Amphibians and reptiles of the Calakmul Biosphere Reserve, México, with new records

Abstract

We provide a list of amphibians and reptiles of the Calakmul Biosphere Reserve in the southern half of the Mexican Yucatan, in the state of Campeche. The study area was sampled through opportunistic, transect and pitfall trap surveys conducted for three successive years. These surveys resulted in a total of 2,359 amphibian and reptile encounters, belonging to 20 amphibian and 69 reptile species from 24 total families. We present herein the records for one snake, one chelonian and two salamander species not previously recorded in the area.

Full text

Check List | www.biotaxa.org/cl Volume 11 | Number 5 | Article 1759 1
Check List the journal of
biodiversity data
Amphibians and reptiles of the Calakmul Biosphere Reserve,
México, with new records
Timothy J. Colston1, 2*, José António L. Barão-Nóbrega2, Ryan Manders2, Alice Lett2, Jamie Willmott2,
Gavin Cameron2, Sidony Hunter2, Adam Radage2, Etienne Littlefair2, Robert J. Williams2,
Antonio Lopez Cen3 and Kathy Slater2
1 University of Mississippi, Biology Department, University, MS, USA
2 Operation Wallacea, Wallace House, Old Bolingbroke, Lincolnshire, PE23 4EX, UK
3 Pronatura Peninsula de Yucatan A. C., Calle 32 # 269 x 47 y 47a Col. Pinzón II. C. P. 97205. Mérida Yucatán, Mexico
* Corresponding author. E-mail: tim@maddreptiles.com
Abstract: We provide a list of amphibians and reptiles
of the Calakmul Biosphere Reserve in the southern half
of the Mexican Yucatan, in the state of Campeche. e
study area was sampled through opportunistic, transect
and pitfall trap surveys conducted for three successive
years. ese surveys resulted in a total of , amphibian
and reptile encounters, belonging to  amphibian and
 reptile species from  total families. We present
herein the records for one snake, one chelonian and two
salamander species not previously recorded in the area.
Key words: Herpetofauna, Campeche, Yucatán
INTRODUCTION
Mexico is one of the most biologically rich countries
on the planet (Mittermeier and Goettsch-Mittermeier
; Gárcia-Frapolli et al. ). is is due in no small
part to the diversity found in the southern region of
the Yucatan Peninsula, which is home to the largest
expanse of mature, seasonal tropical forests remaining
in Mesoamerica (Carr ; Vester et al. ). Unlike
the majority of forest in the Yucatan Peninsula, the
forest in Calakmul Biosphere Reserve, hereafter CBR,
of Campeche, Mexico has not been used for large scale
timber production nor has it been burned for farming
and ranching. As such it is one of the last remaining
stands of virgin forest in Mexico, being classied by
UNESCO as World Heritage Site of Culture and Nature
(UNESCO ). In spite of the recognized cultural
and ecological value of CBR, few surveys have been
conducted on its herpetofauna.
Beginning in , annual biodiversity surveys have
been conducted conducted in the CBR by Operation
Wallacea, a UK-based non-governmental organization
that specializes in biodiversity assessments and
monitoring of protected areas through utilizing
the expertise of university academics and students.
We present herein the results of the herpetological
(amphibian and reptile) surveys and discuss their
conservation implications for the CBR.
MATERIAL AND METHODS
Our study was conducted in the CBR (.ʹ
N, .ʹ W; Figure ), located in the Yucatan
Peninsula in Campeche, México. CBR is a large expanse
of tropical forest (, ha of reserve and , ha
of buer zone) that is part of the Selva Maya (Mayan
Check List 11(5): 1759, 4 October 2015 doi: http://dx.doi.org/10.15560/11.5.1759
ISSN 1809-127X © 2015 Check List and Authors
5
11
1759
4 October 2015
Figure 1. Location of Calakmul Biosphere Reserve within the southern
Yucatán Peninsula. Study sites within the reserve are also indicated. KM27
and KM40 are not shown due to their close proximity to KM20 and Zona
Arqueologica respectively. Green shading on inset represents the pro-
tected reserve (core and buer zones) and orange shading indicates the
connected state reserves of Balam-kim and Balam-ku.
