\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,greek,polish,english]{babel}



\usepackage{float}








\begin{document}

\title{Craniodental traits and interaction in the bite force of tropical
frugivore bats (Phyllostomidae: Stenodermatinae)}



\author[1]{Leidy García Herrera}%
\author[1]{Leidy  Ramirez Francel }%
\author[1]{Giovany Guevara}%
\author[1]{Gladys  Reinoso Florez }%
\author[1]{Alfonso Sánchez Hernández}%
\author[2]{Burton Lim}%
\author[1]{Sergio  Losada prado }%
\affil[1]{Universidad del Tolima}%
\affil[2]{Royal Ontario Museum}%


\vspace{-1em}



  
  \date{May 24, 2024}


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





\selectlanguage{english}
\begin{abstract}
1. Phyllostomid bats exhibit great diversity in skull size and
morphology that reflects the degree of resource division and ecological
overlap in the group. In particular, Stenodermatinae has high
morphological diversification associated with cranial and mandibular
traits that is associated with the ability to consume the full range of
available fruits (soft and hard). In terms of morphology, performance
(bite force) appears to play an important role in niche partitioning
among bat species, however, very few studies have confirmed these
relationships using functional cranial traits. 2. Here, we analyzed
craniodental traits and their relationship to the bite force in 308
specimens distributed in seven species of stenodermatine bats with two
foraging types: nomadic and sedentary frugivorous bats. We evaluated 19
functional traits of the skull and jaw related to feeding and bite force
in live animals by correcting bite force with body size. We used a GLM
model and post hoc tests to determine possible relationships and
differences between cranial traits, species, and sex. 3. The results
showed that there is significant interspecific variation between
stenodermatines that are nomadic and sedentary. The greatest variation
in bite force within species was explained by the mandibular toothrow
length (MANDL) between sexes, which was greater in females. The larger
species of Artibeus, together with Platyrrhinus helleri, Uroderma
convexum and Sturnira giannae, which have a greater length of the skull,
condylo-incisor, condylo-canine, mandibular toothrow and height of the
coronoid, exhibit greater bite force. By contrast, the smaller species
A. anderseni and A. phaeotis have short skulls and the lowest values of
bite force, which suggests that the size of the skull confers a
biomechanical advantage. 4. Our results highlight the usefulness of
analyzing functional traits related to feeding to establish the
performance of bats in terms of the bite force.%
\end{abstract}%



\sloppy


\textbf{Craniodental traits and interaction in the bite force of
tropical frugivore bats (Phyllostomidae: Stenodermatinae)}

Leidy Viviana Garc\selectlanguage{ngerman}ía Herrera\textsuperscript{1*}, Leidy Azucena
Ramírez-Fráncel\textsuperscript{1}, Giovany Guevara\textsuperscript{2},
Gladys Reinoso-Flórez\textsuperscript{2}, Alfonso
Sánchez-Hernández\textsuperscript{3} Burton K. Lim\textsuperscript{4},
Sergio Losada-Prado\textsuperscript{2}

\textsuperscript{1}Programa de Doctorado en Ciencias Biológicas \& Grupo
de Investigación en Zoología (GIZ), Facultad de Ciencias, Universidad
del Tolima, Altos de Santa Elena, Ibagué, Colombia.

\textsuperscript{2}Departamento de Biología \& Grupo de Investigación en
Zoología (GIZ), Facultad de Ciencias, Universidad del Tolima, Altos de
Santa Elena, Ibagué, Colombia.

\textsuperscript{3}Departamento de Matemáticas y Estadística, Facultad
de Ciencias, Universidad del Tolima, Altos de Santa Elena, Ibagué,
Colombia.

\textsuperscript{4}Department of Natural History, Royal Ontario Museum,
100 Queen's Park, Toronto, Ontario M5S 2C6, Canada.

*Corresponding author\emph{lvgarcia@ut.edu.co}

\textbf{Abstract}

\begin{enumerate}
\tightlist
\item
  Phyllostomid bats exhibit great diversity in skull size and morphology
  that reflects the degree of resource division and ecological overlap
  in the group. In particular, Stenodermatinae has high morphological
  diversification associated with cranial and mandibular traits that is
  associated with the ability to consume the full range of available
  fruits (soft and hard). In terms of morphology, performance (bite
  force) appears to play an important role in niche partitioning among
  bat species, however, very few studies have confirmed these
  relationships using functional cranial traits.
\item
  Here, we analyzed craniodental traits and their relationship to the
  bite force in 309 specimens distributed in seven species of
  stenodermatine bats with two foraging types: nomadic and sedentary
  frugivorous bats. We evaluated 19 functional traits of the skull and
  jaw related to feeding and bite force in live animals by correcting
  bite force with body size. We used a GLM model and \emph{post
  hoc}tests to determine possible relationships and differences between
  cranial traits, species, and sex.
\item
  The results showed that there is significant interspecific variation
  between stenodermatines that are nomadic and sedentary. The greatest
  variation in bite force within species was explained by the mandibular
  toothrow length (MANDL) between sexes, which was greater in females.
  The larger species of \emph{Artibeus} , together
  with\emph{Platyrrhinus helleri} , \emph{Uroderma convexum}
  and\emph{Sturnira giannae} , which have a greater length of the skull,
  condylo-incisor, condylo-canine, mandibular toothrow and height of the
  coronoid, exhibit greater bite force. By contrast, the smaller
  species\emph{A. anderseni} and \emph{A. phaeotis} have short skulls
  and the lowest values of bite force, which suggests that the size of
  the skull confers a biomechanical advantage.
\item
  Our results highlight the usefulness of analyzing functional traits
  related to feeding to establish the performance of bats in terms of
  the bite force.
\end{enumerate}

KEYWORDS

Ecomorphology, frugivores, morphometry, Stenodermatinae, skull, trophic
performance.

\textbf{1.} \textbf{INTRODUCTION}

The skull of vertebrates is a complex assembly that is closely related
to the resource collection, food processing, behavior, and ecology of
the species (Bels \& Herrel, 2019). Examining the patterns and
mechanisms that lead to cranial variation, including shape and traits,
enables an understanding of the morphology, ecology, and general fitness
of animals (Santana, Dumont, \& Davis, 2010). Bats have variation in
cranial morphology attributed to evolutionary processes of ecological
specialization, which result in a niche division between
ecomorphologically similar species (Santana, Grosse, \& Dumont, 2012).
This variation responds mainly to functional requirements related to
nutritional performance and the sensory system (Thiagavel et al., 2018).

Among chiropterans, New World leaf-nosed bats (Phyllostomidae)
represents one of the largest and most morphologically diverse mammal
families (Rossoni et al., 2017). Ecological diversification in
Phyllostomidae is related to bite performance and mechanical demands of
different diets, including frugivores, insectivores, nectarivores,
carnivores, and sanguinivores (Dumont, 2007, Nogueira, Peracchi, \&
Monteiro, 2009; Manhães, Nogueira, \& Monteiro, 2017). Dietary
differences require specific mechanical modifications, including
variation in the rostral length and height of the skull (Santana,
Dumont, \& Davis, 2010). However, there is a lack of understanding about
the patterns in the variation of the shape and performance
(López-Aguirre \& Pérez-Torres, 2015). The remarkable specializations
seen in these bats provide a unique opportunity for studying the
relationship between cranial morphology, feeding performance, and
dietary ecology (Aguirre, Herrel, Van Damme, \& Matthysen, 2002; Rossoni
et al., 2017).

The bite force is a biomechanical trait that determines various vital
functions, such as feeding, mating, defense, and competition, and is
closely related to body size and feeding performance (Santana, Dumont,
\& Davis, 2010). Therefore, an increase in bite force allows for a
broader spectrum of food available (Santana \& Dumont, 2009).
Phylostomid exhibit bite forces associated with the size and hardness of
food (Aguirre, Herrel, Van Damme, \& Matthysen, 2003), resulting in
strong morphological heterogeneity due to the particularities of the
diet (Dumont \& O'Neil, 2004).

Although various studies have addressed the relationships of
craniodental morphology, bite force and diet (e.g., Aguirre, Herrel, Van
Damme, \& Matthysen, 2002; Dumont, Herrel, Medellín, Vargas-Contreras,
\& Santana, 2009: Santana, Grosse, \& Dumont, 2012; Santana \& Miller
2016), the evaluation of these variables has been carried out
separately, evaluating each variable independently (Shi et al., 2020).
Likewise, the information related to the craniodental traits responsible
for generating bite forces in bats is limited without much attention to
the functionally relevant components of the craniodental system and the
musculature of the jaw (Herrel, Smet, Aguirre, \& Aerts, 2008). The
specific hardness of food and the bite force required to process food
play an important role in the division of resources within frugivore bat
communities (Dumont, 2007). According to Soriano (2000) and Giannini \&
Kalko (2004), fruit bats can be categorized as nomadic or sedentary,
with preference for a specific type of plant, morphological diversity
reflected in the physical properties of the fruits, and a tendency of
specialization towards a specific fruit or group of fruits (Santana,
Dumont, \& Davis, 2010; Santana, Grosse, \& Dumont, 2012).

