\documentclass[10pt]{article}

\usepackage{fullpage}
\usepackage{setspace}
\usepackage{parskip}
\usepackage{titlesec}
\usepackage[section]{placeins}
\usepackage{xcolor}
\usepackage{breakcites}
\usepackage{lineno}
\usepackage{hyphenat}





\PassOptionsToPackage{hyphens}{url}
\usepackage[colorlinks = true,
            linkcolor = blue,
            urlcolor  = blue,
            citecolor = blue,
            anchorcolor = blue]{hyperref}
\usepackage{etoolbox}
\makeatletter
\patchcmd\@combinedblfloats{\box\@outputbox}{\unvbox\@outputbox}{}{%
  \errmessage{\noexpand\@combinedblfloats could not be patched}%
}%
\makeatother


\usepackage{natbib}




\renewenvironment{abstract}
  {{\bfseries\noindent{\abstractname}\par\nobreak}\footnotesize}
  {\bigskip}

\titlespacing{\section}{0pt}{*3}{*1}
\titlespacing{\subsection}{0pt}{*2}{*0.5}
\titlespacing{\subsubsection}{0pt}{*1.5}{0pt}


\usepackage{authblk}


\usepackage{graphicx}
\usepackage[space]{grffile}
\usepackage{latexsym}
\usepackage{textcomp}
\usepackage{longtable}
\usepackage{tabulary}
\usepackage{booktabs,array,multirow}
\usepackage{amsfonts,amsmath,amssymb}
\providecommand\citet{\cite}
\providecommand\citep{\cite}
\providecommand\citealt{\cite}
% You can conditionalize code for latexml or normal latex using this.
\newif\iflatexml\latexmlfalse
\providecommand{\tightlist}{\setlength{\itemsep}{0pt}\setlength{\parskip}{0pt}}%

\AtBeginDocument{\DeclareGraphicsExtensions{.pdf,.PDF,.eps,.EPS,.png,.PNG,.tif,.TIF,.jpg,.JPG,.jpeg,.JPEG}}

\usepackage[utf8]{inputenc}
\usepackage[T2A]{fontenc}
\usepackage[ngerman,polish,english]{babel}



\usepackage{float}








\begin{document}

\title{A Rapid Inventory of Amphibians, Squamates, and Bats of Mata de Pl\selectlanguage{ngerman}átano
Field Station and Nature Reserve, Arecibo, Puerto Rico\selectlanguage{english}}



\author[1]{Justin M. Bernstein}%
\author[2]{Camilo Calderón-Acevedo}%
\author[2]{Pedro Mônico}%
\author[3]{Lázaro Viñola-Lopez}%
\author[2]{J. Soto-Centeno}%
\affil[1]{University of Kansas College of Liberal Arts and Sciences}%
\affil[2]{Rutgers University Newark}%
\affil[3]{University of Florida}%


\vspace{-1em}



  
  \date{May 22, 2024}


\begingroup
\let\center\flushleft
\let\endcenter\endflushleft
\maketitle
\endgroup





\selectlanguage{english}
\begin{abstract}
Puerto Rico harbors a diverse vertebrate fauna with high levels of
endemism. However, while several books for vertebrate diversity and
local checklists for birds have been published, checklists of
amphibians, reptiles, and bats are lacking or nonexistent at both local
and regional scales. In this study, we documented the amphibian,
reptile, and bat faunas at Mata de Pl\selectlanguage{ngerman}átano Field Station and Nature
Reserve, in Arecibo, Puerto Rico. We document four species of
amphibians, seven lizards, three snakes, and nine bats. Despite prior
works using samples from this nature reserve, this represents the first
vertebrate checklist for the Mata de Plátano Field Station and Nature
Reserve. We provide additional natural history observations based on our
survey results and highlight the importance of including local and
regional checklists of species for downstream research and conservation.%
\end{abstract}\selectlanguage{ngerman}%



\sloppy


\selectlanguage{ngerman}\textbf{A Rapid Inventory of Amphibians, Squamates, and Bats of Mata de
Plátano Field Station and Nature Reserve, Arecibo, Puerto Rico}

Justin Matthew Bernstein\textsuperscript{1, 2, 3, *}, Camilo Andrés
Calderón-Acevedo\textsuperscript{4}, Pedro Ivo
Mônico\textsuperscript{4}, Lázaro Willian
Viñola-Lopez\textsuperscript{5}, J. Angel
Soto-Centeno\textsuperscript{4,6}

\textsuperscript{1} Department of Biological Sciences, Rutgers
University-Newark, Newark, New Jersey, United States of America

\textsuperscript{2} Center for Genomics, University of Kansas, Lawrence,
Kansas, United States of America; email: jmbernst223@gmail.com; ORCID
id: 0000-0002-5249-3340

\textsuperscript{3} Current address: Department of Biology, University
of Texas at Arlington, Arlington, Texas, United State of America

\textsuperscript{4} Department of Earth and Environmental Science,
Rutgers University-Newark, Newark, New Jersey, United States of America
(CAC, PIM, ASC): emails: calderon.mammalogy@gmail.com (CAC),
pm859@scarletmail.rutgers.edu (PIM), angelo.soto@rutgers.edu (ASC);
ORCID ids: 0000-0003-2342-3981 (PIM), 0000-0002-1468-3565 (CAC),
0000-0002-1729-1153 (JASC)

\textsuperscript{5} Department of Biology and Florida Museum of Natural
History, University of Florida, Gainesville, Florida, United States of
America: email: lvinolalopez@ufl.edu; ORCID id: 0000-0002-2956-6759

\textsuperscript{6} Department of Mammalogy, American Museum of Natural
History, New York, New York, United States of America.

* Corresponding author

\textbf{Abstract}

Puerto Rico harbors a diverse vertebrate fauna with high levels of
endemism. However, while several books for vertebrate diversity and
local checklists for birds have been published, checklists of
amphibians, reptiles, and bats are lacking or nonexistent at both local
and regional scales. In this study, we documented the amphibian,
reptile, and bat faunas at Mata de Plátano Field Station and Nature
Reserve, in Arecibo, Puerto Rico. We document four species of
amphibians, seven lizards, three snakes, and nine bats. Despite prior
works using samples from this nature reserve, this represents the first
vertebrate checklist for the Mata de Plátano Field Station and Nature
Reserve. We provide additional natural history observations based on our
survey results and highlight the importance of including local and
regional checklists of species for downstream research and conservation.