Lists of species

Check List | www.biotaxa.org/cl Volume 11 | Number 5 | Article 1759 2
Colston et al. | Herpetofauna of the Calakmul Biosphere Reserve
owing water in the form of a stream rather than semi-
permanent aguadas (temporary pond). KM served
as the primary base camp and operation headquarters
due to the park infrastructure already in place and was
located in the transition zone from medium to tall
canopy forest. KM and KM contained the largest and
most numerous aguadas among our camps; while Dos
Naciones, the southernmost camp, was characterized by
the largest and most numerous fruiting trees.
Semi-permanent camps were constructed and utilized
at ve of the sites (Dos Naciones, KM, Mancolona,
Nadzca’an and Hormiguero). Camp locations were
chosen for their accessibility during the wet season
and because they cover the full geographical and
vegetation range of the reserve. Each camp contained
four, two km long transect lines for data collection.
Transect surveys were conduct both in the day and at
night and all surveys were replicated a minimum of
four times. Transect surveys were alternated such that
each transect had a minimum of  hours rest between
surveys. All animals encountered on transect surveys
were immediately captured, identied, photographed,
when possible marked to avoid repeating counts and
identify recaptures, measured and then released at the
point of capture. Common, easily identiable species
Jungle) which encompasses parts of Mexico, Guatemala
and Belize, spans over . million ha and is the largest
continuous section of tropical forest in Mesoamerica
(Vester et al. ). ere is a notable precipitation
gradient in the reserve, from  mm annually in the
north to , mm annually in the south of the reserve;
this has signicant eect on forest structure and tree
species composition (Vester et al. ). e majority
of the reserve is composed of tropical semi-deciduous
forest with a canopy ranging from  to  m, whereas
the northern parts of the reserve are composed of
tropical deciduous forest, with canopy of  to  m
(Chowdhury ).
Data collection was carried out between June and
August of – and included surveys in seven
dierent sampling localities (Dos Naciones, KM,
KM, KM, Mancolona, Nadzca’an, and Hormiguero)
within the CBR (Figure ). Mancolona and Nadzca’an
are located in the dry north of the reserve, and are
proximal to buer zones for the reserve. Both of these
northern camps have few of the typical large fruiting
trees (i.e., ramon (Brosimium alicastrum) and zapote
(Manilkara zapota)) relative the core and southern areas
of the reserve. Although situated in the relatively dry
north, Nadzca’an was the only site with permanent
Figure 2. (A) Example of pitfall trap and drift fence array. (B) Typical semi-permanent aguada. (C) Graphical representation of transect and trap design
where camp is indicated by yellow triangle, transects by dashed lines, aguadas by blue circles and pitfall and drift fence arrays by Xs.

Check List | www.biotaxa.org/cl Volume 11 | Number 5 | Article 1759 3
Colston et al. | Herpetofauna of the Calakmul Biosphere Reserve
(e.g., Ameiva undulata, Incillius valliceps) were simply
counted after a minimum of ten individuals had been
measured during a given survey.
At each camp two pitfall trap arrays composed of four
buckets each disposed in a Y shape connected by  m
long by  m high drift fences were constructed. In each
location, one trap array was placed adjacent to a transect
near an aguada, and one adjacent to a transect at least
one km from an aguada (Figure ). Traps were checked
each morning and were left open for a minimum
of three weeks. All animals captured in traps were
immediately removed, identied, photographed, when
possible marked to avoid repeating counts and identify
recaptures and measured prior to release a safe distance
away from the traps while still in the immediate area.