Frugivorous bats provide an excellent model to study the relationship
between craniodental morphology and bite force because they consume both
hard and soft fruits, the result of adaptive pressures related to
mandibular morphology (see Murillo-García, \& De la Vega, 2018). We
selected the members of the family Phyllostomidae due to the fact that
they have the highest diversity of mammals with more than 70 species
that are found sympatrically and serve as nocturnal predators,
pollinators and predominant seed dispersers (Giannini \& Kalko, 2004;
Reid et al., 2015), and within this family, the Stenodermatinae
subfamily, contains \textgreater{} 43\% of all described New World
leaf-nosed bat species (Shipley \& Twining, 2020) and in central
Colombia, department of Tolima, these chiropterans represent near the
13\% in the sampled areas (see García-Herrera, Ramírez-Fráncel, \&
Flórez, 2019).

Our objective was to identify the functional traits associated with bite
force in seven representative species of the Stenodermatinae subfamily
occurring in Colombia. We hypothesize that large species and small-sized
nomadic frugivorous species with narrow faces, wide palates and shorter
length between molars will have a greater biomechanical advantage of
stronger bite force in relation to sedentary frugivorous bats.

\textbf{2.} \textbf{MATERIALS AND METHODS}

\textbf{2.1. \emph{Samples and classification}}

From February 2019 to January 2020, we registered bite force and skull
morphology data of seven species of Stenodermatinae fruit bats (Appendix
I). These bats were divided into two categories: (i) nomadic
frugivores:\emph{Artibeus anderseni, A. phaeotis, A. lituratus, A.
planirostris, Platyrrhinus helleri, Uroderma convexum} ; (ii) sedentary
frugivores:

\emph{Sturnira giannae.}

\textbf{2.2. \emph{Field collection and laboratory analysis}}

We follow two procedures for collecting our data: i) fieldwork in
selected areas of the Colombian tropical dry forest-TDF in the
department of Tolima (Figure 1, red triangle marks) using conventional
methodology, i.e., mist-nets placed along trails within forest areas, at
the edge of forest remnants, and near waterbodies. Each sampling night
consisted of four standard size mist nets (12 x 2.5m), eight nets (6 x
2.5m) in the sub-canopy and a Triple High net (30 X 7 m), with a
sampling intensity of 36.288 m\textsuperscript{2} nets/h, corresponding
to 864 h in 144 nights. The captured bats were handled according to the
American Society of Mammalogists guidelines for the use of wild animals
for research purposes (Sikes \& Animal Care and Use Committee of the
American Society of Mammalogists, 2016). After capture, the age, sex,
and reproductive status were evaluated, and only adult males and adult
non-pregnant, non-lactating females were used for the measurements. Age
and sex were identified based on the degree of ossification of the coat
and tibia. Reproductive status in females was determined by examining
the nipples and palpitation of the abdomen. Then, bats were put into
cloth bags and transferred to measure the bite force using a portable
digital fruit hardness tester (Lutron, Indian) with a capacity of 196.10
Newton and precision ± 0.05.

We follow the method of Freeman \& Lemen (2008), and bite force was
recorded at the molars, measurements were repeated five times for each
bat with an inter-trial interval of at least 5\selectlanguage{english} min. The maximum value of
the five measurements was considered as the maximum bite force produced
by that individual. The bite force of the species was calculated by
averaging the maximum bite force of each individual. The captured
specimens were used for direct measurements of the forearm length, body
mass, then they were euthanized and handled for craniodental morphometry
(Figure 2; see Table S1), and entry to biological collection of the
University of Tolima CZUT-M (Ibague, Colombia). ii) The second procedure
consisted of measurements of 16 craniodental traits and two body traits
(Table 1, Figure 2) obtained of voucher specimens from the Zoological
Collection of the University of Tolima (CZUT; Ibague, Colombia), Museo
Javeriano de Historia Natural ''Lorenzo Uribe, SJ'' (MPUJ; Bogota,
Colombia) and Royal Ontario Museum (ROM; Ontario, Canada). The revised
specimens from these biological collections belong to 24 localities in
Colombia (Figure 1; Appendix I). We verified that all adult specimens
according to the ossification of the growth plates of the epiphysis of
the phalanges of the fingers (Dietz et al., 2007).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

\textbf{FIGURE 1} Geographic distribution of the locations of the
Stenodermatinae bat samples used for the biometric analyzes. Triangle
black marks represent sampling points where capture of Colombian
Stenodermatinae bats was carried out in the department of Tolima; other
circle marks represent records of other sampling sites obtained from
voucher specimens (see text for more details).

\textbf{TABLE 1} The craniodental, mandibular and external measurements
used as functional traits in this study.\selectlanguage{english}
\begin{longtable}[]{@{}llllll@{}}
\toprule
Variable & Main trait & Description & Abbreviation & Unit & Key
supporting reference\tabularnewline
\midrule
\endhead
Bite force & Head & Maximum bite force produced by molars & BF & N/g &
Shi et al., 2020\tabularnewline
Forearm length & Body & Distance from the olecranon process to anterior
surface of carpals in the folded wing & FA & mm & Garc\selectlanguage{ngerman}ía-Morales et al.,
2016\tabularnewline
Mass & Body & Physical quantity that allows to indicate the quantity of
matter contained in a body. & MAS & g & Cisneros et al.
(2014)\tabularnewline
Greatest length of skull & Head & Distance from the posterior-most point
of the occiput to the anterior-most point of the premaxilla, including
incisors & GLS & mm & Cisneros et al. (2014),\tabularnewline
Condyloincisive length & Head & Distance between a line connecting the
posterior-most margins of the occipital condyles and the anterior most
surface of the upper incisors & CIL & mm & Cisneros et al.
(2014)\tabularnewline
Condylocanine length & Head & Distance between a line connecting the
posterior-most margins of the occipital condyles and a line connecting
the anterior-most surface of the upper canines. & CCL & mm & Cisneros et
al. (2014)\tabularnewline
Braincase breadth & Head & Breadth of the braincase, excluding mastoid
and paraoccipital processes & BB & mm & Cisneros et al.
(2014)\tabularnewline
Zygomatic breadth & Head & Breadth across the zygomatic arches & ZB & mm
& Cisneros et al. (2014)\tabularnewline
Postorbital breadth & Head & Breadth at the postorbital constriction &
PB & mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la vega, 2018\tabularnewline
Mastoid breadth & Head & Greatest breadth across the mastoid region & MB
& mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la vega, 2018\tabularnewline
Palatal length & Head & Distance between the posterior palatal notch and
the anterior border of the incisive alveolus & PL & mm & Murillo-Garc\selectlanguage{ngerman}ía
\& De la vega, 2018\tabularnewline
Maxillary toothrow length & Head & Distance from the anterior-most
surface of the upper canine to the posterior-most surface of the crown
of M3 & MTRL & mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la vega, 2018\tabularnewline
Width at M1 & Head & Greatest width of palate across M1s & M1M1 & mm &
Murillo-Garc\selectlanguage{ngerman}ía \& De la vega, 2018\tabularnewline
Width at M2 & Head & Greatest width of palate across M2s & M2M2 & mm &
Murillo-Garc\selectlanguage{ngerman}ía \& De la vega, 2018\tabularnewline
Palatal width at canines & Head & Least width across palate between
alveoli of upper canines & C-C & mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la vega,
2018\tabularnewline
Dentary length & Head & Length between midpoint of condyle to
anterior-most point of dentary & DEL & mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la
vega, 2018\tabularnewline
Mandibular toothrow length & Head & Distance from the anterior-most
surface of the lower canine to the posterior-most surface of m3 & MANDL
& mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la vega, 2018\tabularnewline
Coronoid height & Head & Perpendicular height from ventral margin of
mandible to tip of coronoid process & COH & mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la
vega, 2018\tabularnewline
Width at mandibular condyles & Head & Greatest width between inner
margins of mandibular condyles & WMC & mm & Murillo-Garc\selectlanguage{ngerman}ía \& De la
vega, 2018\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{ngerman}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

\textbf{FIGURE 2} Lateral and dorsal views of the cranium and mandible
(\emph{Platyrrhinus helleri} , adult male), and ventral view of the
cranium, measurements used in the craniodental morphometry.
Abbreviations: GLS, greatest length of skull; CIL, condyloincisive
length; CCL, condylocanine length; BB braincase breadth; ZB, zygomatic
breadth; PB, postorbital breadth; C--C, palatal width at canines; MB,
mastoid breadth; PL, palatal length; MTRL, maxillary toothrow length;
M1--M1, width at M1; M2--M2, width at M2; DENL, dentary length; MANDL,
mandibular toothrow length; COH, coronoid height; WMC, width at
mandibular condyles.