\textbf{Keywords}

Bats, Caves, Frogs, Lizards, Karst, Snakes, Tropical moist forest

\textbf{Introduction}

Puerto Rico is an island of the Greater Antilles comprising
\textasciitilde{}9100 km\textsuperscript{2}, and located at 18.2° N,
66.6° W between the Virgin Islands and Hispaniola. The island
constitutes a small archipelago of over 125 islands and cays that are
geologically complex. About 28\% of the terrain of the main island of
Puerto Rico is covered in limestone cliffs, valleys, and hills with a
dichotomy between mesic and xeric forests in the northern and southern
regions, respectively (Monroe 1976; Lugo et al. 2001). The heterogeneity
of Puerto Rico's landscape and habitats harbor diverse vertebrate fauna,
especially in areas associated with forests and karst formations. Among
the focal taxa in this study, the diversity of this island includes a
total of 18 amphibians, 72 squamates (lizards and snakes) (Rivero 1998;
Uetz et al. 2023) and 13 bats (Gannon et al. 2005). For the
herpetofauna, about 60 species (43\%) are endemic to Puerto Rico; out of
the bat species, 2 (15\%) and 6 (46\%) are locally endemic to Puerto
Rico and regionally endemic to the West Indies, respectively. The bats
and herpetofauna of Puerto Rico are continuously being studied and have
served as models for studying natural disaster effects on island
ecosystems (Calderón-Acevedo et al. 2021), hybridization and sex
chromosome research (Pinto et al. 2019, 2022), and niche partitioning
genome-environmental association studies (Wogan et al. 2020; Ingram et
al. 2022). Nonetheless, distribution and taxonomic accounts of these
taxa in locally protected habitats and preserves are generally lacking.
Checklists at local and island-wide scales for Puerto Rico are limited
to algae (Ballantine and Aponte 1997), insects (Ramírez et al. 2020),
birds (Arendt et al. 2015), and arthropods (Vélez Jr. Jr. 1967;
Pérez-Reyes et al. 2013; Ospina-Sánchez et al. 2020), but are lacking
for bats, squamates, and amphibians. The need for more regional and
up-to-date checklists in Puerto Rico is important for biogeographic,
evolutionary, and conservation studies, as many taxa have restricted
ranges and are isolated to regional caves and/or forests (Rivero 1998;
Kurta and Rodriguez-Durán 2005). Here, we provide the first faunal
checklist for Mata de Plátano Field Station and Nature Reserve
(collectively referred to as Mata de Plátano here), an area in the
north-central subtropical moist forests of Arecibo that contains
multiple caves, including Cueva de los Culebrones, a cave system
well-known for the predator-prey interactions between the endemic boa
(\emph{Chilabothrus inornatus} ) and several bat species.

\textbf{Study Area}

Mata de Plátano consists of a 53-hectare, protected nature reserve,
about 7 km southwest of the city of Arecibo. It is located in the
northern karst belt region of Puerto Rico (18.414, -66.729) at 151 m
above sea level (Figure 1). The reserve is part of the largest
unfragmented and species rich forests of Puerto Rico. The sub-tropical
moist forested areas of the reserve consist of a few undisturbed areas
amongst different successional stages of other forests. These areas have
canopies up to \textasciitilde{} 20 m that receive precipitation that
ranges from 100--220 cm per year, with dry months being from
December--March, and wet months from April--November (Ewel and Whitmore
1973). The study area in this checklist includes two cave systems: Cueva
Larvas (18.414° N, -66.727° W) and Cueva de los Culebrones (18.414° N,
66.725° W). Cueva Larvas is a small cave with a wide entrance, while
Cueva de los Culebrones is a 182-meter (m) long hot cave with an
entrance that is 5 x 3 x 8 (width x height x depth, m; Puente-Rolón
1999).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

\textbf{Figure 1.} Map of Puerto Rico, with Arecibo shaded in yellow.
The blue dots represent the location of Mata de Pl\selectlanguage{ngerman}átano Field Station.

\textbf{Methods}

\emph{Herpetofauna} ---. Field surveying was conducted from 1--9 June
2022. Opportunistic sampling was performed to find lizards and snakes
under logs, rocks, in leaf litter, on trees, and throughout cave
systems. Non-protected species were caught by hand, while taxa that are
protected and listed as endangered by U.S. Fish \& Wildlife Service
(see\emph{Results} ) were identified with photo vouchers and observed
from a distance. Photographs were taken with a Nikon 70D DSLR camera
with a Tamron AF 90 mm macro lens. Species identification was done using
morphological criteria from (Rivero 1998). No specimens were collected
but were easily identifiable based on distinguishing morphology from
syntopic taxa.

\emph{Bat fauna} ---. Field surveys for bats were conducted during 1--9
June 2022. Bats were captured in mist nets (6--9 m long by 3 m high,
Avinet 38mm mesh) set along forest corridors near the field station.
Captured bats were identified to species and standard measurements
(i.e., forearm and weight) were taken. Photographs were taken with a
Nikon 70D DSLR camera with a Tamron AF 90 mm macro lens and with a
Cannon G12. No specimens were collected during surveys and all
individuals were released at the site of capture. This study was
conducted following the Departmento de Recursos Naturales de Puerto Rico
(permit no. 2022-IC-034 issued to JASC) and under compliance with
Rutgers IACUC (permit no. 201900087 issued to JASC). Coordinates for
sites of capture were taken with a Garmin GPSMap 65 series to the
nearest 0.00001 degree.

\textbf{Results}

During our field work we found a total of 14 species of herpetofauna
(four amphibians, seven lizards, three snakes) and 8 species of bats. We
list our inventory of amphibians, reptiles, and bats below.

\textbf{Amphibians}

\textbf{Family Bufonidae}

\emph{Rhinella marina} (Linnaeus, 1758)

Figure 2A

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found in leaf litter on south side of field station at
11:30 PM.

\textbf{Identification.} One adult individual, diagnosed by its large
size, enlarged parotid glands, and drier, warty skin distinguish this
amphibian from all others on the island. It is easily distinguished from
\emph{Peltophryne lemur} by its shorter rostrum that is not upturned.

\textbf{Family Eleutherodactylidae}

\emph{Eleutherodactylus coqui} Thomas, 1966

Figure 2B

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found in leaf litter on west side of field station at
6:00 PM; Individual 2 -- found on east wall of field station at 9:30 PM;
Individual 3 -- found in leaf litter at far end of open field 73 m south
of field station.

\textbf{Identification.} Three adults, identified by a combination of
morphology and geographic distribution. Diagnosed by eye color
(tan/brown) and color pattern (no dorsal stripe, saddle pattern on back
and dorsolateral lines). \emph{Eleutherodactylus monensis} , \emph{E.
cooki, E. cochranae} , \emph{E. portoricensis} , \emph{E. karlschmidti}
,\emph{E. gryllus} , \emph{E. locustus} , \emph{E. wightmanae} ,
\emph{E. eneidae} , \emph{E. richmondi} , \emph{E. unicolori} , and
\emph{E. jasperi} are all outside the range of our study
area.\emph{Eleutherodactylus brittoni} found in meadows and has a narrow
black streak at the tip of the snout, stretching and continuing along
the flanks (absent in our observed individuals).\emph{Eleurtherodactylus
portoricensis} has silver or chalky white eyes (tan/brown eyes in our
individuals) and restricted to mountains.\emph{Eleutherodactylus
hedricki} has a pronounced constriction behind the head and a shallow
furrow on the middle of the back from the head to the sacrum (absent in
our individuals). \emph{Eleutherodactylus richmondi} has yellow or white
lines on each side of the head, that range from the tip of the snout to
near the vent (absent in our individuals). \emph{Eleutherodactylus
antillensis} has a dorsal stripe that stretches from the tip of the
snout to the vent (absent in our individuals) and reddish eyes
(tan/brown in our individuals).