All data were collected by teams of students led by
university academics and local indigenous experts. Due
to permit restrictions within the Biosphere reserve no
specimens were collected but digital photographs were
taken and represent digital vouchers for all but two
species. ese digital vouchers are curated by Operation
Wallacea, TJC and Pronatura Peninsula de Yucatan (PPY).
Tissue samples in the form of scale, toe or tail clips that
resulted from marking individuals were retained and
deposited at the tissue collection of the lead author at
the University of Mississippi. Additionally, collections of
tissues as a by-product of marking individuals followed
the University of Mississippi Institute for Animal Use
and Care protocols: SOP - and SOP -.
RESULTS
We recorded a total of , encounters during our
surveys, representing  amphibian species included in
seven families (Table ) and  reptile species from 
families (Table ). All  and  species of amphibians and
reptiles, respectively, previously described for CBR were
observed (Lee ; Kholer ; Mandujuano et al.
; Cedeño-Vasquéz et al. ). e most species-rich
Table 1. List of amphibians encountered in the CBR. *New records not previously reported for the area. **Digital voucher only.
Family Species Field ID Locality
Bufonidae Incilius valliceps TJC618 KM20, KM40, Dos Naciones, Hormiguero, Mancolona, Nadzca’an
Bufonidae Rhinella marina TJC733 KM20, KM40, Dos Naciones, Hormiguero, Mancolona, Nadzca’an
Hylidae Agalychnis callidryas TJC935 KM20, Dos Naciones, Hormiguero, Mancolona
Hylidae Dendropsophus ebraccatus TJC1042 KM20, Dos Naciones
Hylidae Hyla microcephala TJC1197 KM20, Hormiguero, Mancolona
Hylidae Hyla picta TJC1202 KM20
Hylidae Scinax staueri TJC605 KM20, Hormiguero, Mancolona
Hylidae Smilisca baudinii TJC649 KM20, Dos Naciones, Hormiguero, Mancolona, Nadzca’an
Hylidae Tlalocohyla loquax TJC693 KM20, KM40, Dos Naciones, Hormiguero, Mancolona
Hylidae Trachycephalus venulosus TJC710 KM20, Hormiguero, Mancolona
Leptodactylidae Leptodactylus fragilis TJC950 KM20, KM40, Mancolona
Leptodactylidae Leptodactylus melanonotus TJC951 KM20
Microhylidae Gastrophryne elegans TJC995 KM20
Microhylidae Hypopachus variolosus TJC626 KM20, KM40, Hormiguero, Mancolona, Nadzca’an
Ranidae Lithobates berlandieri TJC720 KM20, KM40, Dos Naciones, Hormiguero, Mancolona
Ranidae Lithobates vaillanti TJC607 KM20, Hormiguero
Rhinophrynidae Rhinophrynus dorsalis TJC749 KM20, Dos Naciones, Hormiguero
Plethodontidae Bolitoglossa mexicana* TJC1041 KM20, Dos Naciones
Plethodontidae Bolitoglossa rufescens ** KM20
Plethodontidae Bolitoglossa yucatana ** KM20, Dos Naciones
Table 2. List of reptiles encountered and localities in the CBR. *New records not previously reported for the area.
