Rica: Based on the review of specimens housed

Fig. 1. Cumulative number of amphibian and reptile species that inhabit Costa Rica by year of description. Figura 1. Número

acumulado de especies de anfibios y reptiles que habita en Costa Rica por año de descripción.

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64536, mayo 2025 (Publicado May. 15, 2025)

at the MZUCR and our literature review, we

recognize 221 species of amphibians and 257

reptiles that inhabit Costa Rica (Appendix 1).

Amphibians are grouped into 48 genera and 17

families, while 113 genera and 36 families of

reptiles are recognized (Table 1).

Our findings reveal a critical situation: 10

species of anurans are now considered extinct

in the country, with no individual found in

the last 25 years. This list includes five species

in the genera Craugastor, three Atelopus, and

two Incilius species including the emblematic

golden toad I. periglenes (Appendix 1). Fur-

thermore, three species of anurans (genera

Eleutherodactylus and Osteopilus), six species

of lizards (genera Hemidactylus and Anolis),

and a single snake (Virgotyphlops braminus) are

among the exotic species currently inhabiting

the country (Appendix 1).

Herpetological collection at MZUCR:

The Herpetological collection at MZUCR cur-

rently houses over 24 000 specimens represent-

ing 218 species of amphibians and 248 reptiles

from Costa Rica. This represents a little over

99% and 97% of the known species for each

group, respectively. The specimens come from

localities that cover almost the entire surface

of the country (Fig. 2). However, there are still

regions that have not been adequately sampled,

especially the lowlands in the northwest of the

country and the foothills of the Talamanca

Mountain range on the Caribbean side (Fig. 2).

In addition, the collection is not restricted to

species distributed in Costa Rica but includes

material from other countries (Table 1) that is

used mainly for educational purposes in spe-

cialized courses offered at the School of Biology.

The herpetological collection is predomi-

nantly wet, consisting of specimens fixed in

10% formalin and preserved in jars with 70%

ethanol. The collection, handling, and preser-

vation of specimens strictly adhere to Costa

Rican legislation, following permits from the

Table 1

Number of families, genera and species represented in the amphibian and reptile collection of the Museum of Zoology of

the University of Costa Rica.

Taxa from Costa Rica Taxa from other countries

# Families # Genera # Species # Families # Genera # Species

Amphibians 17 48 217 43 93 152

Caecilians 2 4 9 0 0 0

Salamanders 1 3 56 11 27 47

Anura 14 41 152 32 66 105

Reptiles 36 106 248 38 103 157

Crocodilians 2 2 2 1 1 1

Turtles 6 9 14 4 5 10

Sauria 16 33 94 23 51 81

Snakes 12 62 138 8 43 62

Amphisbaenians – – – 2 3 3

Total taxa 53 154 465 81 196 309

Fig. 2. Collection locations of amphibians and reptiles

cataloged and protected in the MZUCR.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64536, mayo 2025 (Publicado May. 15, 2025)

National System of Conservation Areas of the

Ministry of Environment and Energy of Costa

Rica and protocols approved by the Institution-

al Committee for the Care and Use of Animals

(CICUA) of the University of Costa Rica. Once

preserved, specimens are labeled with a unique

collection number that retains their identity

and collection information, ensuring a well-

organized and managed collection. The jars

are stored on metal shelves, organized by class,

family, and genus in alphabetical order, within

a standard room that the herpetology collection

shares with wet collections of other taxonomic

groups (Angulo et al., 2016). The collection

room has a constant temperature of 15–17 ºC

and a relative humidity of 60–65% to safeguard

the integrity of the collection. Periodic checks

are made of the alcohol level in the collection,

and partial changes of this preservative are

made to protect the specimens. Cleaning and

fumigation of the collection room are also car-

ried out periodically.

The herpetology collection also guards

a significant number of type-specimens that

confirm the descriptions, variation, and type

localities of described species in the country.

This includes 39 holotypes (Table 2) and 190

paratypes of 47 species. This material is essen-

tial for the biodiversity catalog and to ensure

adequate management of the species’ identity in

the country. The collection also includes at least

3 000 tissue samples from preserved specimens,

usually muscle or skin. This material is fixed

in 95% EtOH and stored in a freezer at -20°C

(Gamble, 2014).