\textbf{2.3.} \textbf{Statistical analyses}

Data analyses were carried out by including both fieldwork and voucher
specimen's data. A linear regression was performed to assess the effects
of body parameters on the changes in bite force. In the models, the
averages of the bite force and body size (forearm length, greatest
length of skull) and mass of each species were used. Ln (y) = \selectlanguage{greek}β\selectlanguage{english}0 + \selectlanguage{greek}β\selectlanguage{english}1x1
+ \selectlanguage{greek}β\selectlanguage{english}2x2 + \selectlanguage{greek}ε, \selectlanguage{english}where y corresponds to the bite force, x1 and x2 are the
length of the forearm and greatest length of skull, respectively. \selectlanguage{greek}β\selectlanguage{english}0 is
the intercept, \selectlanguage{greek}β\selectlanguage{english}1, \selectlanguage{greek}β\selectlanguage{english}2 are the regression coefficients (for x1 and x2),
and \selectlanguage{greek}ε \selectlanguage{english}is the random error.

In order to analyze the intra and interspecific morphological variation,
the mean \selectlanguage{ngerman}± SD was calculated for all morphometric variables and to
determine sexual dimorphism. The assumptions of normality and
homoscedasticity were corroborated with the Shapiro-Wilk and Levene's
test, respectively. The test results showed that our data fit a normal
distribution, and accomplish the equality of variances, thus we used a
two-way analysis of variance (ANOVA) to test differences between species
and sexes. Post-hoc comparisons between sexes within species were
performed to identify species with significant dimorphism.

To explore the differences in skull morphology and body traits among
different species and sexes, we performed a Principal Component
Analysis- PCA based on a correlation matrix. A threshold value of
\selectlanguage{greek}λ\selectlanguage{english}\textgreater{} 1 was used to determine the relevant traits that explain
most of the observed variance. Based on the preliminary results of the
PCA, the variables forearm length and mass were excluded because they
presented a correlation close to one, while the other variables (17)
were used for the Canonical Variate Analysis- CVA. A CVA was carried
out, in order to establish the largest axes of discrimination between
the groups identified \emph{a priori} ; find the linear combinations of
the starting variables with maximum discriminating power between the
groups; test whether the means of these groups along these axes are
significantly different to find an ordering of the groups of individuals
each represented by the vector of the means in all the variables; and to
study the dimensionality of the data.

In order to determine the existence of significant variation between
species and morphological traits, an analysis of variance was carried
out using a generalized linear model (GLM). We used the bite force as
the response variable and the cranial and body traits as covariates. The
model used was: Yij\selectlanguage{polish}ķ \selectlanguage{english}= \selectlanguage{greek}µ \selectlanguage{english}+ \selectlanguage{polish}Ţ\selectlanguage{english}i + \selectlanguage{greek}δ\selectlanguage{english}j + \selectlanguage{greek}Ω\selectlanguage{polish}ķ \selectlanguage{english}+ \selectlanguage{greek}ε\selectlanguage{english}ij\selectlanguage{polish}ķ, \selectlanguage{english}where Yij\selectlanguage{polish}ķ \selectlanguage{english}represents
the response of the bite force at the jth level of sex and ith species;
\selectlanguage{greek}µ \selectlanguage{english}general average, \selectlanguage{polish}Ţ\selectlanguage{english}i effect produced by the i-th species, \selectlanguage{greek}δ\selectlanguage{english}j effect
produced by the j-th sex, \selectlanguage{greek}Ω\selectlanguage{polish}ķ \selectlanguage{english}effect due to the R-th trait and \selectlanguage{greek}ε\selectlanguage{english}ij\selectlanguage{polish}ķ \selectlanguage{english}the
random error. For detecting the masked variability, an intuitive and
qualitative procedure based on graphical representation was used, then
we performed \emph{post hoc} tests using the Fisher's Least Significant
Difference (LSD) pairwise comparison procedure. We set the statistical
significance for all tests of P ? 0.05. All analyzes were performed in R
3.5.3 (R Core Team 2019).

\textbf{3. RESULTS}

Comparisons of morphological traits between males and females showed
significant differences by species (F = 72.04, d.f. = 92, p
\textless{}0.001), sex (F = 4.22, d.f. = 17, p \textless{}0.001) and
their interaction (F = 2.8, d.f. = 94, p \textless{}0.001, Table 2). Los
rasgos craneales que presentaron mayor variabilidad fueron la longitud
dentaria (DENL), la longitud de los dientes de la mand\selectlanguage{ngerman}íbula (MANDL) con
un coeficiente de variación de 47.2\% y 18.50\% respectivamente,
mientras que los demás rasgos morfológicos presentaron una menor
variación baja en 10\%.

The PCA had the first two axes explaining 91.8\% of all the variance.
The first component (PC1) explained 87.4\% of the total variance and is
determined by the width across the upper first molars (M1-M1), zygomatic
breadth (ZB), forearm length (AB), dentary length (DENL) and
condylo-incisor length (CIL). The second component (PC2), explained
4.4\% of the variance, and was associated with the traits of maxillary
toothrow length (MTRL), width between the cingulate of the upper canines
(C-C), greater length of the skull (GLS), postorbital amplitude (PB),
braincase breadth; (BB), and the bite force (BF) (Figure 3).

The CVA had two main groups containing species of small-sized nomadic
frugivorous and sedentary frugivorous bats; and large-sized nomadic
frugivorous bats. Five subgroups were identified for \emph{A.
anderseni}and \emph{A. phaeotis} ; \emph{S. giannae} ; \emph{U.
convexum} and\emph{Platyrrhinus helleri} ; \emph{A. planirostris} ; and
\emph{Artibeus lituratus} (Figure 3). The traits that most contributed
to the discrimination of the species in the CVA were MANDL, PL and WMC,
which explained the greater morphological variation between species
(Figure 4).