\emph{Eleutherodactylus antillensis} (Reinhardt \& Lutken, 1863)

Figure 2C

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found in leaf litter near entrance of open field, 73 m
south of field station; Individual 2 -- caught in mist net placed at
path of open field 75 m south of field station 7:40 PM.

\textbf{Identification.} Two adults, identified by a combination of
morphology and geographic distribution; diagnosed by its thin,
mid-dorsal stripe on the head and body, and reddish eyes. Similar to the
identification of \emph{E. coqui} , \emph{E. monensis} , \emph{E. cooki,
E. cochranae} , \emph{E. portoricensis} , \emph{E. karlschmidti} ,
\emph{E. gryllus} , \emph{E. locustus} , \emph{E. wightmanae} , \emph{E.
eneidae} ,\emph{E. richmondi} , \emph{E. unicolori} , and \emph{E.
jasperi} are all outside the range of our study area.
\emph{Eleutherodactylus brittoni}found in meadows and has a narrow black
streak at the tip of the snout, stretching and continuing along the
flanks (absent in our observed individuals). \emph{Eleurtherodactylus
portoricensis} has silver or chalky white eyes (tan/brown eyes in our
individuals) and restricted to mountains. \emph{Eleutherodactylus
hedricki} has a pronounced constriction behind the head and a shallow
furrow on the middle of the back from the head to the sacrum (absent in
our individuals).\emph{Eleutherodactylus richmondi} has yellow or white
lines on each side of the head, that range from the tip of the snout to
near the vent (absent in our individuals). \emph{Eleutherodactylus
coqui} may or may not have the dorsal stripe that stretches from the tip
of the snout to the vent that is seen in \emph{E. antillensis} ,
\emph{E. coqui} has tan/brown eyes (reddish in our individuals).

\emph{Eleutherodactylus cochranae} Grant, 1932

Figure 2D

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found on east wall of field station at 9:10 PM.

\textbf{Identification.} One adult. Identified by its light coloration,
two concave dorsal pigmentation lines, and a faint line of pigmentation
in the middle of the snout.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

\textbf{Figure 2.} Frogs found at the Mata de Pl\selectlanguage{ngerman}átano Field Station and
Nature Reserve.\textbf{A.} \emph{Rhinella marina} . \textbf{B.}
\emph{Eleutherodactylus coqui} . \textbf{C.} \emph{Eleutherodactylus
antillensis} . \textbf{D.}\emph{Eleutherodactylus cochranae} . All
photographs taken by Justin M. Bernstein.

\textbf{Lizards}

\textbf{Family Dactyloidae}

\emph{Anolis cristatellus} Duméril \& Bibron, 1837

Figure 3A

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found on karst wall on southwest side of field station
at 6:00 PM; Individual 2--3 -- found on karst wall next to steel
entrance gate of field station, 138 m southwest of field station
building at 6:00 PM; Individuals 4--5 -- on walls, beneath second-level
deck of field station at 8:23 AM; Individual 6 -- Outside the entrance
of Cueva Larvas (18.414deg N, -66.727degW) in forest with cooler
microclimate, found on vines with spotty sunlight, at 12:25 PM;
Individuals 8 -- on palm tree at \textasciitilde{} 1.5 m off ground at
8:00 AM, performing territorial `push up' behavior; Individuals 9--10 --
on south wall of field station mating with each other at 4:58 PM;
Individuals 11--13 -- sleeping on plants on forest edge on south side of
field station at 12:30 AM; Individuals 14--17: found on plants on forest
edge on south side of field station at 8:00 AM; Individuals 18--19 on
branches \textasciitilde{}1.5 m above ground, near karst wall on
southwest side of field station at 2:45 PM; Individual 20 -- in forest
on leaf litter, \textasciitilde{}50 m south of Cueva de los Culebrones.

\textbf{Identification.} Identified based on morphology and geographic
distribution. The brown/tan and coloration, brown eyes, and crest of
this species distinguishes it from all other \emph{Anolis} in Puerto
Rico. \emph{Anolis evermanni} has green coloration, and \emph{A.
stratulus} has large, dorsal, saddle-shaped blotches on the back
distinguished. \emph{Anolis gundlachi} has blue eyes, \emph{A. krugi}has
yellow or cream-colored lateral stripes (absent in our individuals), and
\emph{A. cuvieri} is a large, green crown giant anole. All
other\emph{Anolis} are found outside the range of Mata de Platano.

\emph{Anolis cuvieri} Merrem, 1820

Figure 3B

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found sleeping on U-shaped vine \textasciitilde{} 4.5 m
above ground at southeast edge of field station at 9:00 PM.

\textbf{Identification.} This species is readily distinguished from all
congeners by being one of two crown giant anoles in the Puerto Rico
Bank. It's bright green coloration, large dorsal body and tail crests,
and large size (\textasciitilde{}125 mm; 5 mm snout-vent-length; Rivero
1998) separate it from all other \emph{Anolis} . The other crown
giant,\emph{A. roosevelti} , was distributed to the Puerto Rico
satellite islands of Culebra and Vieques, and Tortola and St. John of
the Virgin Islands, but is presumed to be extinct (Rivero, 1998).

\emph{Anolis krugi} PETERS, 1877

Figure 3C

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1--2 -- found on plants on forest edge on south side of field
station at 5:00 PM; Individual 3 -- on branch \textasciitilde{}1.5 m
above ground, over karst wall on southwest side of field station at 3:00
PM; Individual 4 -- on branch \textasciitilde{}1.7 m above ground on
forest edge \textasciitilde{}50 m south of field station; Individual 5
-- in forest on ground, \textasciitilde{}50 m south of Cueva de los
Culebrones.

\textbf{Identification.} Distinguished from all other species by a
combination of the presence of prominent yellow/cream-colored lateral
lines that extend from the eye to the groin (also found in \emph{A.
pulchellus} ) and numerous black specks on the sides of the back and
flanks (absent in \emph{A. pulchellus} ), and the absence of a brown
band behind the eye. All other \emph{Anolis} are found outside the range
of Mata de Platano or lack the lateral lines and black specks on the
back and flanks.

\emph{Anolis stratulus} Cope, 1861

Figure 3D

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- On south wall of field station at 11:00 AM; Individuals
2--3 -- found on karst wall on southwest side of field station at 8:23
AM; Individuals 4--6 -- found on karst wall on southwest side of field
station 11:07 AM; Individuals 7--8 -- sleeping on plants on forest edge
on south side of field station at 12:30 AM.

\textbf{Identification.} Readily identified from all other
\emph{Anolis}on the island by the prominent saddle-shaped blotched on
the dorsum. The lack of a crest on the dorsum or tail distinguish it
from all other Puerto Rican anoles, with the exception of \emph{A.
evermanni} and\emph{A. occultus} . \emph{Anolis evermanni} has bright
green coloration (tan/brown in our individuals) and \emph{A. occultus}
has a downward taping snout and a smaller body size and tail than
\emph{A. stratulus} ; both \emph{A. evermanni} and \emph{A. occultus}
lack saddle-shaped blotches on the dorsum.