DISCUSSION

Our species list presents novel findings,

increasing the number of amphibians previous-

ly reported in the country by between 5.7% and

8.9%, while the number of reptiles increased by

between 1.3% and 5.5% (Leenders, 2016; Leen-

ders, 2019; Sasa et al., 2010). Equally important

Table 2

Holotypes of amphibians and reptiles housed in the Museum of Zoology of the University of Costa Rica.

UCR

voucher Species (Caudata) UCR

voucher Species (Anura) UCR

voucher Species (Squamata)

22842 Bolitoglossa aurae 8042 Atelopus chirripoensis 999 Drymobius margaritiferus

19893 B. aureogularis 22961 Craugastor aenigmaticus 23185 Lachesis melanocephala

22965 B. bolanosi 21864 C. gabbi 5579 Anolis riparius

22420 B. chiquitica 22703 C. zunigai 2463 A. robinsoni

5217 B. diminuta 21843 Diasporus amirae 11642 Porthidium volcanicum

9378 B. gracilis 20493 D. ventrimaculatus 13983 Sibon lamari

20852 B. kamuk 14119 Ecnomiohyla sukia 3592 Sphaerodactylus graptolaemus

2445 B. obscura 22038 Hyalinobatrachium dianae 1888 Tretanorhinus nigroluteus

11788 B. pygmaea 8717 Incilius aucoinae

11216 B. robinsoni 16855 I. chompipe

20489 B. silentium 12877 I. guanacaste

19835 B. splendida 23700 Tlalocohyla celeste

22900 Nototriton costaricense

23685 N. kenorum

23694 N. lateomuscus

23689 N. máximo

23681 N. vereh

22845 Oedipina berlini

8391 O. nimaso

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64536, mayo 2025 (Publicado May. 15, 2025)

are recent taxonomic changes that have led to

discrepancies in the nomenclature between our

list and those previously published, a key point

that we address here.

Gymnophiona: Dubois et al. (2021) placed

Dermophis in synonymy with Gymnopis.

However, Frost (2024) avoided validating this

change due to the lack of information on the

specimens used by San Mauro et al. (2014) who

recovered Gymnopis as paraphyletic concerning

Dermophis. We recognize both genera pending

new data on these phylogenetic relationships.

Caudata: Boza-Oviedo et al. (2012) pro-

pose that Bolitoglossa sombra could be consid-

ered a junior synonym of B. nigrescens, given

the absence of sequence divergence between

the two taxa and the subtle morphological

differences noted in the original description

by Hanken et al. (2005). There is also a pos-

sibility that B. obscura is synonymous with B.

nigrescens. However, to definitively establish

the taxonomic status of these species, new evi-

dence is urgently needed. Until such evidence

emerges, we acknowledge only B. nigrescens.

Furthermore, we do not acknowledge B. indio

as a species present in the country, in con-

trast to the claims of Sunyer et al. (2012) and

Leenders (2016). This is primarily due to its

morphological resemblance to B. alvaradoi, the

polymorphism observed within B. alvaradoi,

and the crucial lack of molecular data that

could provide clarity on the status of these spe-

cies in the country.

Anura: Ryan et al. (2010) described Pris-

timantis educatoris as a closely related but

distinct species to P. caryophyllaceus based on

specimens from the El Cope region of Panama.

However, these authors suggest its presence in

southeastern Costa Rica. We follow Batista et al.

(2014), who synonymized P. educatoris under

P. caryophyllaceus until we have more evidence

that both species occur in our country.

In our species list, we follow Dubois et

al. (2021) and Duellman et al. (2016), who

recognized Phyllomedusidae and Hylidae as

separate families. Within this last family, Sala-

zar-Zúñiga et al. (2019) recorded Ecnomio-

hyla veraguensis in Costa Rica. However, the

specimens reviewed by Salazar-Zúñiga et al.

(2019) seem to correspond more to E. sukia.

Therefore, E. veraguensis in Costa Rica has yet

to be confirmed.

Ron et al. (2016) recognized the polytypic

species Trachycephalus typhonius as paraphy-

letic relative to other species in the genus. In

resolving the identity of the populations of this

species from western Ecuador and Peru and

distinguishing them from T. typhonius (sensu

stricto) from Suriname, these authors left the

populations from Mexico and Central America

without an assigned name. There is one name

available for these populations: Trachycephalus

vermiculatus, although we recommend its for-

mal assignment until a more robust analysis of

the Mesoamerican populations and their phy-

logenetic position is in place (see commentary

in Frost, 2024).