\textbf{Table 2.} Species, sample size (n) bats used to investigate bite
force and jaw-skull shape relationships.\selectlanguage{english}
\begin{longtable}[]{@{}llllllllllllllllllllllllllll@{}}
\toprule
\textbf{Species} & \textbf{Sex} & \textbf{n} & \textbf{GLS} &
\textbf{CIL} & \textbf{CCL} & \textbf{ZB} & \textbf{BB} & \textbf{PB} &
\textbf{MB} & \textbf{PL} & \textbf{MTRL} & \textbf{M1-M1} &
\textbf{M1-M1} & \textbf{M2-M2} & \textbf{M2-M2} & \textbf{C-C} &
\textbf{C-C} & \textbf{DENL} & \textbf{DENL} & \textbf{MANDL} &
\textbf{MANDL} & \textbf{COH} & \textbf{COH} & \textbf{WMC} &
\textbf{WMC} & \textbf{BS} & \textbf{BS}\tabularnewline
\midrule
\endhead
\emph{A. anderseni} & \emph{} & 20 & 18.26 (0.46) & 13.98 (1.50) & 13.9
(1.40) & 10.56 (0.57) & 8.87 (0.39) & 4.60 (0.37) & 8.73 (0.46) & 7.6
(0.43) & 5.38 (0.28) & 7.74 (0.41) & 7.74 (0.41) & 7.39 (0.18) & 7.39
(0.18) & 4.29 (0.44) & 4.29 (0.44) & 10.9 (0.54) & 10.9 (0.54) & 5.90
(0.40) & 5.90 (0.40) & 4.42 (0.26) & 4.42 (0.26) & 5.92 (0.58) & 5.92
(0.58) & 35.50 (4.53) & 35.50 (4.53)\tabularnewline
& \emph{} & 21 & 18.11 (1.83) & 14.21 (0.58) & 14.12 (0.39) & 10.71
(0.58) & 8.88 (0.40) & 4.54 (0.47) & 9.11 (0.46) & 7.54 (0.38) & 5.41
(0.28) & 7.59 (0.50) & 7.59 (0.50) & 7.49 (0.16) & 7.49 (0.16) & 4.15
(0.33) & 4.15 (0.33) & 10.93 (0.47) & 10.93 (0.47) & 5.87 (0.33) & 5.87
(0.33) & 4.52 (0.27) & 4.52 (0.27) & 5.68 (0.55) & 5.68 (0.55) & 55
(5.94) & 55 (5.94)\tabularnewline
\emph{A. lituratus} & \emph{} & 33 & 30.69 (0.74) & 24.35 (062) & 23.76
(0.56) & 18.34 (0.44) & 14.12 (0.35) & 6.58 (0.51) & 14.58 (0.63) &
14.63 (0.43) & 10.51 (0.51) & 12.78 (0.29) & 12.78 (0.29) & 12.74 (0.37)
& 12.74 (0.37) & 6.41 (0.25) & 6.41 (0.25) & 20.41 (0.58) & 20.41 (0.58)
& 12.65 (0.34) & 12.65 (0.34) & 9.64 (0.30) & 9.64 (0.30) & 9.58 (0.43)
& 9.58 (0.43) & 61.31 (10.48) & 61.31 (10.48)\tabularnewline
& \emph{} & 31 & 31.34 (0.55) & 25.06 (0.67) & 24.44 (4.16) & 18.62
(0.51) & 13.93 (0.56) & 7.12 (0.38) & 14.72 (0.57) & 15.13 (0.43) & 11.1
(0.41) & 12.77 (0.43) & 12.77 (0.43) & 12.87 (0.50) & 12.87 (0.50) &
6.33 (0.37) & 6.33 (0.37) & 20.68 (0.81) & 20.68 (0.81) & 12.59 (0.48) &
12.59 (0.48) & 9.56 (0.49) & 9.56 (0.49) & 9.87 (0.38) & 9.87 (0.38) &
77.5 (4.24) & 77.5 (4.24)\tabularnewline
\emph{A. planirostris} & \emph{} & 26 & 27.87 (4.97) & 22.58 (0.77) &
22.07 (0.81) & 16.78 (0.73) & 13.32 (0.56) & 6.63 (0.46) & 12.63 (0.53)
& 13.54 (0.75) & 10.41 (0.53) & 11.46 (0.59) & 11.46 (0.59) & 10.65
(0.83) & 10.65 (0.83) & 5.74 (0.39) & 5.74 (0.39) & 19.26 (0.70) & 19.26
(0.70) & 11.15 (0.72) & 11.15 (0.72) & 7.89 (0.56) & 7.89 (0.56) & 8.42
(0.38) & 8.42 (0.38) & 76.15 (4.58) & 76.15 (4.58)\tabularnewline
& \emph{} & 38 & 28.12 (0.85) & 22.71 (0.83) & 22.21 (0.77) & 17.19
(0.40) & 13.47 (0.70) & 6.98 (0.34) & 13.19 (0.38) & 13.88 (0.47) &
10.61 (0.65) & 11.53 (0.55) & 11.53 (0.55) & 11.50 (0.81) & 11.50 (0.81)
& 6.28 (0.32) & 6.28 (0.32) & 19.32 (0.34) & 19.32 (0.34) & 11.49 (1.21)
& 11.49 (1.21) & 8.48 (0.45) & 8.48 (0.45) & 8.56 (0.35) & 8.56 (0.35) &
93.51 (3.00) & 93.51 (3.00)\tabularnewline
\emph{A. phaeotis} & \emph{} & 17 & 18.68 (0.64) & 15.11 (0.78) & 14.89
(1.00) & 11.03 (0.48) & 9.27 (0.47) & 4.33 (0.32) & 8.56 (0.55) & 7.65
(0.77) & 5.69 (0.55) & 7.5 (0.51) & 7.5 (0.51) & 7.34 (0.40) & 7.34
(0.40) & 4.11 (0.25) & 4.11 (0.25) & 11.14 (0.57) & 11.14 (0.57) & 6.14
(0.64) & 6.14 (0.64) & 4.35 (0.43) & 4.35 (0.43) & 6.24 (0.59) & 6.24
(0.59) & 33.75 (4.77) & 33.75 (4.77)\tabularnewline
& \emph{} & 11 & 18.33 (0.77) & 14.21 (0.58) & 14.24 (0.52) & 10.71
(0.41) & 8.95 (0.39) & 4.45 (0.21) & 9.35 (0.42) & 7.66 (0.68) & 5.54
(0.42) & 8.02 (0.52) & 8.02 (0.52) & 7.45 (0.21) & 7.45 (0.21) & 4.35
(0.33) & 4.35 (0.33) & 11.36 (0.55) & 11.36 (0.55) & 6.02 (0.51) & 6.02
(0.51) & 4.69 (0.25) & 4.69 (0.25) & 5.12 (0.85) & 5.12 (0.85) & 62.5
(3.13) & 62.5 (3.13)\tabularnewline
\emph{P. helleri} & \emph{} & 9 & 21.25 (0.48) & 17.58 (0.40) & 17.21
(0.45) & 10.44 (0.45) & 8.58 (0.48) & 4.89 (0.37) & 8.81 (0.29) & 9.73
(0.44) & 7.55 (0.37) & 7.92 (0.35) & 7.92 (0.35) & 7.69 (0.51) & 7.69
(0.51) & 3.85 (0.28) & 3.85 (0.28) & 13.65 (0.44) & 13.65 (0.44) & 8.32
(0.64) & 8.32 (0.64) & 4.96 (0.43) & 4.96 (0.43) & 5.96 (0.44) & 5.96
(0.44) & 44.5 (9.50) & 44.5 (9.50)\tabularnewline
& \emph{} & 20 & 21.88 (0.68) & 17.99 (0.60) & 17.33 (0.52) & 11.16
(0.47) & 9.5 (0.62) & 5.24 (0.32) & 9.15 (0.38) & 9.99 (0.33) & 7.72
(0.45) & 8.02 (0.31) & 8.02 (0.31) & 8.12 (0.41) & 8.12 (0.41) & 3.84
(0.39) & 3.84 (0.39) & 13.74 (0.63) & 13.74 (0.63) & 8.64 (0.41) & 8.64
(0.41) & 4.60 (0.44) & 4.60 (0.44) & 6.13 (0.42) & 6.13 (0.42) & 72.5
(2.13) & 72.5 (2.13)\tabularnewline
\emph{U. convexum} & \emph{} & 19 & 22.12 (0.65) & 18.32 (0.67) & 17.56
(0.49) & 11.69 (0.52) & 9.40 (0.36) & 5.66 (0.65) & 9.25 (0.41) & 11.23
(0.56) & 7.85 (0.43) & 8.12 (0.62) & 8.12 (0.62) & 8.22 (0.44) & 8.22
(0.44) & 4.00 (0.53) & 4.00 (0.53) & 14.24 (0.28) & 14.24 (0.28) & 8.25
(0.49) & 8.25 (0.49) & 5.04 (0.55) & 5.04 (0.55) & 6.28 (0.27) & 6.28
(0.27) & 45.00 (6.25) & 45.00 (6.25)\tabularnewline
& \emph{} & 21 & 22.34 (0.44) & 17.58 (0.70) & 17.25 (0.54) & 11.67
(0.68) & 10.24 (0.48) & 5.32 (0.47) & 10.34 (0.70) & 11.39 (0.43) & 7.67
(0.42) & 8.16 (0.43) & 8.16 (0.43) & 7.69 (0.49) & 7.69 (0.49) & 3.87
(0.45) & 3.87 (0.45) & 13.67 (0.43) & 13.67 (0.43) & 8.25 (0.47) & 8.25
(0.47) & 4.69 (0.44) & 4.69 (0.44) & 6.19 (0.62) & 6.19 (0.62) & 79.25
(5.04) & 79.25 (5.04)\tabularnewline
\emph{S. giannae} & \emph{} & 13 & 22.25 (0.62) & 17.81 (0.69) & 17.22
(0.63) & 12.09 (0.34) & 10.45 (0.26) & 6.08 (0.37) & 10.38 (0.56) &
10.25 (0.53) & 6.52 (0.21) & 7.58 (0.28) & 7.58 (0.28) & 7.64 (0.27) &
7.64 (0.27) & 5.30 (0.44) & 5.30 (0.44) & 13.99 (0.53) & 13.99 (0.53) &
7.60 (0.34) & 7.60 (0.34) & 5.25 (0.36) & 5.25 (0.36) & 7.22 (0.48) &
7.22 (0.48) & 33.00 (2.24) & 33.00 (2.24)\tabularnewline
& \emph{} & 17 & 22.11 (0.49) & 17.65 (0.47) & 17.22 (0.41) & 12.26
(0.41) & 10.47 (0.36) & 6.25 (0.40) & 10.25 (0.28) & 10.24 (0.30) & 6.55
(0.52) & 7.59 (0.29) & 7.59 (0.29) & 7.58 (0.22) & 7.58 (0.22) & 5.16
(0.40) & 5.16 (0.40) & 14.01 (0.41) & 14.01 (0.41) & 7.86 (0.34) & 7.86
(0.34) & 5.47 (0.23) & 5.47 (0.23) & 7.25 (0.44) & 7.25 (0.44) & 58.00
(6.02) & 58.00 (6.02)\tabularnewline
Manova (Wilks) & & & \textbf{Value} & \textbf{Value} & F & \textbf{d.f.}
& P & & & & & & & & & & & & & & & & & & & &\tabularnewline
& Specie & Specie & 5.20e-05 & 5.20e-05 & 72.04 & 92 & \textless{}0.001
& & & & & & & & & & & & & & & & & & & &\tabularnewline
& Sex & Sex & 0.79 & 0.79 & 4.22 & 17 & \textless{}0.001 & & & & & & & &
& & & & & & & & & & & &\tabularnewline
& Specie*Sex & Specie*Sex & 0.39 & 0.39 & 2.8 & 94 & \textless{}0.001 &
& & & & & & & & & & & & & & & & & & &\tabularnewline
\bottomrule
\end{longtable}

* All measurements are presented in millimeters except for BS which are
presented in Newtons.

Values are described as mean (\selectlanguage{ngerman}± SD).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\textbf{FIGURE 3} Distribution of body and craniodental traits for seven
species of fruit bats based on a Principal Components Analysis (PCA).

The GLM determined significant differences both for species and for all
traits (F = 27.14; P \textless{}\textless{} 0.001) and variation between
sexes (F = 5.73, P = 0.017; Figure 5). All pairwise comparisons were
significant with P [?] 0.01.