\textbf{Family Teiidae}

\emph{Pholidoscelis exsul} (Cope, 1862)

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1--2 -- on ground of trail \textasciitilde{}6m south of field
station at 8:40 AM; Individua 3 -- on ground of trail \textasciitilde{}
15 m south of field station.

\textbf{Identification.} \emph{Pholidoscelis exsul} is one of few teiid
lizards found on Puerto Rico. It is distinguished from its island
conspecifics by a combination of morphology (color pattern), but is
primarily identified by geographic distribution. \emph{Pholidoscelis
wetmorei} is only found in the southwest part of the island, \emph{P.
alboguttatus} is endemic to Mona Island, and \emph{P.
desechensis}endemic to Desecheo Island.

\textbf{Family Sphaerodactylidae}

\emph{Sphaerodactylus grandisquamis guarionex} Stejneger, 1904

Figure 3E

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- under palm trash on side of trail \textasciitilde{}15 m
south of field station at 10:30 PM; Individual 2 -- under palm trash on
side of trail \textasciitilde{}15 m south of field station at
\textasciitilde{}8:30 AM; Individual 3 -- found on top of leaf litter in
open field 75 m south of field station at 6:20 PM; Individuals 4--10 --
found on top of leaf litter in open field 85 m south of field station at
6:30 PM; Individual 11 -- found on top of leaf litter in open field 85 m
south of field station at 7:35 PM; Individuals 12 -- 16 -- found on top
of leaf litter in open field 85 m south of field station at 9:50 AM;
Individuals 18--22 -- found on top of leaf litter in open field 85 m
south of field station at 2:50 PM

\textbf{Identification.} The only species of \emph{Sphaerodactylus} on
the northern part of mainland Puerto Rico are \emph{S. g. guarionex}
and\emph{S. klauberi} (sensu Daza et al. 2019). \emph{Sphaerodactylus g.
guarionex} is distinguished from \emph{S. klauberi} by its smaller size
and lighter color (larger total length and snout-vent-length and dark,
near-black color in \emph{S. klauberi} ); \emph{S. g. guarionex} is
distinguished by other subspecies of \emph{S. grandisquamis} by its
reduced number of dorsal body scales (\textasciitilde{}14 for \emph{S.
g. guarionex} , 17--23 in \emph{S. g. grandisquamis} , \emph{S. g.
spanius} , \emph{S. g. mimetes} , and \emph{S. g. ateles} ) and rounded
snout scales (Daza et al. 2019). Molecular and morphological data from
recent studies support the claim that \emph{Sphaerodactylus} in the
northern part of the mainland are \emph{S. g. guarionex} or \emph{S.
klauberi} (Daza et al. 2019; Reynolds et al. 2021).

\textbf{Family Typhlopidae}

\emph{Anillotyphlops} sp.

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found in abandoned termite nest on side of trail
\textasciitilde{} 2 m above the ground, \textasciitilde{}125 m southwest
of field station, at 11:00 PM. Individual quickly escaped and fell down
an escarpment upon discovery.

\textbf{Identification.} Individual could not be identified to the
species level, but is distinguished from all other snakes on the island
by its smooth, shiny scales, extremely small size with cylindrical body
(\textasciitilde{}15 cm total length), neck not visibly distinct from
head, and the utilization of termite nest microhabitats.

\textbf{Family Dipsadidae}

\emph{Borikenophis portoricensis} (Reinhardt \& Lutken, 1862)

Figure 3F

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1 -- found under tarp from an old harp net bat trap on side
of trail, \textasciitilde{}10 m from the entrance of Cueva de los
Culebrones, at 12:52 PM.

\textbf{Identification.} Easily diagnosed from blind snakes
(\emph{Anillotyphlops} , \emph{Typhlops} ), by its significantly larger
size total length and circumference, and enlarged eyes. As a racer, it
is not similar in appearance to \emph{Xenocrophis} or\emph{Chilabothrus}
. \emph{Borikenophis portoricensis} is distinguished from
\emph{Magliophis} by its larger size (\emph{Magliophis}\textasciitilde{}
0.5 m; \emph{Borikenophis} from this study \textasciitilde{}1--1.2 m
total length). \emph{Borikenophis portoricensis} can be distinguished
from its congeners by distribution and morphology: \emph{B. prymnus} is
smaller and paler in color and is found in southern Puerto Rico and
\emph{B. variegatus} is endemic to Mona Island.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\textbf{Figure 3.} Lizards and snakes found at the Mata de Pl\selectlanguage{ngerman}átano Field
Station and Nature Reserve. \textbf{A.} \emph{Anolis cristatellus}
.\textbf{B.} \emph{Anolis cuvieri} , perched on a high branch; inset
image zooms in on individual, showing the prominent crests and bright
green color. \textbf{C.} \emph{Anolis kugri} ; note the lateral lines
and flecks of dark pigmentation on the flanks in the inset
image.\textbf{D.} \emph{Anolis stratulus} ; inset image better shows the
saddle-shaped blotches on the dorsum. \textbf{E.} \emph{Sphaerodactylus
grandisquamis guarionex} . \textbf{F.} \emph{Borikenophis portoricensis}
. All photographs taken by Justin M. Bernstein.

\textbf{Family Boidae}

\emph{Chilabothrus inornatus} (Reinhardt, 1843)

Figure 4A--H

\textbf{Materials examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
Individual 1--8 -- adult individuals found hanging on branches or on
rock ledges of entrance of Cueva de los Culebrones, either perched or
dangling vertically. One individual observed eating \emph{Mormoops
blainvillei} . All individuals seen at once at 7:00 PM; Individuals
9--11 -- juvenile individuals found hanging verticlles from branches or
rock ledges of entrance of Cueva de los Culebrones at 7:00 PM. All
juveniles seen at the same time as Individuals 1--8; Individual 12 --
found inside Cueva de los Culebrones \textasciitilde{}17.6 m depth
moving along the ground; Individuals 13--14 -- adult skeletons, found at
\textasciitilde{}17.6 m depth in Cueva de los Culebrones; one skeleton
\textasciitilde{}30 m past cave entrance; second skeleton
\textasciitilde{}15 m before cave thermocline at \textasciitilde{}20 m
depth. Individuals 15--16 -- live adults, found coiled on limestone wall
ledges \textasciitilde{}1 m above the ground at the point of the
thermocline inside Cueva de los Culebrones, at \textasciitilde{}20 m
depth.