We include the microhylid Elachistocleis

pearsei in our checklist of anurans from Costa

Rica, based on the recent report by Vargas &

Barrio-Amoros (2023). These authors regis-

tered the species for Laurel de Corredores (far

southeast of the country) based on a photo-

graph. However, to date, no voucher specimen

has been collected, and it remains to be con-

firmed whether this is an isolated case or an

effectively established population in the area.

This uncertainty underscores the need for fur-

ther research on this species.

Our list includes Lithobates forreri as one of

the leopard frog species present in the country.

Researchers agree that L. forreri consists of a

species complex (see, for example, Zaldívar-

Riverón et al., 2004). However, there is still no

clarity regarding the identity of several popula-

tions assigned to this species throughout its

distribution, including those in Costa Rica. As a

result, we continue to refer to them as L. forreri,

recognizing the need for further research.

Testudines: The Pacific population of Che-

lonia mydas has been referred to as a subspe-

cies, Chelonia mydas agassizi Bocourt 1868.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64536, mayo 2025 (Publicado May. 15, 2025)

However, some authors argue that morpho-

logical differences allow it to be recognized as a

separate species (Okamoto & Kamezaki, 2014).

However, we prefer to follow the more conser-

vative position most specialists adopt, consider-

ing this population as a more melanistic one of

the same nominal species (Bowen & Karl, 2000;

Rhodin et al., 2021).

We follow the suggestions of Fritz et al.

(2012), who propose the synonymy of Trache-

mys emolli with T. grayi, distributed in Costa

Rican Pacific lowlands. Two subspecies are

recognized: T. gray i emolli (in the Lake Nicara-

gua basin and adjacent lands in northern Costa

Rica) and T. grayi panamensis McCord, Joseph-

Ouni, Hagen, & Blanck, 2010 in the South

Pacific of Costa Rica (Rhodin et al., 2021). On

the other hand, the status of T. venusta is a sub-

ject of intense debate. However, we align with

Rhodin et al. (2021) suggestion, recognizing T.

venusta uhrigi (McCord, et al., 2010) from the

Caribbean lowlands of Costa Rica. In contrast,

Fritz et al. (2012) propose that populations of

these subspecies be recognized as T. ornata,

adding to the intrigue and complexity of the

taxonomic landscape.

Squamata. Sauria: Until recently, only one

species of the subfamily Mabuyinae (Scincidae)

was considered to occur in Costa Rica (Sav-

age, 2002). Hedges & Conn (2012) erected the

genus Marisora to include several skinks previ-

ously in Mabuya. These authors recognized

three species of Marisora present in Costa Rica

(Table 1), indicating features that distinguish

between them.

We accepted Alopoglossidae as the fam-

ily that includes Alopoglossus plicatus (Tay-

lor, 1949). This family was recently erected

(Goicoechea et al., 2016) to resolve a problem

of nomenclature and priority of names within

the family Gymnophthalmidae Fitzinger, 1826.

This move is based on previous work (Castoe

et al., 2004; Pellegrino et al., 2001; Pyron et al.,

2013) that showed that Alopoglossus and Ptycho-

glossus Boulenger, 1890, formed the sister clade

to the group Gymnophthalmidae + Teiidae.

Our placement of the microteiid species in

the subfamilies Cercosaurinae and Gymnoph-

thalminae (Table 1) follows Goicoechea et al.

(2016). Neusticurus apodemus Uzzell, 1966 was

transferred to the genus Potamites by Doan &

Castoe (2005). In contrast, Marques de Souza

et al. (2018) recovered P. apodemus as a sister

species to the clade containing members of

Echinosaura Boulenger 1890. Later, Vásquez-

Restrepo et al. (2020) found that Echinosaura

was not monophyletic, with E. apodema being

closer to members of the genus Andinosaura.

Consequently, these authors erected the genus

Centrosaura, with C. apodema as the type spe-

cies in this new combination. We have followed

that suggestion.

We also recognize the genus Loxopholis

Cope 1866 following Goicoechea et al. (2016),

who revalidated this genus to include several

species of microteiids previously recognized

in Leposoma. Loxopholis rugiceps is the type

species of that genus, first described from

the Colombian Choco. The species has been

reported from the Caribbean of Honduras and

Costa Rica (Mora et al., 2019).