\textbf{3.1 Variation in the bite force associated with craniodental
traits}

Most of the variation in the bite force data according to test LSD was
mainly explained for both males and females by the MANDL trait (t =
8.00, df = 12, P \textless{} 0.01; Figure 5). It was identified that 15
traits are significantly associated with a high bite force, including
greatest length of the skull, condylo-incisor length, condylo-canine
length, postorbital breadth, mastoid breadth, zygomatic arch breadth,
palatal length, mandibular toothrow length, and height of the coronoids
(Table 2).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

\textbf{FIGURE 4} Canonical variate analysis with the description of
body and craniodental traits and seven species of Stenodermatinae bats.
The five ellipses (groups) are highlighted.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

\textbf{FIGURE 5} Pairwise relationships of craniodental traits with
bite force for seven Colombian species of Stenodermatinae bats. Blue
squares correspond to males and yellow to females. Abbreviations as in
Table 1.

\textbf{3.2 Interspecific bite force variation}

The bite force varied between species; big-sized nomadic bats had the
highest bite force, followed by small-sized nomadic bats (\emph{P.
helleri} and \emph{U. convexum} ), and then by the only sedentary bat
(\emph{S. giannae} ). The lowest force was recorded by small-sized
nomadic bats of the species \emph{A. anderseni} and \emph{A. phaeotis}(P
\textless{} 0.001). Significant intersexual differences between species
were detected with the females registering a greater bite force compared
to the males (P \textless{} 0.04; Figure 6).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}

\textbf{FIGURE 6} Box plot showing differences in relative bite force
among bat species and sex. \emph{A. lit} : \emph{Artibeuslituratus. A.
pla} : \emph{A. planirostris. A. and} : \emph{A. anderseni. A. pha} :
\emph{A. phaeotis. U. con: Uroderma convexum} .\emph{P. hel} :
\emph{Platyrrhinus helleri} and \emph{S. gia: Sturnira giannae.}

Our results revealed that big-sized nomadic bats, \emph{A. lituratus} ,
and \emph{A. planirostris} differed from the remaining species, as
established by post-hoc tests (P \textless{}0.001). Furthermore, it was
established that \emph{A. anderseni} and \emph{A. phaeotis} are
significantly different from \emph{S. giannae} and \emph{U. convexum} ,
while these last two species are similar to \emph{P. helleri} and the
small species of \emph{Artibeus} (Table 3).

\emph{Differences between bite force between bat species} .

In this study, significant differences in skull morphology and bite
strength were observed between species. LSD post-hoc testing provided
clustering indicators, the first group consisted of the large nomadic
bats, \emph{A. lituratus} and \emph{A. planirostris} ; the second by a
nomadic species U. convexum; the third formed by the small
species\emph{P. helleri} , \emph{S. giannae} , \emph{A. pheotis} and the
fourth formed by \emph{A. anderseni} , species that share cranial and
corporal features in the third group (Table 4).

\textbf{Table 3.} Significance between pairs of species and relationship
of bite force with other traits.\selectlanguage{english}
\begin{longtable}[]{@{}llll@{}}
\toprule
Species & P (value significant) & Traits & P (value
significant)\tabularnewline
\midrule
\endhead
\emph{A. lituratus - A. phaeotis} & \textless{}0,001 & Bf - C-C &
\textless{}0,001\tabularnewline
\emph{A. lituratus - A. anderseni} & \textless{}0,001 & Bf - CCL &
\textless{}0,001\tabularnewline
\emph{A. lituratus - A. planirostris} & 0,486 & Bf - CIL &
\textless{}0,001\tabularnewline
\emph{A. lituratus - P. helleri} & \textless{}0,001 & Bf - COH &
\textless{}0,001\tabularnewline
\emph{A. lituratus - S. giannae} & \textless{}0,001 & Bf - DENL &
\textless{}0,001\tabularnewline
\emph{A. lituratus - U. convexum} & \textless{}0,001 & Bf - GLS &
\textless{}0,001\tabularnewline
\emph{A. phaeotis - A. planirostris} & \textless{}0,001 & Bf - M1-M1 &
\textless{}0,001\tabularnewline
\emph{A. phaeotis - A. anderseni} & 0,84 & Bf - M2-M2 &
\textless{}0,001\tabularnewline
\emph{A. phaeotis - P. helleri} & 0,13 & Bf - MANDL &
\textless{}0,001\tabularnewline
\emph{A. phaeotis - S. giannae} & 0,04 & Bf - MASA &
\textless{}0,001\tabularnewline
\emph{A. phaeotis - U. convexum} & 0,05 & Bf - MB &
\textless{}0,001\tabularnewline
\emph{A. planirostris - A. anderseni} & \textless{}0,001 & Bf - MTRL &
\textless{}0,001\tabularnewline
\emph{A. planirostris - P. helleri} & \textless{}0,001 & Bf - PB &
\textless{}0,001\tabularnewline
\emph{A. planirostris - S. giannae} & \textless{}0,001 & Bf - PL &
\textless{}0,001\tabularnewline
\emph{A. planirostris - U. convexum} & \textless{}0,001 & Bf - WMC &
\textless{}0,001\tabularnewline
\emph{A. anderseni - P. helleri} & 0,08 & Bf - ZB &
\textless{}0,001\tabularnewline
\emph{A. anderseni - S. giannae} & 0,08 & &\tabularnewline
\emph{A. anderseni - U. convexum} & 0,04 & &\tabularnewline
\emph{P. helleri - S. giannae} & 0,99 & &\tabularnewline
\emph{P. helleri - U. convexum} & 0,73 & &\tabularnewline
\emph{S. giannae -U. convexum} & 0,73 & &\tabularnewline
\bottomrule
\end{longtable}

\textbf{Table 4.} Post hoc test Least Significant Difference (LSD)
pairwise comparison of bat species and body / cranial traits.\selectlanguage{english}
\begin{longtable}[]{@{}ll@{}}
\toprule
Species & Group\tabularnewline
\midrule
\endhead
\emph{A. lituratus} & a\tabularnewline
\emph{A. planirostris} & a\tabularnewline
\emph{U. convexum} & b\tabularnewline
\emph{P. helleri} & bc\tabularnewline
\emph{S. giannae} & bc\tabularnewline
\emph{A. pheotis} & bc\tabularnewline
\emph{A. anderseni} & c\tabularnewline
\bottomrule
\end{longtable}

\textbf{4. DISCUSSION}

We found that skull size and inter-sex morphology was significantly
different, variation that can be associated with the diet of the
selected bat species; possibly as a result of differences in energy
requirements during the reproductive season (de Camargo \& de Oliveira,
2012). Although sexual dimorphism has been relatively well documented in
vespertilionid bats, being the females larger and heavier than males
(Bornholdt, Oliveira, \& Fabian, 2008), in phylostomid bats this
information is less documented despite it being the most diverse and
distributed family in the Neotropics (Gardner 2008). However,
L\selectlanguage{ngerman}ópez-Aguirre \& Pérez-Torres (2015) identified that Artibeus lituratus
females in Colombia had greater fluctuating asymmetry in the
splanchnocranium presenting a differential on bite force.

\textbf{4.1 Variation in the bite force associated with craniodental
traits}

Bite force has been established as an important performance trait for
vertebrates that is associated with both cranial morphology and trophic
ecology (Santana, Dumont, \& Davis, 2010). However. studies in bats have
based their conclusions on biomechanical models of the bite force
without adequately testing the assumed correlation of it with the shape
of the skull (Aguirre, Herrel, Van Damme, \& Matthysen, 2002; Dumont,
2007; Herrel, De Smet, Aguirre \& Aerts, 2008). Our study shows, for the
first time, strong quantitative evidence of such a correlation by using
a dataset of cranial traits involved in bite force within a
morphologically diverse clade of New World fruit bats.

\textbf{4.2 Interspecific bite force variation}

Stenodermatinae represents a subfamily of Phyllostomidae bats with a
diet that contains significantly more fruit than other family's bats
(Santana, Grosse, \& Dumont, 2012). They are morphologically diverse
with cranial and mandibular traits that overlap with other feeding
guilds (Santana, Grosse, \& Dumont, 2012), which is reflected in the
variety of foods as well as in the tendency of these species to
specialize in a group of particular fruits (Rojas, Vale, Ferrero, \&
Navarro, 2012; Rossoni, Assis, Giannini, \& Marroig, 2017). Several
craniodental traits contribute to generate a greater bite force and the
differences between these particular traits had showed a clear
segregation between the species (Santana, Grosse, \& Dumont, 2012).

The association in craniodental features and bite force by species can
be explained by the type of fruit consumed. \emph{A. phaeotis} ,
\emph{P. helleri} and \emph{S. giannae} (small-sized and sedentary
nomadic fruit bats) formed an independent group. These species have a
short skull and a shorter distance from the teeth to the jaw joint,
which allows them to consume hard and soft fruits (Dumont et al., 2012),
P. helleri and S. giannae are characterized by elongated and narrow
snouts (Santana, Grosse, \& Dumont, 2012), which is associated with the
consumption of soft and smaller fruits such as those of Solanaceae
(Arias \& Pacheco, 2019; García-Herrera, Ramírez-Fráncel and Flórez,
2019). Species that share cranial and nutritional traits with \emph{D.
anderseni.} For its part, \emph{Uroderma convexum} was distinguished
from the other species studied here by having a parallel rostrum and a
short face that abruptly expands from the apex to the edges of the
lacrimal bone (Mantilla-Meluk, 2014). These characteristics allow them
to exploit mainly hard fruits of plant species such as \emph{Ficus} spp.
(Sagot and Stevens, 2012).