Family Species Field ID Locality
Crocodylidae Crocodylus moreletii ** KM27, Hormiguero, Mancolona
Boidae Boa constrictor TJC928 KM20, Nadzca’an
Colubridae Dryadophis melanolomus TJC971 KM27, KM40, Dos Naciones, Mancolona, Nadzca’an
Colubridae Drymarchon corais TJC843 Hormiguero, Mancolona
Colubridae Drymobius margaritiferus TJC617 KM20, KM27, KM40, Hormiguero, Mancolona, Nadzca’an
Colubridae Elaphe avirufa TJC698 KM20, KM27, KM40, Hormiguero
Colubridae Ficimia publia TJC780 KM20, Hormiguero
Colubridae Lampropeltis triangulum TJC786 KM20, Hormiguero
Colubridae Leptophis ahaetulla ** KM20, KM40, Dos Naciones
Colubridae Leptophis mexicanus TJC647 KM20, KM27, KM40, Hormiguero, Mancolona
Colubridae Mastigodryas melanolomus ** KM20
Colubridae Oxybelis aeneus TJC1032 KM20, Nadzca’an
Colubridae Oxybelis fulgidus ** Mancolona
Continued

Check List | www.biotaxa.org/cl Volume 11 | Number 5 | Article 1759 4
Colston et al. | Herpetofauna of the Calakmul Biosphere Reserve
Family Species Field ID Locality
Colubridae Pseustes poecilonotus TJC766 KM20
Colubridae Senticolis triaspis ** Mancolona
Colubridae Spilotes pullatus ** KM27, KM40, Hormiguero
Colubridae Symphimus mayae ** KM20, Dos Naciones
Colubridae Xenodon rabdochephalus TJC992 KM20, Mancolona
Corytophanidae Basiliscus vittatus TJC816 KM20, KM27, KM40, Hormiguero, Mancolona
Corytophanidae Corytophanes cristatus TJC707 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona
Corytophanidae Corytophanes hernandezii TJC811 Dos Naciones, Hormiguero
Corytophanidae Laemanctus serratus TJC659 KM20, Hormiguero, Nadzca’an
Dactyloidae Norops biporcatus TJC1191 Dos Naciones, Hormiguero
Dactyloidae Norops cristellatus ** KM20
Dactyloidae Norops lemurinus TJC1033 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona,
Nadzca’an
Dactyloidae Norops pentaprion ** KM20
Dactyloidae Norops rodriguezii TJC1043 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona,
Nadzca’an
Dactyloidae Norops sagrei ** KM20, Dos Naciones, Mancolona
Dactyloidae Norops sericeus TJC638 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona
Dactyloidae Norops tropidonotus TJC662 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona
Dipsadidae Coniophanes imperialis TJC616 KM20, KM27, KM40, Hormiguero, Mancolona, Nadzca’an
Dipsadidae Coniophanes schmidti TJC725 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona,
Nadzca’an
Dipsadidae Dipsas brevifacies TJC674 KM20, KM27, KM40, Hormiguero, Mancolona
Dipsadidae Imantodes cenchoa TJC954 KM20, Dos Naciones, Hormiguero, Mancolona
Dipsadidae Imantodes gemnistratus TJC836 KM20, Hormiguero
Dipsadidae Imantodes tenuissimus TJCA13 KM20, KM27, KM40
Dipsadidae Leptodeira frenata TJC643 KM20, Hormiguero, Mancolona, Nadzca’an
Dipsadidae Leptodeira septentrionalis TJC1211 Dos Naciones
Dipsadidae Ninia diademata* TJC1208 KM20, KM40
Dipsadidae Ninia sebae TJC827 KM20, Hormiguero
Dipsadidae Sibon sanniola TJCA35 KM20
Dipsadidae Tropidodipsas fasciata TJC666 Hormiguero
Dipsadidae Tropidodipsas nebulata ** KM20
Dipsadidae Tropidodipsas sartorii TJC682 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona,
Nadzca’an
Elapidae Micrurus diastema TJC753 KM20, KM27, KM40, Mancolona, Nadzca’an
Gekkonidae Coleonyx elegans TJC640 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona,
Nadzca’an
Gekkonidae Hemidactylus frenatus ** KM20, Dos Naciones, Hormiguero, Mancolona
Gekkonidae Thecadactylus rapicauda TJC632 KM20, Dos Naciones, Hormiguero, Mancolona