\textbf{Identification.} As the only boa on Puerto Rico, this snake is
unequivocally identifiable as \emph{E. inornatus} . Its large, stout
body, grey/silver scales, and the lack of labial thermic receptors are
key characters of this species. \emph{Chilabothrus inornatus} is
distinguished from \emph{C. granti} by the latter only being found on
satellite islands (Culebra, and islets to the east) and \emph{C.
monensis} only found on Mona Island. Bones found in Cueva de los
Culebrones were identified by the enlarged, recurved teeth
characteristic of boids; all other snakes on the island lack these and
have much smaller teeth.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

\textbf{Figure 4.} \emph{Chilabothrus inornatus} from Cueva de los
Culebrones. \textbf{A.} Lateral view of \emph{C. inornatus} perches on a
branch. \textbf{B.} Lateral shot of head, perched on a high
branch.\textbf{C.} Entrance of Cueva de los Culebrones at night, as bats
fly out of the cave and six individuals (white arrows) of \emph{C.
inornatus} wait to catch their prey. \textbf{D--E.} Two difference
individuals of \emph{C. inornatus} hanging vertically to catch bats
flying out of the cave; Panel D shows an individual of \emph{Mormoops
blainvillei} flying in front of a Puerto Rican Boa. \textbf{F.} A Puerto
Rican Boa constricting its prey, \emph{M. blainvillei} . \textbf{G.} A
dark pigmented individual of \emph{C. inornatus} from the thermocline
inside Cueva de los Culebrones. \textbf{H.} Bones of \emph{C.
inornatus}found inside Cueva de los Culebrones. Cranial elements,
vertebrae, and ribs were found. All photographs taken by Justin M.
Bernstein.

\textbf{Chiroptera}

\textbf{Family Mormoopidae}

\emph{Mormoops blainvillei} Leach, 1821

Figure 5A

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} * Mata de
Pl\selectlanguage{ngerman}átano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andrés Calderón-Acevedo,
Pedro Ivo Mônico, Lázaro Willian Viñola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 1 \selectlanguage{english}, ASC 1490; 1 \selectlanguage{english}, ASC 1485.

\textbf{Identification.} The Antillean ghost-faced bat (\emph{M.
blainvillei} ) has a unique facial morphology that easily identifies it
in the field. Nose-leaf absent, snout is short and adorned with skin
folds, the skin flap in the lower lip is split in two, giving the
appearance of paired leaflets. Fur bright orange/reddish, the ears are
joined by a membrane across the head. The ears are broad and round, with
the outer rims connecting to the lower lip which gives the eyes the
appearance of being immersed in the ear, inside a funnel. Size small,
total length = 78-87 mm (tail length =21 mm), forearm= 45--48 mm)
(Gannon et al., 2005; Wilson, 2023a).

\emph{Pteronotus portoricensis} Miller, 1902

Figure 5B

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 1 \selectlanguage{english}, ASC 1492.

\textbf{Identification.} The Puerto Rican mustached bat (\emph{P.
portoricensis} ) has dark grey to greyish-brown fur coloration, with
ventral hairs being paler. Nose-leaf absent, the snout protrudes more
than in \emph{M. blainvillei} , with the skin flap in the lower lip
continuous and not split. The ears are pointed and with straight edges,
not joined in the middle, eye position normal. It can only be confused
with \emph{P.} \emph{quadridens} , however its considerably larger with
a total length of 78--83 mm, and a forearm over 44 mm. Additionally it
possesses a fleshy knob above the snout, which is lacking in \emph{P.
quadridens} (Gannon et al., 2005; Pavan, 2023).

\emph{Pteronotus quadridens} Gundlach, 1840

Figure 5C

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 2 \selectlanguage{english}, ASC 1484, ASC 1486; 3 \selectlanguage{english}, ASC 1484,
1489, ASC 1491.

\textbf{Identification.} The sooty mustached bat (\emph{P. quadridens} )
has greyish-brown fur with tricolored individual hairs. Nose-leaf
absent, as in \emph{P.} portoricensis, the snout protrudes more than
in\emph{M. blainvillei} , with the skin flap in the lower lip continuous
and not split. The ears are pointed and with straight edges, not joined
in the middle, eye position normal. Considerably smaller than \emph{P.
portoricensis,} total length = 59--80 mm, forearm length = 36--39
millimeters. Can be discriminated from \emph{P. portoricensis} by having
a row of three to five small bulbs above each nostril, and the lack of a
fleshy knob above the snout (Gannon et al., 2005; Wilson, 2023b).

\textbf{Family Phyllostomidae}

\emph{Artibeus jamaicensis} Leach, 1821

Figure 5D

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 1 \selectlanguage{english}, ASC 1487; 1 \selectlanguage{english}, ASC 1493.

\textbf{Identification.} Jamaican fruit-eating bats (\emph{A.
jamaicensis} ) were identified by their ashy-gray coloration and faint
but noticeable lighter color stripes on the rostrum. These bats have a
broad nose leaf, a large central chin papillae surrounded by smaller
ones, and a tail membrane that forms a V shape with no tail. Jamaican
fruit bats are medium sized, one of the stockier bats on the island,
have broad dark brown wings, and are commonly captured on understory
mist nets. We identified a small colony of Jamaican fruit bats present
in Cueva Larvas.

\emph{Brachyphylla cavernarum} Gray, 1834

Figure 5E

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 1 \selectlanguage{english}, ASC 1487; 1 \selectlanguage{english}, ASC 1493.

\textbf{Identification.} Antillean fruit-eating bats (\emph{B.
cavernarum} ) are medium sized and identified by their light brown to
grayish pelage that is lighter color at the base. These bats have a
small and rudimentary nose-leaf that almost give it a pig-like
appearance. Small papillae on the chin of Antillean fruit-eating bats
form a distinctive V shape. The wings are broad and grayish in color.
This species is a year-round resident of Cueva de los Culebrones.

\emph{Erophylla bombifrons} Miller, 1899

Figure 5F

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
observed roosting and flying in first open room of Cueva de los
Culebrones.

\textbf{Identification.} Brown flower bats (\emph{E. bombifrons} ) have
light brown to yellowish colored fur that is very short. We identified
roosting and flying brown flower bats at Cueva de los Culebrones by
their characteristic hair color, which easily reflects light. In the
hand, brown flower bats are small bats with a slightly elongated and
semi-naked rostrum. Their nose leaf is small and pointy. They have a V
shaped tail membrane and a tail shorter than the femur extends beyond
the membrane. These bats are abundant at Cueva de los Culebrones and
often are hunted by the Puerto Rican boa. Although no individuals were
captured in mist nets, brown flower bats can be captured in understory
mist nets set close to fruiting Panama berry trees (\emph{Mutingia
calabura} ), piper plants (\emph{Piper aduncum} ), or turkey berry
plants (\emph{Solanum torvum} ). This species is a year-round resident
of Cueva de los Culebrones.

\emph{Monophyllus redmani} Leach, 1821

Figure 5G

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 1 \selectlanguage{english}, ASC 1488.

\textbf{Identification.} The Greater Antillean long-tongued bat
(\emph{M. redmani} ) is easily identified by being the smallest
leaf-nosed bat on the island. This primarily nectarivorous species has
grayish brown pelage, sometimes with patches of white hairs present on
the dorsal side. The ventral side is lighter grayish brown in color.
This bat has an elongated rostrum culminating in a distinctive arrow
shaped nose leaf. The tail membrane is V shaped and a short tail extends
beyond the tip of the membrane. Greater Antillean long-tongued bats are
captured in understory mist nets near fruiting Panama berry trees
(\emph{M. calabura} ) and flowering endemic Maga trees (\emph{Thespesia
grandiflora} ). This species is a year-round resident of Cueva de los
Culebrones.