All the big and small-sized nomadic frugivorous bat species studied
here, except for \emph{A. anderseni} and \emph{A. phaeotis} had a
stronger bite force, which was correlated to a greater length of the
skull, shorter palate, narrower width between molars, dentary length,
and higher coronoid process. These traits have been recognized as
predictors of a biomechanical advantage for higher feeding performance
(Dumont et al., 2012). Shortening and widening of the rostrum in\emph{A.
anderseni} and \emph{A. phaeotis} was negatively correlated to bite
force, although it should confer a greater biomechanical advantage; a
pattern that has been repeatedly observed in placental and marsupial
carnivores (Wroe \& Milne, 2007). This is probably due to the fact that
the variation in size could be the most dominant factor and it has been
shown that the differential proportions of the skull determine the
maximum value of the bite force (Herrel, Podos, Huber, \& Hendry, 2005;
Nogueira et al., 2009; Dollion et al., 2016). Recent studies have
suggested that among closely related mammals, larger taxa exhibit
relatively longer faces, whereas smaller taxa exhibit paedomorphic
traits such as proportionally shorter faces and larger brains (Cardini
\& Polly, 2013; Cardini, Polly, Dawson, \& Milne, 2015; Tamagnini,
Meloro, \& Cardini, 2017). This pattern called cranial evolutionary
allometry, has been seen in antelopes, squirrels, fruit bats, cats and
kangaroos (Tamagnini, Meloro, \& Cardini, 2017).

Previous studies have demonstrated that skull size is important in
determining interspecific differences in bite force, specifically,
Nogueira, Peracchi \& Monteiro (2009) found that, in phyllostomids,
size\selectlanguage{english}-corrected bite forces were correlated with shorter rostra,
alignment of the molars along the masseter insertion region, and
expansion of the anterior zygomatic arch and angular process. On the
contrary, our study determined that the variation in bite force in
Stenodermatinal bats is better explained by the length of the skull,
condylo-incisor, condylo-canine, mandibular toothrow and height of the
coronoid, results similar to those reported by Herrel \emph{et
al.,}(2008) in a sample of bat species (including non-phylostomids).

Shortening and widening of the rostrum in A. anderseni and A. phaeotis
was negatively correlated to bite force, although it should confer a
greater biomechanical advantage; a pattern that has been repeatedly
observed in placental and marsupial carnivores (Wroe \& Milne, 2007).
This is probably due to the fact that the variation in size could be the
most dominant factor and it has been shown that the differential
proportions of the skull determine the maximum value of the bite force
(Herrel, Podos, Huber, \& Hendry, 2005; Nogueira et al., 2009; Dollion
et al., 2016). Recent studies have suggested that among closely related
mammals, larger taxa exhibit relatively longer faces, whereas smaller
taxa exhibit paedomorphic traits such as proportionally shorter faces
and larger brains (Cardini \& Polly, 2013; Cardini, Polly, Dawson, \&
Milne, 2015; Tamagnini, Meloro, \& Cardini, 2017). This pattern called
cranial evolutionary allometry, has been seen in antelopes, squirrels,
fruit bats, cats and kangaroos (Tamagnini, Meloro, \& Cardini, 2017).

Another factor that could explain our results is that bats modulate the
force of the bite according to the exploited resources of soft and hard
food (Santana \& Dumont, 2009). Artibeus vary in the size and texture of
the fruits they prefer (Wendeln, Runkle, \& Kalko, 2000) and may exhibit
differential maximum bite force, which can potentially increase by more
than 10\% when consuming hard fruits (Santana \& Dumont, 2009). Another
factor to consider is the possible variation in the intraspecific bite
force in response to the temporal or geographical variation of food
resources or the hormonal status of bats, as proposed by Irschick et al.
(2006). Since the bats included in our study are composed of individuals
from different localities and data (fieldwork and voucher specimens;
Appendix I), our data could be susceptible to this source of variation
and therefore a more detailed study is needed to test this hypothesis
and further improve our understanding of the relationships of bite force
and craniodental traits.

Given that nomad bats consume similar fruits, they could require the
same force to process, so we expected that all bats grouped in this
category would present similar bite forces, however our results
established that the species \emph{Artibeus lituratus} , \emph{A.
planirostris} , followed \emph{U. bilobatum} and \emph{P.
helleri}recorded the highest bite force, probably because these bats
feed on food resources that clearly have different mechanical demands
when consuming even other resources other than the fruits.

All Stenodermatinae are frugivores with short and wide rostrum, but our
results show that can occur some divergent morphologies within the
clade, which is seen in the large number of species within the subfamily
and highly diverse genera (Murillo-Garcia \& De la Vega, 2018). As has
been documented in other vertebrates, closely related species exhibit
strong positive relationships between large skulls and greater
biomechanical advantage (Maestri et al., 2016; Santana \& Miller, 2016).
Our findings illustrate that skull size is a determining factor in the
bite force and emphasizes that the use of functional traits is relevant
to establishing the feeding performance of bat species. This highlights
the importance of studying the relationships between morphology, bite
force and the ecology of the species to get a better understanding of
evolutionary adaptions of highly diverse Neotropical bat groups.

\textbf{Acknowledgements}

The authors wish to thank the staff of the Research Group in Zoology
(GIZ), Universidad del Tolima (Ibague, Colombia). LAR-F and LVG-H thank
the Convocatoria No. 755, 2016 para la Formacion de Capital Humano de
Alto Nivel para el Departamento del Tolima, Colfuturo (miniciencias) and
Departamento del Tolima for support (doctoral scholarships). We thank
the biologists Emmanuel Quintero, David Santiago Varon, Carlos Guzman
and Benjamin Polanco for their support during the field activities and
the people from each of the localities surveyed for their hospitality.
Specimens collected are covered through the ``Permiso Marco de
Recoleccion'' granted to the Universidad del Tolima (Resolucion No.
2191. November 27, 2018. Autoridad Nacional de Licencias
Ambientales---ANLA). We thank Dr Jairo Perez for allowing access to the
voucher specimens of the Museo Javeriano de Historia Natural ''Lorenzo
Uribe, SJ''.

\textbf{Authors' contributions}

L.V.G-H., S.L.P. conceived the ideas and designed methodology; L.V.G-H.,
L.A.R-F established the activities in the field, developed the
methodology and took the craniodental measurements in the different
collections and museums evaluated; A.S-H., L.V.G-H., L.A.R-F and S.L.P.
analyzed the data; B.L, G.G and G.R-F. led the writing of the
manuscript. All authors contributed critically to the drafts and gave
final approval for publication.

\textbf{Data Accessibility}

The data are available
at\emph{https://datadryad.org/stash/share/N6vW4UGTQZbdh132I0gaPR2GYsa4iDCy5YdsEeNaerM}

\textbf{REFERENCES}

Aguirre LF. Herrel A. Van Damme R. Matthysen E. 2002. Ecomorphological
analysis of trophic niche partitioning in a tropical savannah bat
community. Proceedings of the Royal Society B. 269:1271.
https://doi.org/10.1098/rspb.2002.2011.

Aguirre. L. F., Herrel, A., Van Damme R., \& Matthysen, E. (2003). The
implications of food hardness for diet in bats. \emph{Functional
Ecology,} \emph{17} , 201-212.

Arias, E., \& Pacheco, V. (2019). Dieta y estructura trofica de un
ensamblaje de murcielagos en los bosques montanos del Santuario Nacional
Pampa Hermosa. Junin. Peru. \emph{Revista Peruana de Biologia}
,\emph{26} , 169-182. http://dx.doi.org/10.15381/rpb.v26i2.16375

Bels, V., \& Herrel, A. (2019). Feeding. a Tool to Understand Vertebrate
Evolution Introduction to ``Feeding in Vertebrates''. In Bels, V. \&
Whishaw, I. Q. (Ed.), \emph{Feeding in Vertebrates: Evolution.
Morphology} (pp.1-18). Berlin, Germany: Springer.

Bornholdt, R., Oliveira, L. R. D., \& Fabian, M. E. (2008). Sexual size
dimorphism in \emph{Myotis nigricans} (Schinz, 1821) (Chiroptera:
Vespertilionidae) from south Brazil. \emph{Journal of Biology}
,\emph{68} , 897-904. http://dx.doi.org/10.1590/S1519-69842008000400028

Cardini, A., \& Polly, P. D. (2013). Larger mammals have longer faces
because of size -related constraints on skull form. \emph{Nature
communications} , \emph{4} , 2458.