Iguanidae Ctenosaura defensor ** Hormiguero, Nadzca’an
Phrynosomatidae Sceloporus chrysostictus TJC642 KM20, Dos Naciones, Hormiguero, Mancolona, Nadzca’na
Phrynosomatidae Sceloporus lundelli ** Mancolona
Scincidae Mesoscincus schwartzei TJC987 KM20, Mancolona
Scincidae Plestiodon sumichrasti ** KM20
Scincidae Sphenomorphus cherriei TJC1207 KM20
Sphaerodactylidae Sphaerodactylus glaucus TJC927 KM20, Hormiguero, Mancolona
Teiidae Ameiva undulata TJC672 KM20, KM27, KM40, Dos Naciones, Hormiguero, Mancolona,
Nadzca’an
Teiidae Aspidoscelis angusticeps TJC609 KM20, KM27, KM40, Hormiguero, Mancolona,
Viperidae Agkistrodon bilineatus ** KM20, Nadzca’an
Viperidae Bothrops asper TJC791 KM20, KM27, KM40, Dos Naciones, Nadzca’an
Viperidae Crotalus simus TJC823 KM20, Dos Naciones, Hormiguero, Nadzca’an
Emydidae Terrapene carolina TJC754 Hormiguero
Emydidae Trachemys scripta ** KM20, KM27, Hormiguero
Geoemydidae Rhinoclemmys areolata TJC840 KM20, Hormiguero
Kinosternidae Claudius angustatus ** KM20, Hormiguero, Nadzca’an
Kinosternidae Kinosternon acutum* ** Hormiguero
Kinosternidae Kinosternon creaseri ** KM20, Nadzca’an
Kinosternidae Kinosternon leucostomum ** KM20, Dos Naciones, Hormiguero, Nadzca’an
Kinosternidae Kinosternon scorpioides TJC833 KM20, KM27, Hormiguero, Mancolona, Nadzca’an
Table 2. Continued.

Check List | www.biotaxa.org/cl Volume 11 | Number 5 | Article 1759 5
Colston et al. | Herpetofauna of the Calakmul Biosphere Reserve
amphibian family was Hylidae ( of amphibian species
richness), while the most species-rich reptile families
were Colubridae, Dipsadidae and Dactyloidae (, 
and  of reptile species richness, re spec tively).
e snake Ninia diademata; Figure , the turtle
Kinosternon acutum; Figure  and two salamanders
(Bolitoglossa mexicana, Bolitoglossa rufescens; Figure )
were recorded during the study. None of these were
previously reported for the CBR although all have been
recorded  km to the south in Guatemala and to the
west in the Mexican state of Chiapas (Lee ).
DISCUSSION
In our study, the total number of species found in the
CBR represents about  of the herpetofaunal species
richness observed in the Mayan jungle ( species; Lee
). Of this number, nearly  of species occur only
within the Yucatan Peninsula (e.g., Bolitoglossa yucatana;
Coleonyx elegans). Knowledge of the assemblage and
distribution of amphibians and reptiles in the Yucatan
Peninsula is, therefore, of great importance when taking
measures to preserve the fauna of this region, especially
because site conservation may be one of the most eective
means of reducing biodiversity loss (Eken et al. ).
Our results point out that only about  of the
recorded species were recorded in three or more camps,
suggesting signicant dierences on species richness
and diversity within Calakmul, and therefore highlight
the need for specic conservation measures that are
adapted to each habitat type. Latitude-correlated rain-
fall dierences among camps may be responsible for
dierences in local microhabitat dierences (Stevens
; Lyons and Willig ), which may result in species
Figure 6. Lizards encountered in the Calakmul Biosphere Reserve: (A)
Basiliscus vittatus, (B) Laemanctus serratus, (C) Sphenomorphus cherriei, (D)
Corytophanes hernandezii , (E) Corytophanes cristatus. Photos by TJC.
Figure 5. Amphibians encountered in the Calakmul Biosphere Reserve:
(A) Bolitoglossa mexicana, Gastrophryne elegans, (C) Leptodactylus fragilis.
Photos by TJC.