On the other hand, Bachia pallidiceps has

been suggested for Costa Rica. However, given

the subtle differences between this species and

B. blairi (McDiarmid & DeWeese, 1977), we

wait to list its presence until more material

secures this position.

As for the teiids, we follow the conclu-

sions of Harvey et al. (2012), who found the

genus Ameiva sensu lato to be polyphyletic.

These authors resurrected the genus Holcosus

Cope, 1862 to resolve the polyphyly. Ugue-

to & Harvey (2011) recognized A. praesignis

from Isthmian Central America and northern

South America as distinct from A. ameiva.

Meza-Lázaro et al. (2015) elevated several of

the hitherto recognized subspecies of Holco-

sus undulatus (Wiegmann, 1834) to species

level, recognizing Holcosus parvus (Barbour &

Noble, 1915) as a form with a distribution from

the Isthmus of Tehuantepec along the Pacific

coast of Central America to northern Costa

Rica. Similarly, based on this work, Holcosus

pulcher (Hallowell, 1861), a species of the same

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64536, mayo 2025 (Publicado May. 15, 2025)

complex described from Nicaragua, has been

suggested for the northern Pacific of our coun-

try. However, we are refraining from listing it

until we verify the presence of two species in

Costa Rica. As for Cnemidophorus lemniscatus

(Linnaeus, 1758), it has been suggested as a

possible inhabitant of Costa Rica due to its wide

distribution in Central and South America and

the Caribbean (Savage, 2002). However, despite

extensive herpetological studies, no specimens

have been collected in this country, and hence,

it is not included in our list.

We recognize the family Anolidae as

including the diverse genus Anolis (Table 1),

following the suggestion of De Queiroz (2022),

who recently rejected using Dactyloidae Fitz-

inger, 1843. His suggestion is because Anolidae

Cocteau (1836) has priority. Anolis auratus

Daudin, 1802 is the type species of the genus

Norops Wagler, 1830. Some authors favor this

name for the clade containing the anolid spe-

cies reported here (Nicholson et al., 2012; Nich-

olson et al. 2018). However, we follow a more

conservative approach until the monophyly of

that genus is better supported (see Poe, 2013;

Poe et al., 2017). Anolis auratus from Isthmian

Central America is possibly a different species

from the South American populations.

Other considerations regarding ano-

lid lizards are as follows. Anolis quaggulus was

removed from the synonymy of A. humilis

(Köhler et al., 2006). However, we have not

been able to distinguish between both spe-

cies in the country. Therefore, we agree with

Poe (2016) that the diagnostic characters sug-

gested by Köhler are insufficient to distinguish

between them. Anolis laeviventris (Wiegmann,

1834) was initially described from Mexico.

Until recently (Sasa et al., 2010; Savage, 2002),

populations of this species in Costa Rica were

assigned to A. intermedius Peters, 1863, now

considered a junior synonym of the nominal

species. However, given the disjunct distribu-

tion of A. laeviventris at intermediate elevations

from Mexico to Costa Rica, a more thorough

taxonomic revision is required to determine the

species status in Costa Rica. We consider Anolis

salvini Boulenger, 1885, to have priority over

A. vociferans (Myers, 1971). Köhler & Vesely

(2010) reviewed the polytypic species A. seri-

ceus Hallowell, 1856 and concluded that it is a

species complex, recognizing A. unilobatus as

the species in Costa Rica. We have chosen to

follow their suggestion.

We recognize seven anguid lizards (Angui-

dae + Diploglossidae) in the country (Table

1). Gerrhonotus rhombifer Peters 1876 was the

type species of the genus Coloptychon Tihen,

1949. However, García-Vázquez et al. (2018)

found Coloptychon to be nested within Ger-

rhonotus, thus validating the original descrip-

tion. According to Schools & Hedges (2021),

the genera Celestus Gray, 1839 and Diploglossus

Wiegmann, 1834 used until recently to clas-

sify diploglossids in Central America (Savage,

2002) are polyphyletic. Therefore, they propose

a taxonomic reorganization that includes estab-

lishing the genus Mesoamericus and assigning

the Central American species previously in

Celestus to the genus Siderolamprus Cope, 1861.

Squamata. Serpentes: Some modifica-

tions to the nomenclature of snakes previously

reported for the country (Solorzano, 2022)

are the following. Albuquerque & Fernandes

(2022) elevated several of the recognized sub-

species of the nominate species Leptophis

ahaetulla (Linnaeus, 1758) to species status.