Cardini, A., Polly, D., Dawson, R., \& Milne, N. (2015). Why the Long
Face? Kangaroos and Wallabies Follow the Same ``Rule'' of Cranial
Evolutionary Allometry (CREA) as Placentals. \emph{Evolutionary Biology}
, \emph{42} , 169-176. http://dx.doi.org/10.1007/s11692-015-9308-9

Cisneros, L. M., Fagan, M. E., \& Willig, M. R. (2015). Effects of
human\selectlanguage{english}-modified landscapes on taxonomic, functional and phylogenetic
dimensions of bat biodiversity. \emph{Diversity and Distributions,} 21,
523-533. https://doi.org/10.1111/ddi.12277

de Camargo, N. F., \& de Oliveira, H. F. (2012). Sexual dimorphism
in\emph{Sturnira lilium} (Chiroptera. Phyllostomidae): can pregnancy and
pup carrying be responsible for differences in wing shape? \emph{Plos
one} ,~\emph{7} , e49734. http://dx.doi.org/10.1371/journal.pone.0049734

Dietz, C., Dietz, I., \& Siemers, B. M. (2007). Growth of horseshoe bats
(Chiroptera: Rhinolophidae) in temperate continental conditions and the
influence of climate. \emph{Mammalian Biology} , \emph{72} , 129-144.

Dollion, A. Y., Measey, G. J. Cornette, R., Carne, L., Tolley, K. A.,
Silva, J. M., da. Boistel, R., Fabre, A. C., \& Herrel, A. (2017). Does
diet drive the evolution of head shape and bite force in chameleons of
the genus \emph{Bradypodio} n? \emph{Functional Ecology} , \emph{31} ,
671-684. https://doi.org/10.1111/1365-2435.12750

Dumont. E. R. (2007). Feeding mechanisms in bats: variation within the
constraints of flight. \emph{Integrative and Comparative Biology,47} ,
137-146. https://doi.org/10.1093/icb/icm007

Dumont, E. R. \& O'Neil, R. (2004). Food hardness and feeding behavior
in old world fruit bats (Pteropodidae). \emph{Journal Mammal} ,\emph{85}
, 110-116. https://doi.org/10.1644/BOS-107

Dumont, E. R., Herrel, A., Medellin, R. A., Vargas-Contreras, J. A.,
Santana, S. E. (2009). Built to bite: cranial design and function in the
wrinkle-faced bat. \emph{Journal of Zoology} , \emph{279} , 329--337.
https://doi.org/10.1111/j.1469-7998.2009.00618.x.

Dumont, E. R., Davalos, L. M., Goldberg, A., Santana, S. E., Rex, K.,
Voigt, C. C. (2012). Morphological innovation. diversification and
invasion of a new adaptive zone.~\emph{Proceedings of the Royal Society
B} , \emph{279} , 1797-1805. https://doi.org/10.1098/rspb.2011.2005.

Freeman, P. W., \& Lemen, C. A. (2008). Measuring bite force in small
mammals with a Piezo-resistive sensor. \emph{Journal Mammalogy,89} ,
513-517. https://doi.org/10.1644/07-MAMM-A-101R.1.

Garcia-Morales, R., Moreno, C. E., Badano, E. I., Zuria, I.,
Galindo-Gonzalez, J., Rojas-Martinez, A. E., \& Avila-Gomez, E. S.
(2016). Deforestation impacts on bat functional diversity in tropical
landscapes. \emph{PloS one} , 11, e0166765.
https://doi.org/10.1371/journal.pone.0166765

Garcia-Herrera, L. V., Ramirez-Francel, L. A. R., \& Florez, G. R.
(2019). Consumo de plantas pioneras por murcielagos frugivoros en un
fragmento de bosque seco tropical (Colombia). \emph{Ciencia en
desarrollo,} \emph{10} , 22-31.

Giannini, N. P., \& Kalko, E. K. V. (2004). Trophic structure in a large
assemblage of phyllostomid bats in Panama. \emph{Oikos} , \emph{2} ,
209-220. https://doi.org/10.1111/j.0030-1299.2004.12690.x

Herrel, A., Podos, J., Huber, S. K., \& Hendry, A. P. (2005). Bite
performance and morphology in a population of Darwin's finches:
implications for the evolution of beak shape. \emph{Functional Ecology}
,\emph{19} , 43-48. https://doi.org/10.1111/j.0269-8463.2005.00923.x

Herrel, A., De Smet, A., Aguirre, L. F., \& Aerts, P. (2008).
Morphological and mechanical determinants of bite force in bats: do
muscles matter? \emph{Journal of Experimental Biology} , \emph{211} ,
86--91. https://doi.org/10.1242/jeb.012211

Irschick, D. J., Ramos, M., Buckley, C., Esltrot, J., Carlisle, E.,
Lailvaux, S. P., Bloch, N., Herrel, A., \& Vanhooydonck, B. (2006). Are
morphology-performance relationships invariant across different seasons?
A test with the green anole (\emph{Anolis carolinensis} ). \emph{Oikos}
,\emph{114} , 49-59. https://doi.org/10.1111/j.2006.0030-1299.14698.x

Lopez-Aguirre, C., \& Perez-Torres, J. (2015). Cranial and mandibular
asymmetry in \emph{Artibeus lituratus} (Chiroptera. Phyllostomidae) from
Colombia. \emph{Universitas Scientiarum} , \emph{20} , 141-152.

Maestri, R., Monteiro, L. R., Fornel, R., Upham, N. S., Patterson, B.
D., \& de Freitas, T. R. O. (2017). The ecology of a continental
evolutionary radiation: Is the radiation of sigmodontine rodents
adaptive? \emph{Evolution} , \emph{71} , 610-632.

Manhaes, I. A., Nogueira, M. R., \& Monteiro, L. R. (2017). Bite force
and evolutionary studies in phyllostomid bats: a meta-analysis and
validation. \emph{Journal of Zoology, 302(4),} 288-297.
https://doi.org/10.1111/jzo.12457

Mantilla-Meluk, H. (2014). Defining species and species boundaries
in\emph{Uroderma} (Chiroptera: Phyllostomidae) with a description of a
new species. \emph{Occasional Papers} \emph{Museum of Texas Tech
University} , \emph{325} , 1-25.

Murillo-Garcia, O. E., \& De la Vega, M. E. (2018). Divergence.
convergence and phenotypic diversity of neotropical frugivorous
bats.\emph{Diversity} , \emph{10} , 10-12.
https://doi.org/10.3390/d10030100

Nogueira, M. R., Peracchi, A. L., \& Monteiro, L. R. (2009).
Morphological correlates of bite force and diet in the skull and
mandible of phylostomid bats. \emph{Functional Ecology} , \emph{23} ,
715-723. https://doi.org/10.1111/j.1365-2435.2009.01549.x

Reid, J. L., Mendenhall, C. D., Zahawi, R. A., \& Holl, K. D. (2015).
Scale-dependent effects of forest restoration on Neotropical fruit
bats.\emph{Restoration Ecology} , \emph{23} , 681-689. doi:
https://doi.org/10.1111/rec.12235

Rojas, D., Vale, A., Ferrero, V., \& Navarro, L. (2012). The role of
frugivory in the diversification of bats in the Neotropics.\emph{Journal
Biogeography} , \emph{39} . 1948-1960.
https://doi.org/10.1111/j.1365-2699.2012.02709.x

Rossoni, D. M., Assis, A., Giannini, N. P., \& Marroig, G. (2017).
Intense natural selection preceded the invasion of new adaptive zones
during the radiation of New World leaf-nosed bats. \emph{Scientific
Reports} , \emph{7} , 1-11.

Core Team. (2019). \emph{R: A language and environment for statistical
computing} .Vienna, Austria: R Foundation for Statistical Computing.

Sagot, M, \& Stevens, R. (2012). The evolution of group stability and
roost lifespan: perspectives from tent-roosting bats. \emph{Biotropica}
,\emph{44} , 90-97.

Santana, S. E., \& Dumont, E. R. (2009). Connecting behaviour and
performance: the evolution of biting behaviour and bite performance in
bats. \emph{Journal of Evolutionary Biology} , \emph{22} , 2131-2.
https://doi.org/10.1111/j.1420-9101.2009.01827.x

Santana, S. E., Dumont, E. R., \& Davis, J. L. (2010) Mechanics of bite
force production and its relationship to diet in bats. \emph{Functional
Ecology} , \emph{24} , 776-784.
https://doi.org/10.1111/j.1365-2435.2010.01703.x.

Santana, S. E., Grosse, I. R., \& Dumont, E. R. (2012). Dietary
hardness. loading behavior. and the evolution of skull form in
bats.\emph{Evolution} , \emph{66} , 2587-2598.
https://doi.org/10.1111/j.1558-5646.2012.01615.x

Santana, S. E., \& Miller, K. (2016). Extreme postnatal scaling in bat
feeding performance: a view of ecomorphology from ontogenetic and
macroevolutionary perspectives. \emph{Integrative and Comparative
Biology} , \emph{56} , 459-468. https://doi.org/10.1093/icb/icw075

Sikes, R. S., \& Animal Care and Use Committee of the American Society
of Mammalogists. (2016). Guidelines of the American Society of
Mammalogists for the use of wild mammals in research and
education.\emph{Journal of mammalogy} , \emph{97} , 663-688.