Figure 3. Snakes encountered in the Calakmul Biosphere Reserve: (A)
Ninia diadema, (B) Ninia sebae, (C) Coniophanes imperalis, (D) Ficimia pub-
lia. Photos by TJC.
Figure 4. Turtles encountered in the Calakmul Biosphere Reserve: (A)
Kinosternon leucostomum; adult and juvenile, (B) Claudius angustatus, (C)
Rhinoclemmys areolata, (D) Kinosternon acutum (Photo by JAB). Photos by
TJC.

Check List | www.biotaxa.org/cl Volume 11 | Number 5 | Article 1759 6
Colston et al. | Herpetofauna of the Calakmul Biosphere Reserve
variation among areas (Vitt et al. ; Garda et al. ).
However, when considering family level variation (e.g.,
Colubridae, Hylidae) and, to some extent, genus level
variation (e.g., Norops, Kinosternon), richness among our
camps was even.
Overall species abundance and diversity appeared to
follow the known precipitation gradient in the reserve.
Camps in the wetter, central and southern portions of the
CBR contained more herpetofaunal species as expected.
Interestingly when only considering reptile species
diversity the dry northern camp of Nadzca’an had a
similar number of reptile species as the southern camps.
We suspect this is due to the permanent stream located
in the camp which provides suitable habitat to a variety
of turtles as well as a water source for other reptiles
while not providing suitable breeding habitat for many
amphibian species that reproduce in temporary ponds.
Species which favour open or disturbed habitat
(e.g., Sceloporus chrysostictus) were more abundant
at the dry (open canopy) northern camps that are
also situated along the edge of the buer zone of the
reserve. Although some frogs were found in or near
human-made structures (e.g. Incillius valliceps) or small
water reservoirs (e.g., Smilisca baudinii), the majority
of individuals were found near aguadas. On the other
hand, some species were recorded away from major
water sources (e.g., Triprion petatsatus) in seemingly dry
habitat. Future studies will specically test the eect of
forest structure, habitat disturbance and distance from
aguadas on herpetofaunal community composition.
Guides to species of amphibians (Cedeño-Vasquez et
al. ) and reptiles (Calderon-Mandujano et al. )
of the CBR have already been produced. However, these
guides are not indexed and are only available in Spanish.
ereby, our study is the rst published indexed species
list for the region, and includes four new species that
were not previously reported from CBR, and signicant
range extensions for these species. Future studies
should be conducted in other seasons and considerable
eorts should be directed toward fossorial amphibians
and reptiles, which are hard to detect and may have not
been encountered previously.
ACKNOWLEDGEMENTS
We would like to thank the communities of Calakmul,
Dos Naciones, , Mancolona, Nadzca’an, and Hormiguero
for granting us the opportunity to work in their ajidos
as well as for providing guide services and logistical
support. We especially think Don Oliverio Platas
Vargas, for without his assistance none of these studies
would have been possible. We also thank the rest of
the Operation Wallacea sta and student volunteers
for logistical assistance and data collection. We thank
SEMARNAT and the National Commission of Natural
Protected Areas for providing permits. Permits issued
by SEMANART and National Commission of Natural
Protected Areas over the period of study as part of
a long-term conservation project with Pronatura
Peninsula de Yucatan were: SPGA/DGVS// and
SPGA/DGVS//. TJC would like to thank the
University of Mississippi Graduate College for funding
to support summer research and Lenin Arias Rodriguez
for assistance with export permits. We thank the Editor
and one anonymous reviewer for their helpful comments
on the manuscript.
LITERATURE CITED
Calderón-Mandujuano, R.R., C. Pozo de la Tijera and J.R. Cedeño-
Vasquez. . Guia Rustica de los Reptiles de la Region de
Calakmul, Campeche. Quintana Roo: ECOSUR, CONABIO.  pp.
Carr III, A. . Monitoring in the Selva Maya; pp. –, in: A.