Leptophis occidentalis is the species that inhab-

its Costa Rica. Mastigodryas alternatus has been

reported as a species possibly present in Costa

Rica (Uetz et al., 2023). In the past, this species

was synonymized with M. melanolomus (Peters

& Orejas-Miranda, 1970), and some authors

point out that no substantive morphological

differences allow the separation of these spe-

cies (Solorzano, 2022). Therefore, we adopt the

more conservative position of recognizing only

M. melanolomus in the country.

Among the Dipsadidae family, the taxon-

omy of the cat-eye snake genus Leptodeira that

inhabits Costa Rica has generated some contro-

versy (Barrio-Amorós, 2019). We kept a con-

servative stance when recognizing Leptodeira

septentrionalis (Kennicott, 1859). This species

was first described from northeastern Mexico

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64536, mayo 2025 (Publicado May. 15, 2025)

and is known for exhibiting extensive morpho-

logical variation throughout its distribution in

North, Central, and South America. In his pio-

neering revision of the genus, Duellman (1958)

describes Leptodeira septentrionalis ornata Boc-

court 1884 to account for this variation partial-

ly. According to this author, “this subspecies is

naturally divided into three populations: one in

lower Central America, one on the Pacific coast

of South America, and one in the inter-Andean

valleys in Colombia.” Costa et al. (2022) showed

the polyphyly of L. septentrionalis, indicating

that the populations of L. s. ornata from Costa

Rica and northern Panama possibly represent

a taxon yet to be described. Solorzano (2022)

agrees with this treatment, avoiding assigning

these populations to L. septentrionalis.

Some considerations regarding the nomen-

clature of the Elapidae snakes are presented

here. Jowers et al. (2023) found that the nomi-

nal species Micrurus nigrocinctus is a species

complex that originated in the Pliocene and is

composed of at least three distinct lineages at

the species level. M. n. nigrocinctus, the taxon

recognized in Costa Rica and much of Central

America (Savage, 2002; Solórzano, 2022), is dis-

tinct from the clade that includes the popula-

tions of central and southern Panama, near the

type locality of the species. Given the potential

public health implications, Jowers et al. (2023)

refrained from taxonomically recognizing this

divergence pending further information. Con-

sequently, we have decided to continue recog-

nizing M. nigrocinctus as the species of coral

snake present in the Central Plateau and the

North and Central Pacific of the country.

Solórzano (2022) recognizes Micrurus yate-

si as a separate species from other Costa Rican

coral snakes. This form was initially described

as a subspecies of M. nigrocinctus (M. nigro-

cinctus yatesi) by Dunn (1942) from specimens

collected in Puerto Armuelles, Panama, close

to the southern border of Costa Rica. Savage

& Vial (1974) recognize the taxon as distinc-

tive but include it within the nominal species

M. alleni Schmidt, 1933 (i.e., M. alleni yatesi).

Several researchers have validated the distinc-

tiveness of this taxon (Campbell & Lamar, 2004;

Mena et al., 2022), although a formal analysis of

its taxonomy is still pending.

Likewise, Solórzano (2022) also recognized

the presence of M. mipartitus in the country and

distinguished it from M. multifasciatus based

mainly on differences in coloration patterns.

We follow this author here, although the status

of both species should be reviewed consider-

ing their evolutionary relationships throughout

their distribution since several authors con-

sider both forms to be the same species (Savage,

2002) or that only one species occurs in the

country (Campbell & Lamar, 2004).

The extensive variation in coloration, mor-

phological and molecular characters exhibited

by the widely distributed eyelash palm pitvi-

per, Bothriechis schlegelii (Berthold, 1846), has

raised suspicions that it is a species complex

(Campbell & Lamar, 2004; Taggart et al., 2001).

Daza et al. (2010) and Townsend et al. (2013)

showed that at least two lineages of the nominal

species are recognizable at the molecular level,

one distributed throughout Mesoamerica and

the other in South America. In their recent sys-

tematic review, Arteaga et al. (2024) identified

several lineages within the complex and recog-

nized them as valid species. Although several

of the proposed species in South America has

been questioned (Reyes-Velasco, 2024), Both-

riechis nigroadspersus, first described by Austri-

an zoologist Franz Steindachner in 1870 as the

species inhabiting Mesoamerica, is considered

valid. We favor this name over the nomencla-

ture used until recently for the species in Costa

Rica (Solórzano, 2022).