\emph{Stenoderma rufum} Desmarest, 1820

Figure 5H

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 1 \selectlanguage{english}, ASC 1494.

\textbf{Identification.} The red fig-eating bat (\emph{S. rufum} ) is
the only other member of the subfamily Stenodermatinae inhabiting Puerto
Rico. Nose leaf present and broad, as in \emph{A. jamaicensis} , its fur
is tan to dark chocolate-brown (its conspecifics inhabiting St. Croix
and St. John having a reddish hue in their fur), the venter is lighter
than the back, and it is the only bat in Puerto Rico with distinctive
white epaulettes. Although it is similar to \emph{A. jamaicensis} , it
can be diagnosed by its smaller size, total length = 60--73 mm and
forearm length = 46--52 mm. Additionally it lacks any facial stripes
(Gannon et al., 2005; Genoways, 2023). The connectivity and abundance of
its populations is affected by seasonal climatic disturbances like
hurricanes (Calderon-Acevedo et al., 2021; Gannon \& Willig, 1994).

\textbf{Family Vespertilionidae}

\emph{Eptesicus dutertreus} Gervais, 1837

Figure 5A

\textbf{Materials Examined.} PUERTO RICO -- \textbf{Arecibo} \selectlanguage{english}* Mata de
Platano Field Station and Nature Reserve; 18.414, -66.729; 151 m above
sea level; Justin Matthew Bernstein, Camilo Andres Calderon-Acevedo,
Pedro Ivo Monico, Lazaro Willian Vinola-Lopez, J. Angel Soto-Centeno;
captured in six-meter mist net in open field 75 m south of field station
between 6:45 PM and 8:30 PM; 1 \selectlanguage{english}, ASC 1483.

\textbf{Identification.} The identity of Greater Antillean serotine bats
(\emph{E. dutertreus} ) was confirmed by their characteristically dark
chocolate brown fur, and fully naked nearly black round ears, wing, and
tail membranes. Both the tail and wing membranes look oily. The rostrum
of Greater Antillean serotines is naked and wide, giving it an
appearance of being inflated, a trait not found in other bats in Puerto
Rico. These bats are fast fliers that forage above the canopy of trees.
Our capture of Greater Antillean serotine bats in a forest corridor
using understory mist nets is attributed to the presence of Cueva Larvas
that serves as a roost nearby.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

\textbf{Figure 5.} Bats caught in the mist nets at Mata de Pl\selectlanguage{ngerman}átano.
Photos and parts of the body have been cropped out for easier viewing of
headshots. \textbf{A.} \emph{Mormoops blainvillei} .
\textbf{B.}\emph{Pteronotus portoricensis} . \textbf{C.}
\emph{Pteronotus quadridens} . \textbf{D.} \emph{Artibeus jamaicensis} .
\textbf{E.}\emph{Brachyphylla cavernarum} . \textbf{F.} \emph{Erophylla
bombifrons} .\textbf{G.} \emph{Monophyllus redmani} . \textbf{H.}
\emph{Stenoderma rufum} . \textbf{I.} \emph{Eptesicus dutertreus} .
Photo credits: A,B,D,G,H,I: Justin M. Bernstein; C,E,F: Angel
Soto-Centeno.

\textbf{Discussion}

Mata de Plátano Field Station and Nature Reserve (MPFS) is best known
for its bat-snake interactions, yet no faunal surveys for either of
these groups have been published. Our study shows that while small
(\textasciitilde{}5100 m\textsuperscript{2}), this reserve has a rich
herpetofauna and bat fauna representative of the overall diversity of
the island. This rapid inventory provides the first local,
taxonomic-focused checklist for these species in north-central Puerto
Rico.

In our 10-day rapid inventory, we found a total of 14 species of
reptiles and amphibians. We documented four amphibians, seven lizards,
and three snakes, totaling up to \textasciitilde{}22\% (4 out of 18) and
\textasciitilde{}14\% (10 out of 72) of the amphibians and squamate
diversity on Puerto Rico. Our observations in during field work also
provide important natural history information. We documented~\emph{A.
cristatellus~} copulating on the field station walls, which is not
surprising as this species is often found in man-made structures
throughout the island and is abundant in urban areas. Additionally, we
documented two morphs of \emph{A. cristatellus} : 1) light grey/tan
(with or without darker mottling), and 2) a morph that is dark or light
brown with a light yellow or cream colored mid-dorsal stripe and
light-colored dorsal and lateral flecks. The dorsum of the first morph
sometimes has dark gray or black spots running down the spine but are
not nearly as distinct or as large as the saddle-shaped botched
of~\emph{A. stratulus}(Fig. 6). The dorsal crest and tail fin in both
morphs is sometimes prominent, while other observations the cutest was
hardly distinct from the rest of the body. We also report notes on the
endemic~\emph{Chilabothrus inornatus} . This species is known for
consumption of bats on the island, and many individuals aggregate at the
entrance and inside caves to opportunistically catch them. We saw one
adult individual attack and devour a ghost faced bat~(\emph{Mormoops
blainvillei} ). In our night surveys, we observed juvenile \emph{C.
inornatus~} snakes (N = 3)~at the entrance of Cueva de los Culebrones,
which suggests an actively breeding population of this endemic snake. We
also found three live (and an additional two separate
skeletons)~\emph{C. inornatus} ~inside the cave, at 17.6 m and
\textasciitilde{}20 m depth. The latter of these two depths is the
thermocline of the cave (\textasciitilde{}35° C), where we found two
live~\emph{C. inornatus} . Individuals found deeper in the cave range
from light grey to dark, charcoal grey (Fig. 5G). Compared to the
ambient temperature (average: 24.4--31.1° F), the endemic Puerto Rican
Boas tolerate a wide range of temperatures.

During this rapid inventory we observed nine species of bat, all of
which were documented in a single 1.75-hour period of mist netting. This
bat diversity represents 70\% (i.e. 9 of 13) of the living alpha
diversity present on the island of Puerto Rico. Most of the bats we
observed are residents of MPFS. While our sampling efforts do not permit
a more thorough examination of species abundance or comparisons with
other reserves in the island, these data highlight the importance of
MPFS as a local center of bat diversity. The moist forest area where
MPFS is located is adjacent to pastures, unprotected rustic forest
areas, and the urbanized center of the municipality of Arecibo to the
north. However, more importantly, the station connects with the larger
Río Abajo State Forest to the south. Together these two protected areas
form a hub in the northcentral part of Puerto Rico extending from
coastal to mountainous areas and linking multiple forested corridors
that connect with reserves along the Karst Conservation Zone
(Calderón-Acevedo et al., 2001).