Soriano. P. 2000. Functional structure of bat communities in tropical
rainforests and Andean cloud forests. \emph{Ecotropicos} , \emph{13} ,
1-20.

Shi, B., Wang, Y., Gong, L., Chang, Y., Liu, T., Zhao, X., Lin, A.,
Feng, J, \& Jiang, T. (2020). Correlation of skull morphology and bite
force in a bird-eating bat (\emph{Ia io} ;
Vespertilionidae).\emph{Frontiers in Zoology} ,~\emph{17} , 1-14.
https://doi.org/10.1186/s12983-020-00354-0

Tamagnini, D., Meloro. C., \& Cardini, A. (2017). Anyone with a
Long-Face? Craniofacial Evolutionary Allometry (CREA) in a Family of
Short-Faced Mammals, the Felidae. \emph{Evolutionary Biology} ,\emph{44}
, 476-495. https://doi.org/10.1007/s11692-017-9421-z

Thiagavel, J., Cechetto, C., Santana, S. E., Jakobsen, L., Warrant, E.
J., \& Ratcliffe, J. M. (2018). Auditory opportunity and visual
constraint enabled the evolution of echolocation in bats. \emph{Nature
communications} , \emph{9} , 1-10.
https://doi.org/10.1038/s41467-017-02532-x

Wendeln, M. C., Runkle, J. R., \& Kalko, E. K.V. (2000). Nutritional
values of 14 fig species and bat feeding preferences in
Panama.\emph{Biotropica} , 32, 489-501.
https://doi.org/10.1111/j.1744-7429.2000.tb00495.x

Wroe, S., \& Milne, N. (2007). Convergence and remarkably consistent
constraint in the evolution of carnivore skull shape. \emph{Evolution} ,
61, 1251-1260. https://doi.org/10.1111/j.1558-5646.2007.00101.x

APPENDIX I. SPECIMENS EXAMINED

The following list includes all specimens examined in this study, with
their respective localities.

\emph{Artibeus anderseni} (41)- AMAZONAS, Leticia (ROM 53614, 53615,
53616, 63061) ANTIOQUIA, Los Remedios (ROM 84982), CUNDINAMARCA, La Gran
Curva, 114 km al oeste de Bogota (ROM 48959, 51790), Melgar (ROM 53613,
75300, 75302), Puerto Salgar (ROM 44956, 44957, 44965) TOLIMA, Alvarado,
vereda Rincon de Chipalo, Parque Nacional del Arroz (CZUT-M 2089, 2090,
2091, 2099, 2100), Armero Guayabal, Centro Universitario Regional del
Norte - Universidad del Tolima (ROM 1128, 1415, 1416, 1418, 1434, 1436,
1481, 1492, 1515, 1516, 1517, 1536 ,1537, 1538, 1541, 1624, 1638, 1639,
1641, 1642), Espinal (ROM 88089), Ibague, vereda Aparco (CZUT-M 2098,
2103).

\emph{Artibeus lituratus} (64)- CAUCA, Bellavista (ROM 63232, 63063,
64066, 64067, 64068, 64069, 64070, 64071, 67247, 67253, 67255, 67256,
67261), CUNDINAMARCA, Puerto Salgar (ROM 44811), META, Villavicencio,
Puerto Lopez (ROM 88083), TOLIMA, Ambalema, vereda Chorrillo (CZUT-M
1410, 1546, 1567, 1569, 1668), Armero Guayabal, Centro Universitario
Regional del Norte - Universidad del Tolima (CZUT-M 1124, 1413, 1545,
1546, 1564, 1565, 1566, 1596, 1628, 1629, 1631, 1704), Ibague, vereda
Chucuni (CZUT-M 0763, 0764, 0766, 0835), Buenos Aires (ROM 44882,
44885), Melgar, Santo Tomas (MUJ 00826, 00827, 00828, 00829, 00830,
00832, 00833, 00834) Suarez, vereda Batatas (CZUT-M 0299, 0301, 0303,
0305, 0344, 0346), vereda Aguas Claras (CZUT-M 0363), vereda Los
Arrayanes (CZUT-M 1230, 1231), PUTUMAYO, vereda Guascayaco (ROM 46356,
49206, 49209, 49211), VAUPES, Mitu (ROM 45258, 45259, 45260), VALLE DEL
CAUCA (ROM 44889, 44890).

\emph{Artibeus planirostris} (75)- META, Fuente de Oro, Km 9 Carretera
Puerto Limon (ROM 90109, 91409), TOLIMA, Ambalema, vereda Chorrillo
(CZUT-M 1468, 1469, 1470, 1471, 1472, 1473, 1474, 1475, 1476, 1477,
1494, 1544, 1568, 1583, 1584, 1585, 1586, 1587, 1588, 1589, 1590, 1591,
1592, 1599, 1600, 1605, 1632, 1643, 1644, 1645, 1646, 1669, 1670, 1671,
1672, 1673, 1695, 1696, 1697, 1699, 1700, 1701), Ibague, vereda Aparco
(CZUT-M 1994, 1995, 1996), Armero Guayabal, Centro Universitario
Regional del Norte - Universidad del Tolima (CZUT-M 1412, 1414, 1493,
1513, 1514, 1542, 1543, 1593, 1594, 1595, 1598, 1698, 1705, 1712, 2008,
2049), Suarez, vereda Batatas (CZUT-M 0243, 0244, 0245, 0298, 0300,
0302, 0304, 0345, 0347, 2070), PUTUMAYO, vereda Guayaco (ROM 49211),
Vereda San Miguel (67246).

\emph{Artibeus phaeotis} (28)- AMAZONAS, Leticia (ROM 53610, 53611,
53612), CAUCA, Bellavista (ROM 64064, 64059, 64057), MAGDALENA, Santa
Marta (ROM 79885), TOLIMA, Alvarado, vereda Rincon de Chipalo, Parque
Nacional del Arroz (CZUT-M 2106), Ambalema, vereda Chorrillo (CZUT-M
1346, 1549, 1572, 1647) Armero Guayabal, Centro Universitario Regional
del Norte - Universidad del Tolima (CZUT-M 1127, 1435, 1437, 1518, 1571,
1625, 1640, 1702, 1703, 1706, 1710, 1711), Mariquita (MUJ 0269, 0285,
0286), Melgar (MUJ 00835).

\emph{Platyrrhinus helleri} (31) -MAGDALENA, Santa Marta (ROM 79882),
TOLIMA, Alvarado, vereda Rincon de Chipalo, Parque Nacional del Arroz
(CZUT-M 2009, 2062), Ambalema, vereda Chorrillo (CZUT-M 1709), Armero
Guayabal, Centro Universitario Regional del Norte - Universidad del
Tolima (CZUT-M 1539, 1547, 1570, 1602, 1626, 1630, 2010), Libano,
Hacienda La Trinidad (ROM 88082), Ibague, vereda Martinez (CZUT-M 0027,
0116), San Sebastian de Mariquita (CZUT-M 1063, 1065, 1072), Melgar
(CZUT-M 1928), vereda Santo Tomas (MUJ 00837, 00838, 00839, 00841),
Suarez, vereda Batatas (CZUT-M 0341, 0364, 1914), PUTUMAYO (ROM 403559,
46375, 46353, 63239), VAUPES, Mitu (ROM 45274, 45273),

\emph{Uroderma convexum} (40)- MAGDALENA, Santa Marta (ROM 79886),
TOLIMA, Alvarado, vereda Rincon de Chipalo, Parque Nacional del Arroz
(CZUT-M 2093, 2101, 2104, 2105), Armero Guayabal, Centro Universitario
Regional del Norte - Universidad del Tolima (CZUT-M 1126, 1438, 1511,
1512, 1540, 1597, 1601, 1622, 1623, 1633, 1634, 1635, 1636, 1637, 1694,
1707), Ibague, vereda Aparco (CZUT-M 2095, 2096, 2097, 2102), San
Sebastian de Mariquita (CZUT-M 1054, 1070), Melgar (CZUT-M 1927, MUJ
00842, 00843, 00844, ROM 62508), Suarez, vereda Batatas (CZUT-M 1915),
PUTUMAYO, vereda Guascayaco (ROM 46360), vereda Horno (ROM 63240,
63240), VAUPES, Mitu (CZUT-M 45268, 45267, 45366, 45265).

\emph{Sturnira giannae} (30)- TOLIMA, Ambalema, vereda Chorrillo (CZUT-M
1116, 1117, 1118, 1296, 1310, 1318, 1342, 1343), Armero Guayabal, Centro
Universitario Regional del Norte - Universidad del Tolima (CZUT-M 1129,
1403), Ibague, vereda Aparco (CZUT-M 2014, 2015, 2016, 2021, 2022, 2023,
2028), Mariquita (CZUT-M 1056, 1061), Suarez, vereda Batatas (CZUT-M
0234, 0246, 0307, 0362, 1236), PUTUMAYO (ROM 40374, 40313, 40349, 40375,
46373, 49184).

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