Historical account of the herpetological

collection at MZUCR

In the early 1960s, an essential academic

relationship was established between Dr. Jay

M. Savage, then a professor at the University

of California, and Dr. Rafael Lucas Rodríguez

Caballero, the first director of the then Depart-

ment of Biology at the University of Costa

Rica (UCR) (García, 2009). This relationship

encouraged the participation of professors from

United States universities in the Department

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64536, mayo 2025 (Publicado May. 15, 2025)

activities and provided logistical support of

this unit for their field courses, mainly directed

to their graduate students. The integration

of these efforts by national and foreign pro-

fessors constituted the origin of the Orga-

nization for Tropical Studies (OTS; https://

tropicalstudies.org/), currently a consortium

of universities that offer courses and facilitates

research and exploration of tropical ecosystems

(Stone et al., 1988).

During those early years, the link between

Savage and the UCR brought several zoologists

as faculty members. They made small collec-

tions initially maintained in their own offices.

The first of the herpetologists, Dr. James Vial,

served as a professor in the Department in

1963 (García, 2009). Vial focused his interest on

amphibians, particularly salamanders, and ini-

tiated a personal collection of his study subjects

(Vial, 1966; Vial, 1967). Nearly 90 specimens

collected by Vial form the basis of the current

amphibian collection at the Museum. Dr. Nor-

man Scott succeeded Vial from 1964 to 1966

(García, 2009), who studied the distribution of

litter herpetofauna and the factors determining

their abundance and extended his interests to

reptiles (Scott, 1976). Scott collected broad-

ly while attending OTS field courses, adding

about 360 specimens to the collection started

by Vial. The emerging collection of amphibians

and reptiles was significantly expanded thanks

to the efforts of these dedicated individuals and

contributions from collectors such as Carlos

Valerio and Jim Brockett.

In 1966, Dr. Douglas C. Robinson joined

the Department, succeeding Scott. Robinson’s

academic legacy at UCR is indelible and his

role in developing herpetology as a formal

discipline in Costa Rica was a significant mile-

stone. The same year of his incorporation, the

herpetology and ichthyology collections were

moved to a small space assigned by Dr. Jorge

Mora, then director of the Department (García,

2009). This action is considered the begin-

ning of the Museum of Zoology in said unit

(Springer, 2019).

Robinson served as a professor for the next

twenty-five years. In 1974, the Department

was transformed into the School of Biology,

and he was actively involved in developing

the new academic curriculum, teaching vari-

ous specialized courses, including herpetology,

amphibian biology, reptile biology, comparative

anatomy, and sea turtle biology, a subject that

was particularly close to his heart. He focuses

on improving the museum’s conditions, par-

ticularly the herpetology collection, and hiring

support staff such as Roger Saenz and Federico

Valverde (Sasa et al., 2022). By 1976, the collec-

tions had been moved to a larger space in the

basement of the Biology School building, where

they remain today (Angulo et al., 2016; García,

2009). The improvement in space allowed the

inclusion of new specimens. More than 3 580

specimens collected by Robinson or his stu-

dents were incorporated into the collection,

and many more were added by other collectors,

including Sergio Salas, Peter Siegfried, Roy W.

McDiarmid, Luis Diego Gómez, Michael M.

Johanboecke, and Wayne van Devender. At the

time of Robinson’s death in 1991, 12 500 speci-

mens were exceeded (Fig. 3), a testament to his

impressive efforts to ensure the collection.

Robinson’s most significant contribution

was to train the first generation of Costa Rican

herpetologists. Like no other of the scientists

who preceded him, Robinson had the vision

and patience to consolidate interest in herpe-

tology among his students (Chaves, 2018), a

transcendental fact for the local development

of studies on amphibians and reptiles in the

country and for the development of the collec-

tion that he helped found at MZUCR.