Two endemic bat species documented in this rapid survey are of
particular importance. The endemic Puerto Rican moustached bat
(\emph{Pteronotus portoricensis} ) specifically roosts in hot cave
chambers where temperatures often exceed 35°C, such as those present in
Cueva de los Culebrones (Rodríguez-Durán and Soto-Centeno, 2003).
Previous studies show that the population this species at Cueva de los
Culebrones was not affected by hurricanes (Jones et al., 2001).
Nonetheless, \emph{P. portoricensis} seems to be a new resident to the
cave, and thus effects of natural phenomena on this species at this
locality are not well known. A single individual of another endemic, the
red fig-eating bat (\emph{Stenoderma rufum} ), was captured during our
survey. This endemic frugivorous bat is known to roost in the thick
foliage of trees and occurs in low abundance throughout the island
(Genoways 2023; Calderón-Acevedo et al., 2021). Given these
characteristics, \emph{S. rufum} is a species particularly affected by
strong hurricanes (Gannon and Willig 1994), which can decrease the
structural habitat connectivity through the island. The protection of
MPFS in the wider northcentral habitat connectivity hub of Puerto Rico
(Calderón-Acevedo et al., 2021), may be key for the recovery and
population connectivity of this rare bat.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}

\textbf{Figure 6.} Variation seen in \emph{Anolis cristatellus} from the
Mata de Pl\selectlanguage{ngerman}átano Field Station. \textbf{A.} Grey/tan patterning with
heavily mottling. \textbf{B.} Brown morph with mid-dorsal yellow stripe
and light dorsal and lateral flecks. \textbf{C.} Brown morph with no
mottling or mid-dorsal spots, light flecks present. \textbf{D.} Lateral
shot of the head of the individual from Panel C. \textbf{E.} Brown morph
with light mottling and striping, and dark spots running down the spine.
Note the slightly raised dorsal crest (more prominent at the
tail).\textbf{F.} Lateral shot of the head of the individual from Panel
F. All photographs taken by Justin M. Bernstein.

To our knowledge, this is the first faunal checklist on the bat and
herpetofauna of~Puerto Rico. Our findings expand the current knowledge
of biodiversity and species richness in this protected region. Although
more standardized methods over greater periods of time will provide more
insight on species abundances, our results show that \emph{A.
cristatellus} and \emph{S. grandisquamis guarionex} likely contribute
large portions to the vertebrate biomass here, as has been found in
other studies (Rodda et al. 2001). There are likely more species in Mata
de Plátano. For example, there have been several accounts of \emph{A.
occultus} from the field station (pers. comm. Abel Vale, 2022). Although
we did not see these species during our time at the reserve, future
surveys will be needed to better understand the density of \emph{A.
occultus} and \emph{N. leporinus} in this area. The Puerto Rican fauna
has been studied well, yet new species have recently described (e.g.,
Díaz-Lameiro et al. 2022), thus it is critical to perform local,
intensive surveys and document faunal inventories. These local intensive
surveys and faunal checklists can provide a steady supply of data to
understand the current status of Caribbean species. The main threats to
Caribbean bats and herpetofauna stem from habitat loss and fragmentation
related to human development, and climatic events that can drastically
re-shape the local population connectivity of bats species
(Calderón-Acevedo et al., 2021; Soto-Centeno \& Calderón-Acevedo, 2023).
Therefore, having updated local lists can provide a year-to-year
comparison of local faunal abundance that can inform conservation
strategies, and eventually strengthen the predictive power when
modelling the effect climate change of species richness and abundance.
We encourage more researchers to publish faunal and floral lists, even
for common taxa, to better understand species diversity, abundance, and
interactions for downstream research and conservation.

\textbf{Data Availability Statement}

All research here was observational and no data beyond what is presented
in this manuscript is available.

\textbf{Acknowledgements}

This rapid survey was sponsored by an NSF award (DEB 2135257) to JASC.
We thank Armando Rodríguez-Durán and Abel Vale for providing access to
Mata de Plátano Field Station. Work by CAC was partly funded by a
postdoctoral scholarship at the Soto Lab of Bat Biology (SLaBB) in
Rutgers University. JMB was funded by the University of Kansas Center
for Genomics, and an NSF Postdoctoral Research Fellowships in Biology
(Grant No. 2208959).

\textbf{References}

Arendt W.J., J. Faaborg, M. Canals, and J. Bauer. 2015. Bird checklist,
Guánica Biosphere Reserve, Puerto Rico. Research Note SRS-23. US Forest
Service, Southern Research Station, Ashville, NC, USA. 23 pp.

Ballantine D.L. and N.E. Aponte. 1997. A revised checklist of the
benthic marine algae known to Puerto Rico. \emph{Caribbean Journal of
Science} 33:150--179.

Calderón-Acevedo C.A., A. Rodríguez-Durán, and J.A. Soto-Centeno. 2021.
Effect of land use, habitat suitability, and hurricanes on the
population connectivity of an endemic insular bat. \emph{Scientific
Reports} 11:9115.

Daza J.D., B.J. Pinto, R. Thomas, A. Herrera-Martinez, D.P. Scantlebury
D, L.F.P. García, R.P. Balaraman, G. Perry, and T. Gamble. 2019. The
sprightly little sphaerodactyl: Systematics and biogeography of the
Puerto Rican dwarf geckos \emph{Sphaerodactylus} (Gekkota,
Sphaerodactylidae). \emph{Zootaxa} 4712:151--201.

Díaz-Lameiro A.M., C.I. Villamil, T. Gamble, B.J. Pinto, A.
Herrera-Martínez, R. Thomas, J.M. Bernstein, J.E. Titus-McQuillan, S.V.
Nielsen, E. Agosto-Torres, A.R. Puente-Rolón, F.J. Bird-Picó, T.K.
Oleksyk, J.C. Martínez-Cruzado, and J.D. Daza. 2022. A new species of
Sphaerodactylus (Gekkota: Sphaerodactylidae) from the Northwest
Limestone Region of Puerto Rico. \emph{Ichthyology \&
Herpetology}110(3):449--465.

Ewel J.J. and J.L. Whitmore. 1973. The Ecological Life Zones of Puerto
Rico and the U.S. Virgin Islands. USDA Forest Service, Institute of
Tropical Forestry, Research Paper ITF-018.

Gannon, M.R., A. Kurta, A. Rodríguez-Durán, and M.R. Willig.
2005.\emph{Bats of Puerto Rico: an island focus and a Caribbean
perspective} . 224 pp. Texas Tech University Press, Lubbock.

Gannon, M.R., and M.R. Willig. 1994. The effects of Hurricane Hugo on
bats of the Luquillo Experimental Forest of Puerto
Rico.\emph{Biotropica} 26(3): 320--331.

Genoways, H.H. 2023. \emph{Stenoderma rufum} . In: A. Kurta and A.
Rodríguez-Durán (Eds.), \emph{Bats of the West Indies: A Natural History
and Field Guide} , pp. 297--301. Cornell University Press, Ithaca.

Ingram T., S.T. Giery, J.B. Losos. 2022. Hierarchical partitioning of
multiple niche dimensions among ecomorphs, species and sexes in Puerto
Rican anoles. \emph{Journal of Zoology} 318(2): 127--134.

Jones, K.E., K.E. Barlow, N. Vaughan, A. Rodríguez-Durán, and M.R.
Gannon. 2001. Short-term impacts of extreme environmental disturbance on
the bats of Puerto Rico. \emph{Animal Conservation} 4:59--66.