Carr III and A.C.D. Stoll (eds.) Biological monitoring in the
Selva Maya. Gainsville: US Man and the Biosphere and
Figure 7. Amphibians encountered in the Calakmul Biosphere Reserve:
(A) Triprion petastatus, (B) Smilisca baudinii, (C) Rhinophrynus dorsalis, (D)
Dendropsophis ebracatta. Photos by TJC.
Figure 8. Snakes encountered in the Calakmul Biosphere Reserve: (A)
Xenodon rhabdocephalis, (B) Bothrops asper, (C) Leptodeira frenata, (D)
Leptodeira septentrionalis. Photos by TJC.

Check List | www.biotaxa.org/cl Volume 11 | Number 5 | Article 1759 7
Colston et al. | Herpetofauna of the Calakmul Biosphere Reserve
Wildlife Conservation Society.
Cedeño-Vasquez, J.R., R.R. Calderón-Mandujuano and C. Pozo
de la Tijera. . Guia Rustica de los Anbios de la Region de
Calakmul, Campeche. Quintana Roo: ECOSUR, CONABIO. pp.
Chowdhury, P.K. . Landscape change in the Calakmul Biosphere
Reserve, México: modeling the driving forces of smallholder
deforestation in land parcels. Applied Geography (): –.
doi: ./j.apgeog...
Gárcia-Frapolli, E., G. Ramos-Fernandéz, E. Galicia and A. Serrano.
. e complex reality of biodiversity conservation through
Natural Protected Area policy: three cases from the Yucatan
Peninsula, México. Land Use Policy (): –. doi: ./j.
landusepol...
Garda, A.A., H.C. Wiederhecker, A.M. Gainsbury, G.C. Costa,
R.A. Pyron, G.H.C. Vieira, F.P. Werneck and G.R. Colli. .
Microhabitat Variation Explains Local-scale Distribution of
Terrestrial Amazonian Lizards in Rondônia, Western Brazil.
Biotropica (): –. doi: . / j.-...x
Kohler, G. . Reptiles of Central America. Oenbach: Herpeton
Verlag.  pp.
Lee, J.C. . A eld guide to amphibians and reptiles of the Maya
World: the lowlands of México, Northern Guatemalan and
Belize. Ithaca: Cornell University Press.  pp.
Lyons, S.K. and M.R. Willig. . Species richness, latitude, and
scale-sensitivity. Ecology (): –. doi: ./-
()[:SRLASS]..CO;
Mittermeier, R. and C. Goettsch-Mittermeier. . Megadiversity:
the biologically richest countries of the world. México City:
Conservation International/CEMEX.
Stevens, G. C. . e latitudinal gradient in geographical range: how
so many species coexist in the tropics. e American Naturalist
: –. doi: http://www.jstor.org/stable/
Vester, H.F., D. Lawrence, J.R. Eastman, B.L. Turner, S. Calme, R.
Dickson and F. Sangermano. . Land change in the southern
Yucatan and Calakmul Biosphere Reserve: eects on habitat
and biodiversity. Ecological Applications (): –. doi:
./-
Vitt, J.L., G.R. Colli, J.P. Caldwell, D.O. Mesquita, A.A. Garda and
F.G.R. França. . Detecting variation in microhabitat use
in low-diversity lizard assemblages across small-scale habitat
gradients.Journal of Herpetology(): –. doi:./
-.
UNESCO. . Ancient Maya city and protected tropical forests of
Calakmul, Campeche. Accessed at http://whc.unesco.org/en/
list/,  October .
Authors’ contribution statement: KS and TJC carried out the
design of the study, coordinated the data collection and revised and
edited the manuscript. JALBN participated in the herpetological
surveys and species identication and prepared the rst draft of the
manuscript. RM, AL, JW, GC, SH, AR, ET, and RJW participated in
the herpetological surveys and species identication. All authors
read and approved the nal manuscript.
Received:  April 
Accepted:  August 
Academic editor: Ross MacCulloch