Over the next thirty years, the collection of

amphibians and reptiles experienced a signifi-

cant expansion, doubling in size. This growth

was not a solitary effort, but a collaborative

achievement, thanks to the dedication of the

curatorial team Federico Bolaños and Gerardo

Chaves, and the contributions of their students

and associates, including Adrián García, Ale-

jandro Zamora, José Hernández, and Roberto

Brenes. The Bolaños-Chaves duo played a key

role, adding more than 3,000 specimens to

the collection and accelerating its growth by

facilitating the deposit of material collected

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64536, mayo 2025 (Publicado May. 15, 2025)

by other researchers, including Jens Ehmckey,

Gad Perry, Mason Ryan, Mahmood Sasa, and

Alejandro Solórzano. A significant milestone

during this period was the assignment of geo-

graphic coordinates to all collection localities, a

task that was initiated by Douglas C. Robinson.

By 2000, the herpetological collection con-

tained 67% of the total number cataloged today

(Fig. 3). During the first years of this cen-

tury, the digitalization of specimen information

was completed, and tools from Geographic

Information Systems were incorporated as an

essential resource for analyzing their distribu-

tions. The invaluable contributions of several

researchers, including Adrián García Rodríguez

and Robert Puschendorf, who brought their

unique experience to the project, truly made

this achievement possible. García-Rodríguez

first incorporated spatial analysis tools into the

locality records and used modeling to resolve

biogeographical questions about regional her-

petofauna, while Puschendorf used these tech-

niques to understand the ecology of the chytrid

fungus associated with the decline of amphib-

ians in the country (García-Rodríguez et al.,

2012; Puschendorf et al., 2006). From this point

on, once the information was digitalized, it was

possible to map and model the distribution of

most of the country’s amphibian and reptile

species. These achievements have allowed us

to advance in the knowledge about our herpe-

tofauna and establish a new standard in data

management for the scientific community. As

we look to the future, these advancements open

exciting possibilities for further research, as

evidenced by studies related to niche model-

ing (Granados-Martínez et al., 2021), potential

distribution (Solórzano & Sasa, 2020), and

the progress of pathogens responsible for the

amphibian declines (Puschendorf et al., 2009;

Zumbado-Ulate et al., 2019) that have resulted

from them.

From 2000 to 2020, the number of speci-

mens preserved at the MZUCR increased by

more than 25% (Fig. 3), significantly improv-

ing our research capabilities. This increase

was made possible by the dedicated efforts of

researchers such as David Laurencio, Erick

Arias, Elda Araya, Valerie McKenzie, Eduardo

Boza, Franklin Aguilar, and Robert Lovich,

among others. They incorporated specimens

from environmental studies and conservation

projects (Arias & Bolaños, 2014; Laurencio

& Malone, 2009) or parasitic communities

(McKenzie, 2007), thus expanding the scope

of the collection. By 2020, more than 23,600

specimens had already been cataloged in the

collection. During the first four years of this

last decade, some 400 more specimens have

been added, mainly salamanders and anurans

collected in expeditions carried out to the

Talamanca Mountain range, a region known for

its rich biodiversity and unique herpetofauna

(Arias et al., 2024).

Fig. 3. Number of specimens catalogued per year in the MZUCR amphibian and reptile collection.

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64536, mayo 2025 (Publicado May. 15, 2025)

As of 2020, the Museum of Zoology is no

longer managed by the School of Biology. It

has become part of the Center for Research in

Biodiversity and Tropical Ecology, a unit of the

Vice-Rectorate of Research of the University of

Costa Rica dedicated to research on the biodi-

versity and ecology of terrestrial and freshwater

environments. This measure is hoped to allow

for greater visibility and projection of the work

carried out in the herpetology collection.

Despite the extensive history of herpe-

tological studies, new species continue to be

added to the list of amphibians and reptiles of

Costa Rica. Recent taxonomic changes, many

of them derived from systematic studies that

incorporate new evidence of their evolutionary

relationships and the discovery of new species

from the least studied regions of the country,

guarantee that the number and identity of

species will continue to change in the coming

years. In the face of these changes, the herpe-

tology collection of the Museum of Zoology

constitutes a significant asset since it protects

voucher specimens that represent a very high

proportion of the species in the country. In

addition, the collection is a research unit that

directly manages many of the studies on the

biology of amphibians and reptiles that the

university carries out. The history of this col-

lection, a testament to the collaborative spirit of

the field, reflects the contributions of those who

helped found it, their interactions with other

researchers and students, and the academic col-

laborations that have shaped it over time.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

The authors would like to thank two

anonymous reviewers for their suggestions to

improve the manuscript. This work was par-

tially funded by funds from project C4752

Vicerrectoría de Investigación, Universidad de

Costa Rica.

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