Joglar, R., L.A.O. Álvarez, T.M. Aide, D. Barber, P.A. Burrowes, M.A.
García, A. León-Cardona, A.V. Longo, N. Pérez-Buitrago, A. Puente, N.
Ríos-López, and P.J. Tolson. 2011. Conserving the Puerto Rican
herpetofauna. In: Hailey, A., B.S. Wilson, and J.A. Horrocks
(Eds.),\emph{Conservation of Caribbean Island Herpetofaunas} , pp
339--358, Volume 2. Regional Accounts of the West Indies. Brill, Leiden,
The Netherlands.

Lugo A.E., L.M. Castro, V. Abel, T. del M. López, E.H. Prieto, and A.G.
Martinó, A.R. Puente Rolón, A.G. Tossas, D.D. McFarlane, T. Miller, A.
Rodríguez, J. Lundberg, J. Thomlinson, J. Colón, J.H. Schellekens, O.
Ramos, and E. Helmer. 2001. Puerto Rican Karst - A Vital Resource. U.S.
Department of Agriculture Forest Service General Technical Report WO-65,
2001.

Monroe W.H. 1976. The karst landforms of Puerto Rico. U.S. Geological
Survey Professional Paper.

Ospina-Sánchez C., F. Soto-Adames, and G. González. 2020. Checklist and
distribution of Collembola from Greater Puerto Rico. \emph{Biodiversity
Data Journal} 8:1--33

Pavan, A.C. 2023. \emph{Pteronotus portoricensis} . In: A. Kurta and A.
Rodriguez-Duran (Eds.), \emph{Bats of the West Indies, A Natural History
and Field Guide} , pp. 160--163. Cornell University Press, Ithaca.

Pérez-Reyes O., T.A. Crowl, P.J. Hernández-García, R. Ledesma-Fusté,
F.A. Villar-Fornes, and A.P. Covich. 2013. Freshwater decapods of Puerto
Rico: a checklist and reports of new localities. \emph{Zootaxa} 3717(3):
329--344.

Pinto B.J., S.E. Keating, S.V. Nielsen, D.P. Scantlebury, J.D. Daza, and
T. Gamble. 2022. Chromosome-level genome assembly reveals dynamic sex
chromosomes in neotropical leaf-litter geckos
(Sphaerodactylidae:\emph{Sphaerodactylus} ). \emph{Journal of Heredity}
113:272--287.

Pinto B.J., J. Titus-McQuillan, J.D. Daza, and T. Gamble. 2019.
Persistence of a geographically-stable hybrid zone in Puerto Rican dwarf
geckos. \emph{Journal of Heredity} 110:523--534.

Puente-Rolón, A.R. 1999. Foraging behavior, home range, movements,
activity patterns and habitat characterization of the Puerto Rican boa
(\emph{Epicrates inornatus} ) at Mata de Platano Reserve in Arecibo,
Puerto Rico. Doctoral dissertation, University of Puerto Rico, Mayaguez
(Puerto Rico).

Ramírez A., N. Maldonado-Benítez, A. Mariani-Ríos, and J.
Figueroa-Santiago. 2020. Dragonflies and damselflies (Odonata) from
Puerto Rico: a checklist with notes on distribution and
habitat.\emph{PeerJ} 8:e9711.

Reynolds R.G., A.H. Miller, A. Ríos-Franceschi, C.A. Huffine, J.
Fredette, N.F. Angeli, S.I. Vega-Castillo, L.J. Revell, and A.R.
Puente-Rolón. 2021. Molecular phylogeny of Puerto Rico Bank dwarf geckos
(Squamata: Sphaerodactylidae: \emph{Sphaerodactylus} ). \emph{bioRxiv} ,
https://doi.org/10.1101/2021.03.23.436310.

Rivero J.A. 1998. Los Anfibios y Reptiles de Puerto Rico - The
Amphibians and Reptiles of Puerto Rico. University of Puerto Rico Press,
San Juan.

Rodda G.H., G. Perry, R.J. Rondeau, and J. Lazell. 2001. The densest
terrestrial vertebrate. \emph{Journal of Tropical Ecology} 17:331--338.

Rodríguez-Durán, A. and J.A. Soto-Centeno. 2003. Temperature selection
by tropical bats roosting in caves. \emph{Journal of Thermal
Biology}28:465--468.

Soto-Centeno, J.A., and C.A. Calderón-Acevedo. 2023. Global Change and
the Conservation of Caribbean Bat Communities. In A. Kurta \& A.
Rodríguez-Durán (eds.), \emph{Bats of the West Indies, A Natural History
and Field Guide} (pp. 72--86). Cornell University Press, Ithaca.

Uetz, P., P. Freed, R. Aguilar, F. Reyes, J, Kudera, and J. Ho\selectlanguage{polish}š\selectlanguage{english}ek
(eds.). 2023. The Reptile Database, http://www.reptile-database.org,
accessed 1 March 2023.

V\selectlanguage{ngerman}élez Jr. Jr. M.J. 1967. Checklist of the terrestrial and freshwater
Decapoda of Puerto Rico. \emph{Caribbean Journal of Science} 7:41--44.

Wilson, A.K. 2023a. \emph{Mormoops blainvillei} . In: A. Kurta and A.
Rodriguez-Duran (Eds.), \emph{Bats of the West Indies, A Natural History
and Field Guide} , pp. 139--143. Cornell University Press, Ithaca.

Wilson, A.K. (2023b). \emph{Pteronotus quadridens} . In: A. Kurta and A.
Rodriguez-Duran (eds.), \emph{Bats of the West Indies, A Natural History
and Field Guide} (pp. 167--171). Cornell University Press, Ithaca.

Wogan G.O.U., M.L. Yuan, D.L. Mahler, and I.J Wang. 2020. Genome\selectlanguage{english}-wide
epigenetic isolation by environment in a widespread \emph{Anolis}lizard.
\emph{Molecular Ecology} 29:40--55.

\textbf{Author contribution}

The authors of this manuscript provided the following contributions to
this work: Justin M. Bernstein: Conceptualization (Lead), Data curation
(Equal), Formal analysis (Equal), Investigation (Equal), Methodology
(Equal), Resources (Supporting), Visualization (Lead), Writing --
original draft (Lead), Writing -- review \& editing (Equal). Camilo
Andres Calderon-Acevedo: Investigation (Equal), Methodology (Equal),
Writing -- review \& editing (Equal). Pedro Ivo Monico: Investigation
(Equal), Methodology (Equal), Writing -- review \& editing (Supporting).
Lazaro Willian Vinola-Lopez: Investigation (Equal), Methodology (Equal),
Writing -- review \& editing (Supporting). J. Angel Soto-Centeno: Data
curation (Equal), Funding acquisition (Lead), Investigation (Equal),
Methodology (Equal), Resources (Lead), Supervision (Equal),
Visualization (Supporting), Writing -- original draft (Supporting),
Writing -- review \& editing (Equal).

\selectlanguage{english}
\FloatBarrier
\end{document}

